Plants of the South American Pacific Mangrove Swamps (Colombia, Ecuador, Perú) - Xavier Cornejo by Zhino Design

Xavier Cornejo, editor - 2014

Plants of the South American Pacific Mangrove Swamps (COLOMBIA, ECUADOR, PERU)

Xavier Cornejo, editor 2014

Copyright © Xavier Cornejo, 2014. All rights reserved. No portion of this book may be reproduced in any form or by any means, including electronic storage and retrieval systems, except by explicit, prior written permission of the publisher except for brief passages excerpted for review or critical purposes. ISBN: 978-9942-13-667-1

El libro debe citarse de las siguientes maneras: a. Para el volumen completo: Cornejo, Xavier (ed.). 2014. Plants of the South American Pacific Mangrove Swamps. Publicaciones del Herbario GUAY, Facultad de Ciencias Naturales de la Universidad de Guayaquil. b. Para las contribuciones: <Autor[es]>. 2014. <Contribución>. Páginas xx-xxx en: Cornejo, Xavier (ed.). 2014. Plants of the South American Pacific Mangrove Swamps. Publicaciones del Herbario GUAY, Facultad de Ciencias Naturales de la Universidad de Guayaquil.

Editorial board: Claes Persson: (Herbarium GB, University of Gothenburg) Vicki Breazeale (Great Wilderness) Supervisor and approval: Carmen Bonifaz (Facultad de Ciencias Naturales, Universidad de Guayaquil) Book design: Luis Flores Burgos (zhinodesign@gmail.com) Printed by: Universidad de Guayaquil Cover illustration: Mangroves of Muisne, northwestern Ecuador. Photo by Xavier Cornejo.

Field work has been partially supported by Conservation International and Scott A. Mori (NY). The results presented in this publication contribute to the integrated conservation strategies and the sustainable use of natural resources in the region, that is legally supported in the Constitution of Ecuador (2008) and in the Plan Nacional del Buen Vivir 2013–2017.

FOREWORD

I take great pleasure in introducing the Plants of the South American Pacific Mangrove Swamps by Xavier Cornejo with contributions by Enrique J. Peña-Salamanca, Jaime R. Cantera-Kintz, Philip A. Silverstone-Sopkin, Reynaldo Linares-Palomino, Marisela Monzón-Ramos and Carmen Bonifaz to the botanical community and government institutions. I hope it will stimulate research and conservation of the mangroves.

Mangroves covered in the year 2000 an area of approximately 140.000 km2 worldwide, but had

by then already diminished considerably. The loss of mangroves in Colombia, Ecuador, and Peru varies significantly. The available data from Colombia indicates a 4.5% loss from 1969 to 1996. The loss in Ecuador was 59% from 1969 to 2006, while in Peru it was 30% from 1982 to 1992. Most of the loss was an effect of conversion of land use from localized extraction of wood and protein (fish and crabs principally) to an intensive exploitation of shrimp. To clear the way for massive shrimp farms the mangrove was cleared and basins for shrimp farming established. Mangroves are relatively poor in species, compared to other tropical forest types, for plants to grow successfully in the saline environment requires several physiological changes, and the despite growing with abundant water, the plants struggle to survive high levels of salinity and temperature, leading to low levels of available water for the necessary physiological processes. The present volume contains 49 families, 135 genera, and 222 species. It includes keys of identification and detailed description of all the species. A wealth of information that is particularly useful for an ecosystem that is as threatened as the Pacific mangroves. The book also puts forward a classification of different mangrove types. Some novelties are included as well as many overlooked species have been documented for the first time. The publication is the result of field and herbarium work carried out by Cornejo since 1991. The book is illustrated with 48 plates containing 141 species of plants and constitutes a “must have” volume for field work in the mangroves as well as for any and all environmental studies carried out in these most fragile ecosystems, that for a long time have been undervalued.

Peter Møller Jørgensen Curator Missouri Botanical Garden

PREFACE In recent years, molecular work achieved by several authors yielded in a new systematic arrangement for some well-known mangrove taxa that occur along both coasts of the American continent. Thus, genera as Avicennia and Pelliciera, do not belong to Avicenniaceae or Pelliceriaceae, respectively; and Rhizophora harrisonii is not a species at all. That is without taking into account the taxonomical confusion between R. racemosa and R. x harisonii, and the total absence of the white mangrove Laguncularia racemosa var. glabriflora in the official checklists and conservation or management plans in Ecuador and Peru. Furthermore, the discovery of four new species of plants in mangroves and new distributional records for several other overlooked species from northwestern South America turned out to be great additions to the regional mangrove flora. Facing this scenario and regarding to the absence of publications that deal with all or most of all mangrove plants mainly in Colombia and Ecuador, a volume has been prepared for the region. The present volume is a reliable source of relevant information about plants, providing updated scientific names, keys to identification and descriptions at family and species level, as well as data of distribution, ecology, and IUCN categories for 222 species of mangrove plants that occur on the Pacific coast of northwestern South America. In addition, vernacular names, biological interactions, uses, and etymology have been added for each species if available. The book is divided into three chapters: The first chapter presents details of mangrove swamps on the Pacific coast of Colombia, Ecuador and Peru. The second chapter discusses mangrove plant habits as well as the diversity of pteridophytes, gymnosperms and flowering plants at family and species level, and the dispersal methods regarding to the habit and main taxonomic groups. In this chapter, mangrove trees are classified as major, minor and facultative, and the remaining floristic elements are grouped according to the distributional pattern. As a result, the Chocoan and the Equatorial-Pacific mangroves are here proposed based on the floristic composition. The flora of mangroves is presented in the third chapter which provides full data for each recorded species. It is a useful tool for education at University as well as high school level, habitat restoration and management, conservation, research and tourism. The data and observations have been gathered during many field trips involving plant collection achieved throughout the years since 1991. The research has been enhanced with the review of original publications on cited species and main synonyms and comparing them against the bulk of specimens collected in mangroves of northwestern South America that are housed in eleven herbaria. The following herbaria were visited and the assistance of their curators and staff is here acknowledged: COL, GUAY, L, MO, NY, QCA, QCNE, SEL, US, and WIS (incl. MAD). Mónica Murillo Muñoz and Rubén Darío González Román (Universidad del Valle), collaborated reviewing the herbarium collections in CUVC.

The following plant taxonomists kindly contributed with determinations and useful comments or discussions in their specialties: W. R. Anderson (†), C. C. Berg (†), M. Blanco (FLAS), R. Callejas (HUA), T. Croat, C. H. Dodson (MO), R. Eriksson (GB), P. Fryxell (†), V. Funk (US), D. GiraldoCañas (COL), B. Hammel (MO), E. Hágsater (AMO), B. Hansen (USF), B. Klitgaard (BM), R. Kriebel (INBio), G. P. Lewis (K), R. Moran (NY), R. Kuijt (LEA), S. Lægaard (AAU), C. Luer (SEL), H. Luther (SEL), J. Madsen (AAU), H. Navarrete (QCA), F. Pupulin (Lankester Botanical Garden), J. Ricketson (MO), and L. Skog (US). Thanks to F. Christian Thompson, Adjunct Scientist (Emeritus) at the Department of Entomology, and to Natalia Vandenberg from the Systematic Entomology Lab-USDA of the Smithsonian Institution for the identification of some flower visitors to black and white mangrove as Syrphid flies and Lady beetles, respectively. Gabriel Brito collaborated with the identification of butterflies. Donald Bright, curator at the Department of Bioagricultural Sciences and Pest Management, C. P. Gillette Museum of Arthropod Diversity, Colorado State University, identified the specimens of beetles that infest the hypocotyls of Rhizophora. Field work has been partially supported by Conservation International and Scott A. Mori (NY). Special thanks to brother Máximo Cangá, the motorman and expert field guide at San Lorenzo, Ecuador and to Diana Vitores for useful discussions. Hannah Grochowski and Thomas Hernandez, reviewed the grammar and the text of the manuscript. Most of the bibliography cited was found in the library of The New York Botanical Garden and in the Biology library at the University of Wisconsin-Madison.

TABLE OF CONTENTS 7

INTRODUCTION *

CHAPTER I. MANGROVES ON THE PACIFIC COAST OF SOUTH AMERICA

Mangroves of the Pacific coast of Colombia**

Mangroves of Ecuador*+

Mangroves of Peru***

CHAPTER II. MANGROVE PLANTS*

Plant habits, diversity and dispersal methods

Major, minor, facultative mangroves, and other floristic elements

The Chocoan and Equatorial-Pacific mangrove forests

Conclusions and recommendations

CHAPTER III. FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS*

Format of the Flora

Key to the families

Ferns

Gymnosperms

Seed Plants

LITERATURE CITED

299

INDEX TO NAMES OF PLANTS

307

(*) Xavier Cornejo (+) Carmen Bonifaz (**) Enrique J. Peña-Salamanca, Jaime R. Cantera-Kintz and Philip A. Silverstone-Sopkin (***) Reynaldo Linares-Palomino and Marisela Monzón-Ramos

Xavier Cornejo, editor - 2014

INTRODUCTION The term mangrove is derived from a combination of the word “mangro”, the original common name for Rhizophora mangle in Surinam (Karsten, 1890), and “grove”, the English word for a small stand of trees. Currently, this term is widely applied to a diverse group of tree, palm, and shrub species, which belong to several taxonomically unrelated families. These species are characterized by having specialized morphological and physiological adaptations to successfully settle and live on periodically flooded, saline to brackish, muddy loose soils. They are distributed worldwide along tropical coastlines, inhabiting the intertidal and adjacent supratidal zones, mostly along the estuarine margins, and play an interactive roll in soil formation at the outer-seaward-fringe. The word mangrove is assigned to the vegetal formations and communities1 that inhabit a mangrove, as well as to the entire mangrove ecosystem. According to the theory of continental drift, based on the history of plate tectonics, the separation of the southern supercontinent, the Gondwanaland, occurred in the Late Cretaceous Period. Early Angiosperms were migrating into the then shallow seas between separating continents which allowed the evolution and speciation of mangroves (McCoy and Heck, 1976). The theory is supported by the evidence of fossil pollen in the earliest records of Rhizophoraceae (the red mangrove family), which date back to the Upper Cretaceous period, approximately 20 million years after the first records of angiosperms (Muller, 1981). The taxonomical diversity of mangroves, composed as they are by species of unrelated families, suggests that there have been frequent and independent adaptations and that each species within the mangrove flora has followed a different evolutionary pathway (Duke, 1992). Based on fossil pollen deposits, it has been suggested that the present Western Hemisphere mangrove floras are a relic of historically more diverse mangroves (Gentry, 1982). In northwestern South America, the glacial advances during the Pleistocene restricted many species to the Chocó2 refuge, which “seems only to have preserved more or less the evolutionary dead-ends” (Gentry, 1982). Unfortunately, the majority of living mangrove species are poorly understood in regard to their fossil records (Duke, 1992). The first records of Rhizophora and Pelliciera found in Panama date back to the Eocene (Graham, 1977), approximately 40 million years ago. Rhizophora later became the predominant mangrove element during the Miocene-Pliocene (Germeraad et al., 1968). On the Pacific coast of Ecuador, the presence of Rhizophora spp. in prehistoric mangroves date back to the Early Holocene, between 12.000 to 7.000 years ago (Heusser and Shackleton, 1994).

1 The term “mangal” Macnae (1968) has been proposed to reference communities of mangrove plants. This definition has not

been used here.

2 Chocó, the word, originates from “choque”, which means gold in the Aymara indigenous tongue, used to refer to El Dorado, the fabulous land of gold of the Spaniard conquerors’ tale (Cúneo-Vidal, 1925).

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

On the Pacific coast of the Americas the mangrove ecosystem extends from Puerto Lobos (30º15’N

112º51’W) in the State of Sonora of northwestern Mexico southwards along the coastline to the Manglares de San Pedro de Vice (5º34’10” S 80º54’ W) in the Department of Piura, Peru (Cintrón et al., 1981; CDCUNAML, 1991). On the South American Pacific coast, the mangrove forests are located along Colombia, Ecuador and northwestern Peru. They are commonly characterized by a low alfa diversity of trees and shrubs (Chapman, 1975; Tomlimson, 1986; Duke, 1992), mainly composed of the well-known major mangrove taxa: Rhizophora mangle, R. racemosa, R. x harrisonii, Laguncularia racemosa and Avicennia germinans (plate A). They tend to separate into gregarious societies and display a monotonous canopy due to their relative uniformity, in contrast to the commonly multispecific arborescent composition of the terra firme forests. The mangrove plants diversity is highest to the north of its distributional range, in the Departments of Choco and Valle, in northwestern Colombia, where the increasing epiphytic diversity caused by higher rainfall, estimated average between 4,000 to over 9,000 mm per year (Gentry and Dodson, 1987), makes a remarkable contribution to mangrove flora, providing to those ecosystems a vegetation of great luxuriance. Floristic richness decreases to the south, near latitude 0º, from the Province of Manabi, Ecuador, where the average of annual rainfall is less than 600 mm, and becomes increasingly depauperate southwards to northwestern Peru. The floristic richness that might be correlated to the latitudinal gradient is more likely a response to different levels of precipitation in the several life zones located along the coastal areas from western Colombia to Peru. Most species, excepting major mangroves, located in the rainforest and wet forest life zones in our area, are ecologically restricted in their geographical distribution by the dry forests streching mainly from the Province of Manabi to the south. The few exceptions, mostly epiphytic species (e.g., Epiphyllum rubrocoronatum), seem to occur only in the foggy and/or moist-windy areas of the dry forest life zone, located in the mangrove swamps at the interior of the Gulf of Guayaquil, in western Ecuador. On the other hand, again with exception of most of major mangroves, the species that typically occur in the mangrove ecosystems located in the dry forest, very dry, dry thorn scrub, and desert scrub life zones (e.g., Psittacanthus divaricatus) do not occur in the wetter life zones located to the north. These marked patterns in the associate floristic distribution suggest the presence of at least two floristic groups in the Pacific mangroves of northwestern South America, which are herein proposed as: i) The Chocoan mangroves, distributed in the rainforests, wet forest and to the moist forest life zones; those are geographically centered at the Pacific coast of Colombia, discontinuously extending towards northwest along the adjacent coast of southwestern Mesoamerica to the Peninsula de Osa in southwestern Costa Rica, reaching to the south to the estuary of Cojimies (approximately to 0° 06´ N), in the north of Manabi Province, northwestern Ecuador (plate B: A-C). And, ii) the Equatorial-Pacific mangroves, located in the dry forests of western Ecuador (having less than 600 mm of annual rainfall) from near latitude 0º, nearby the town of Pedernales in the Manabi province, and southwards to the lower precipitation life zones reaching the coast of Peru in the isolated Manglares San Pedro de Vice in Piura, at

Xavier Cornejo, editor - 2014

the end of the whole range of latitudinal distribution for mangrove swamps in the eastern Pacific (5° 34’ S; plate B: D). An exception is the mangrove of Atacames (0° 52´N), located in a patch of discontinuous coastal dry forest in the province of Esmeraldas, overlapping the southern range of distribution for the Chocoan mangroves. Mangrove trees reach their highest structural development in the Choco region, which contains the richest flora of all American mangroves and is one of the wettest places on earth (Prance, 1982; Gentry, 1982). The mangroves in the area of the Choco are exposed to extreme fresh water runoff, as well as high nutrient concentrations from inland ecosystems carried downstream by rivers, resulting in giant mangrove forests commonly dominated by Rhizophora, which reach up to 40 m in height in some areas from Darien, Panama to northwestern Ecuador (Cuatrecasas, 1958a, 1958b; Little, 1969; Cintrón et al., 1981, 1985; Alvarez-León, 1993; Cornejo, 1994). Amazingly, in the northwesternmost region of Ecuador, gigantic individuals of Rhizophora of up to 50 m in height and trunks over 1 m in diameter occur in the village of Majagual (plates B: C; 39: F). These have been reported as the tallest of the American continent and the world (Little, 1969; Actmang, 1995). On the landward side, plants of all habits such as herbs, shrubs, trees, lianas, vines, epiphytes, parasites, and/or stranglers from the adjacent inland ecosystems, whether terra firme forests or freshwater swamps, form a back-mangrove belt of transitional vegetation that is always characterized by an intermixed floristic composition from both adjacent ecosystems. These transitional areas have different extensions and patterns of diversity, which are always directly correlated to the geomorphology, rainfall, soil type, and vegetal formation of each intermixing ecosystem. In the Equatorial-Pacific mangroves the transitional areas tend to be narrower and floristically less diverse than those from the mangroves located in the Chocoan region northwards. The higher salt concentration in the drier life zone soils, a result of lesser precipitation, water runoff, and higher levels of evapotranspiration, acts as an ecological barrier that does not allow most terra firme plant species to colonize on mangrove soils. Thus, in the range from dry forest to thorn woodland life zones, the mangrove vegetation and the adjacent terra firme ecosystems are more marked. The narrow transitional area makes for easy differentiation, even from a distance, among the evergreen and often bright olivegreen mangrove foliage in contrast to the regularly deciduous and rather opaque foliage of the adjacent terra firme, wetland or salt marsh ecosystem. On the other hand, the Chocoan mangroves are more or less continually washed by rainfall, causing a decreased salt concentration in the soil, which allows a broader landward belt of transitional vegetation where the floristic composition gradually morphs from mangrove into the next ecosystem whether a terra firme forest or a fresh water wetland, all of them having a continuous green appearance with no clear boundary. A video of transitional vegetation in the Chocoan mangroves can be seen on this link: http://www.youtube.com/watch?v=XrlEXq0KNoo This book includes only plant species found in the intertidal zone of mangrove communities, whether they are typical of mangroves or better represented in adjacent ecosystems. These comprise 49 families, 135 genera, and 222 species (Tables 4, 8, 9, Fig. 1).

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

MANGROVE SWAMPS DISTRIBUTION (RED LINE) ALONG THE PACIFIC COAST OF NORTHWESTERN SOUTH AMERICA

Plate by Kandis Elliott (2010), modified by Xavier Cornejo

CHAPTER I.

MANGROVES ON THE PACIFIC COAST OF SOUTH AMERICA

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

MANGROVE FORESTS OF THE PACIFIC COAST OF COLOMBIA Enrique J. Peña-Salamanca, Jaime R. Cantera-Kintz and Philip A. Silverstone-Sopkin Departamento de Biología, Facultad de Ciencias Naturales y Exactas, Universidad del Valle, A.A. 25360, Cali, Valle, Colombia enrique.pena@correounivalle.edu.co

INTRODUCTION

In 1992, Colombia was tenth in the world in area of mangrove forests, with 501,300 ha on the

Caribbean and Pacific coasts (Groombridge, 1992). More recent estimates based on aerial photographs indicate a decrease in area to 379,035 ha, which represents only 0.72% of all Colombian forests. It has been estimated that 77% of this area (292,724 ha) is located on the Pacific coast (Zambrano and Rubiano, 1996), 86,214 ha on the Caribbean coast, and 97 ha in the archipelago of San Andrés and Providencia (Saenger et al., 1983). The decrease suggests the presence of several factors that cause destruction of mangroves on the Pacific coast, such as erosive action of waves along the coastline, dynamic geomorphology of rivers, incursion by invasive vegetation, expansion of aquaculture, and anthropogenic deforestation. At present, the mangrove vegetation of the Pacific coast of Colombia is considered moderately disturbed (24%), with very few areas highly disturbed (4%). Mangroves now cover about 300,907 ha, of which 67,504 are on the Caribbean coast and 233,403 on the Pacific coast.

The following species of mangroves have been reported from the Pacific coast of Colombia: Avicennia

bicolor, A. germinans, A. schaueriana (Acanthaceae); Conocarpus erectus, Laguncularia racemosa (Combretaceae); Mora megistosperma, M. oleifera (Fabaceae); Pelliciera rhizophorae (Tetrameristaceae); Rhizophora brevistyla, R. harrisonii, R. mangle, R. racemosa, and R. aff. samoensis (Rhizophoraceae). According to more recent work, however, there are only seven valid species: Avicennia germinans, Conocarpus erectus, Laguncularia racemosa, Mora oleifera, Pelliciera rhizophorae, Rhizophora mangle, and R. racemosa. The other names are considered synonyms or incorrect identifications of species from other parts of the world. Some authors believe that Mora oleifera should not be considered a mangrove (Prahl, 1989; Prahl et al., 1990; Cantera, 1991; Jiménez, 1994; Sánchez-Páez et al., 1997; INVEMAR, 2008). However, in the present publication M. oleifera is regarded as a facultative mangrove.

Xavier Cornejo, editor - 2014

TYPES OF MANGROVE VEGETATION

Mangrove vegetation shows high variability in structural development because of physical factors of the environment. Among these factors are nature and composition of the substrate, topography, tidal amplitude and range, amount of fresh water, protection against wave action, nutrient flow and concentration, climate (amount and periodicity of precipitation and length of dry periods), evaporation, air and water temperature, and salinity. The action of all of these factors determines the structure of a mangrove swamp. Mangrove vegetation has been classified in physiographic types by Lugo and Snedaker (1974), but these types were simplified by Cintrón et al. (1980). According to this classification, on the Pacific coast of Colombia the following types of mangrove vegetation are found (Prahl et al., 1990; Cantera, 2008).

Sand bar mangrove swamps

These mangrove swamps are characterized by developing behind a protective bar (usually sandy), known locally as a “bajo,” which is oriented parallel to the coast and is separated from the coast by a tidal lagoon. This sand bar acts as a sediment trap, which permits the formation of mudflats, protected from the waves, in the shallow zones of the lagoon. The zonation of these mangrove swamps includes an outer belt of young pioneer mangroves, almost always of red mangrove (Rhizophora mangle or R. racemosa). These two species dominate the zone of unstable substrates. More stable substrates, usually with a high content of sand and silt, are normally dominated by stands of black mangrove (Avicennia). The most stable zones, usually with Tertiary substrates exposed to the tides, are occupied by mangle piñuelo (Pelliciera rhizophorae) and nato (Mora oleifera). At the zone of maximum tides, the mangrove swamp adjoins the naidisales (stands of Euterpe palms), and these adjoin fresh water swamps and cuangariales (stands of the myristicaceous tree Otoba). On the highest parts of the internal border of the stabilized sand bar (now partially sedimented) there are populations of mangle jeli (Conocarpus erectus) and majagua (Talipariti tiliaceum, Malvaceae). These associations are continuous with communities of black mangrove, forming broad stands in the zones that are richest in silt and sand. The unstable substrates, usually toward the interior of the basin, have populations of red mangrove, exposed to a continuous tidal interchange in tidal channels (esteros). These esteros break through the beaches at some points, permitting temporary connection with the sea.

Border mangrove swamps

These mangrove swamps depend largely on topography and substrate type. They form on eroded substrates along protected coastal margins (plate B: A). This type of mangrove vegetation is characterized

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

by a frontal zone with unstable substrates dominated by red mangrove, with a stable platform dominated by mangle piĂąuelo. Other species may be present in small numbers, with irregular patterns of distribution.

River mangrove swamps These swamps develop along the banks of rivers up to the high tide level (plate A: B). They

attain their maximum development on the Pacific coast of Colombia because of continuous supply of nutrients. The sediments are of silt and clay and may form layers more than 5 m thick. The zone of the river that is affected by tides and dominated by unstable substrates is colonized by belts of red mangrove, with trees that can reach a height of 30 m. The most elevated zones that are influenced by tides, as well as platforms exposed to reduced tidal interchange, are usually dominated by stands of black mangrove. Behind these belts and in zones where ponds of rainwater form, there are naidisales and cuangariales, which are limited to zones covered with fresh water. Along the low river banks (recent beaches) and in areas exposed to temporary penetration of brackish water (usually during wide-ranging tides or pujas), there appear belts of white mangrove (Laguncularia racemosa) and stands of black mangrove (Avicennia germinans). More stabilized substrates with water of very low salinity generally are dominated by natos.

Mangrove swamps of islets On the Pacific coast of Colombia, there exist two types of mangrove swamps of islets. The most

common form is a modification of the border mangrove swamps. When erosion occurs on several parts of geographical formations such as peninsulas, very small islands may be formed, which are surrounded by mangrove swamps whose distribution resembles that of border mangrove swamps, with R. mangle and P. rhizophorae as dominant species, growing on hard substrates that come from the cliffs. This type of vegetation also can form on muddy beaches that are slightly exposed or permanently submerged, occupied by red mangroves. The other type forms when fine material (mud) is deposited at shallow depth in bays or protected zones; these areas are colonized by embryos of red mangrove, causing retention of material and forming a mangrove island. This type of mangrove vegetation is characterized by continuous leaching of its soils and by the presence of only one species (usually R. mangle).

Dwarf mangrove swamps This name is applied to all mangrove formations that develop on inadequate substrates, such

as platforms of sedimentary rocks exposed to salt water and sandy basins with little tidal flow. The mangroves usually do not surpass 4 m in height and develop abnormally.

Xavier Cornejo, editor - 2014

DISTRIBUTION OF MANGROVE VEGETATION ON THE PACIFIC COAST OF COLOMBIA The Pacific coast of Colombia is separated into two different large physiographic zones (West, 1956). To the north of Cabo Corrientes, in the department of the Chocó, the coast is high, consisting mainly of cliffs and dominated by the foothills of the Serranía de los Saltos and the Serranía de Baudó. To the south of Cabo Corrientes, an alluvial coastal plain begins to form, somewhat narrow at first because of the proximity of the Serranía del Baudó, but becoming broader south of the Bay of Buenaventura, with a low coast characterized by flooded zones formed by old fluvial deposits, until Cabo Manglares on the Ecuadoran frontier (Martínez, 1993). The northern coast is relatively high, not appropriate for the formation of mangrove vegetation, and usually only small mangrove swamps are formed, especially in small colluvial bays, where terrigenous sediments carried down by rain and exposed to tidal action accumulate. Near the Panamanian frontier there is a large mangrove swamp at Juradó, which is the largest on the northern coast. It is protected from direct wave action by a wide sand bar, which runs parallel to the coast, and then is separated from the coast by an estuarine lagoon, which receives water from rivers and tides (Prahl et al., 1990). In Bahía Cupica, Nabugá, Solano, and El Valle there exist small border mangrove swamps (restricted to depressions with terrestrial sediments influenced by the tides), which are not well developed, except at the Ensenada de Utría. This inlet is a large geological fault with well developed mangrove vegetation, principally in its interior, where there are erosion platforms that have been covered with stabilized sediments on which grow border mangrove swamps dominated by Pelliciera rhizophorae, which are bordered inland by moist tropical forest. Toward the borders of these platforms there are some estuarine canals with soft substrates, upon which narrow forests of Rhizophora mangle form. Beginning at the inlet of Utría, the coast is characterized by the presence of rocky cliffs that continue until Punta Jurubidá. A little farther south is the border mangrove of Nuquí, which extends in the form of a horseshoe in a colluvial basin from Jurubidá to Coquí and the Gulf of Tribugá. This mangrove swamp is developed as a large colluvial basin (lateral accumulations of sediments in a valley whose movement and transport are reduced [Bergoeing, 1998]). It is characterized by a wide extensive sand bar, which protects the mangrove swamp from wave action; this bar is interrupted at some points by tidal channells that permit aqueous interchange between the sea and the estuary. From Cabo Corrientes southward, the great Pacific alluvial plain appears, partly occupied by large estuarine systems with very well developed mangrove swamps and fresh water swamps. The first mangrove swamp of this coastal plain is that of Virudó, which extends until the inlet of Catripe and the mouth of the Río Purricha. This zone is characterized by internal lagoons, long sandy frontal barriers, and consolidated beaches. In almost all the mangrove swamps of this type Rhizophora predominates. Other important mangrove swamps are present at the mouths of the Ríos Baudó and Docampadó, with riverine

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

forests in which Rhizophora racemosa is dominant. In the zones at the mouths of the rivers, large sand bars and shoals form. There also are some points on the coast where there are well consolidated bars which, by blocking the penetration of the tides, have permitted the formation of fresh water swamps with cuangariales facing the coast. Farther south are the mangrove swamps of the delta of the Río San Juan, protected by frontal sand bars and stabilized beaches that are not influenced by the tides, especially in the zones of Torogomá, Charambirá, Chavica, and El Choncho. The distribution of the mangrove swamps is distinctive, with a belt of Avicennia germinans after the consolidated beach, while Rhizophora mangle occupies unstable, generally muddy soils bordering estuaries and tidal canals. These mangrove swamps continue along the arms of the river in the form of riverine forests up to the point of maximum penetration of brackish tides. On substrates of consolidated clay, especially in the upper part of the river, but always in zones affected by brackish tides, small forests of Mora oleifera can be seen (Restrepo and Cantera, 1995; Sánchez-Páez et al., 1997). Starting at the Río San Juan and its large delta, the coastal plain is interrupted by cliffs of Tertiary sedimentary rocks, which surround most of the northern part of the bay of Málaga and the northern coast of the bay of Buenaventura. At Bahía Málaga (plate B: A), some border mangrove swamps are developed on platforms of eroded cliffs, with rings of mangroves at the intertidal zone and with elevated central cones covered with tropical moist forest (Peña et al., 2010). These substrates are relatively stable, which permits the proliferation of mangle piñuelo, while on the external borders along the tidal channels belts of red mangroves form. Even in rocky zones, dwarf mangrove forests grow. Toward the back of the bay and the southern part of the Bahía de Málaga, there are river type mangrove swamps, dominated by red mangroves and black mangroves, while on elevated and stabilized zones, corresponding to remains of old abrasion platforms, there are forests of mangle piñuelo. Toward the headwaters, in zones that are occasionally inundated with brackish water, there are well developed populations of mangle nato, which form homogeneous forests known as natales (Cantera, 1991; Cantera and Arnaud, 1995). To the south of the bay of Málaga, the isthmus of Pichidó extends, which continues up to the northern coast of the bay of Buenaventura. Along this coast, bordered by cliffs, there are isolated mangrove swamps. These border mangrove swamps develop in eroded zones with depressions, as at Piangua Grande, La Bocana, Isla Cangrejo, and the tidal channel of Gamboa. In these a distinct zonation is not evident, and generally there are mixed populations of red mangrove and black mangrove occupying unstable protected substrates, produced by local erosion and Tertiary eroded platforms exposed to tidal action. Well developed mangroves are found on the low southern coast, where the rivers Dagua, Limones, and Anchicayá empty into the sea; these rivers carry sediments and create wide mud flats. The unstable zones are colonized by red mangrove, while the consolidated muddy-sandy substrates have forests of black mangroves and white mangroves. In this internal zone of the bay, with broad muddy beaches, there

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is no definite zonation of mangrove swamps, and intercalated populations of Rhizophora and Avicennia appear, with red mangrove dominant on recent more unstable borders. At Punta Soldado, a large sand bar develops, which encloses in its interior a complex system of tidal channels bordered by river mangrove swamps (Peña, 1998; Cantera and Blanco, 2001). This type of formation is preponderant along the entire southern coast, where there are large, well formed mangrove swamps, such as those of the estuarine mouths of the rivers Cajambre, Yurumanguí, Naya, Micay, Saija, Timbiquí, Guapi, and Iscuandé. These rivers have river mangrove swamps and empty into the sea by a network of tidal channels, with broad frontal bars or sandy beaches, such as those of Candelaria, which protects an extensive interior mangrove swamp that is continuous with natales and a cuanganal, which penetrate considerably up the Río Micay. This same pattern of zonation continues up to the mouths of the Ríos Guapi and Iscuandé. Starting at the Río Iscuandé, and continuing to the Río Patía, are the broadest mangrove formations of Colombia, influenced by the rivers Tapaje, La Tola, Amarales, Satinga, Sanquianga, and Patía. In this zone, the mangrove swamps are protected frontally by a wide complex of bars (bajos) and stabilized beaches (mulatos). Behind these formations there is an intricate network of tidal channels with inverted mangrove swamps, on islands and the borders of rivers. This zone is characterized by a large alluvial sedimentary platform, crossed by many firm sand bars, tidal channels, creeks, and tidal lagoons, which form estuarine or delta systems, where there are large bar, basin, border, river, and islet mangrove swamps. In some regions, upon old sand bars, moist tropical forest can be seen advancing toward the water. The mangrove forests extend to 35 km inland, depending on the maximum penetration of the tides. The mangrove swamp in these zones borders on large fresh water swamps, where there are forests on low hills dominated by species of trees known as Virola (Myristicaceae), Sajo (Campnosperma panamense, Anacardiaceae), Sande (Brosimum utile, Moraceae), and Cuángare (Otoba sp., Myristicaceae). This continues along the coast up to the zone of Salahonda, on the island of Gallo, and there it is interrupted by the presence of cliffs. The mangrove swamps reign on most of the northern coast of the inlet of Tumaco, although there are small mangroves in depressions flooded with brackish water, such as those of Curay (Sánchez-Páez et al., 1997). The internal part of the inlet of Tumaco, protected by El Morro and the island of Tumaco, an old stabilized sand bar, forms a wide border mangrove swamp, with internal basins and multiple tidal channels, such as those of Aguaclara and Las Varas. The borders of the tidal channels are covered by red mangroves, like a gallery forest, while the basins and elevated zones, with sandy-muddy substrates, are characterized by forests of Avicennia. Also, there are dwarf trees of Rhizophora in internal depressions covered with sandy soils. Apparently these soils do not retain nutrients, which are leached continually by strong rains; this causes poor development of these mangroves, which do not surpass 4 m in height.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

South of Tumaco appears a complex system of border and bar mangrove swamps, with sandy frontal beaches, such as those of Bocagrande, Papayal, and Güinulero. At the mouth of the Río Mira, toward Cabo Manglares, riverine forests form, dominated by Rhizophora racemosa. The mangrove swamps extend to Ancón de Sardinas and the Río Mataje, on the Ecuadorean border, where they occupy depressions flooded with sea water and low sedimented flats (Prahl, 1989; Prahl et al., 1990; SánchezPáez et al., , 1997).

PREHISPANIC CULTURES AND PRESENT COMMUNITIES The populations that inhabit the Pacific littoral are mainly blacks and native Americans. Their

settlement is ancestral and, for their historical records, information extends from the first pre-Columbian settlements with late archaeological evidence, up to documents of the history of the discovery, conquest, colonization, and the republican period (Sánchez-Páez et al., 1997). The archaeological and ethnohistorical record of the settlements of the Pacific coast indicate that the Tumaco-La Tolita cultural complex flourished between the fourth century B.C. and the third century A.D. from northwestern Colombia to extreme northwestern Ecuador. Some migrations reached the rivers Timbiquí and Guapi between the second century B.C. and the second century A.D.; this occupation continued during periods after the Spanish conquest. The Afrocolombian population is distributed along all the Pacific coast of Colombia, and the majority inhabits scattered villages constructed parallel to the coastline or on riverbanks. They seek spaces to plant coconut palms, which serve to mark the boundaries of family property. The other sector of the population that inhabits the Pacific littoral is formed by the native American communities Eperara-Siapidara, Emberá, Waunana, Cuna, and Awa-Kwáiquer, which inhabit the banks of rivers near the mouths or in the upper parts of the rivers. Small numbers of another population group, the mestizos (people of mixed race, native AmericanCaucasian), inhabit the most important urban centers. They carry out different economic activities, such as exploitation and transformation of timber resources, mining, commerce, and tourism. The mangrove swamps also provide a permanent refuge for fishermen, who obtain fish for the day, the minimum supply for their families, using small trammel nets, lines, wooden net supports, or hooks. The men catch several kinds of fishes, as well as camarón tití (small shrimps) and langostino (prawns). The women capture camarones (shrimps) and jaibas (crabs) using small nets or traps (catangas or esteradas), for domestic consumption or exchange for other products. Also (more often in some zones than in others) people enter the mangrove swamps to collect and remove molluscs and crustaceans (Sánchez-Páez et al., 1997).

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USES OF MANGROVE VEGETATION In mangrove forests, the population traditionally has used the wood for housing construction

or for boats and instruments; the wood also is used for fuel in the form of firewood or charcoal, and for obtaining medicinal substances. Moreover, the fauna associated with mangroves has been used as a primary source of animal protein. Exploitation and cutting of mangrove forests have been encouraged by the incessant demand for products for the construction industry or for infrastructure projects. In all coastal towns and mangrove zones, there is a history of foreign exploitation going back almost thirty years, and mangrove bark has been used for forty years before that (Sánchez-Páez et al., 1997) for the extraction of tannins used in tanneries, because of their preservative action on the proteins of animal skin (Sierra, 2001).

Mangrove forests also are used for posts for electrical cables, in construction, and as ties

for railroad tracks. These industrial uses continued for several years, until the governmental agency that was in charge of administration and management of natural resources (INDERENA), based on evidence of mass destruction by aquaculture that eliminated large areas of mangrove forests, suspended extraction of wood for industrial uses in the entire country in 1977 (Sánchez-Páez et al., 2003).

PRINCIPAL HUMAN IMPACT ON THE MANGROVE VEGETATION

OF THE PACIFIC COAST OF COLOMBIA Since the 1960s, the Colombian Pacific coastal region has undergone a process of concentration

of the population in urban centers. Turbo, Quibdó, Buenaventura, and Tumaco are centers that absorb population that migrates from the rivers that cross the Pacific zone. The greatest cultural wealth of the Pacific region is in the rivers and forests of the rural zone. It is there that the rural black and native American communities, which since a short time ago share their territory with mixed-race communities, have succeeded in creating ways of life that indicate a close relationship with nature (Montaño, 1998). The use of wood from mangrove forests, mainly for construction of boats, tools, houses, and for fuel, caused by the demand in the cities of the interior for establishing an economy based on commercial use of forests, has been the main factor in deforestation of mangrove zones (Sierra, 2001). The history of the Pacific coast of Colombia is included in the general history of the use of the natural resources of the country. This use was in many cases managed by the government, by means of granting licences to private companies or by tacit consent, which has been done in poor rural areas in which the inhabitants, who have no titles to their land, have participated only as unsalaried workers in extraction of resources.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

It has been calculated that in the last 20 years, about 57,276 ha of mangrove forests have been cut. On the Pacific coast of Colombia, about 8,400 ha have been destroyed (Sierra, 2001) (Table 1).

Table 1. Hectares of mangrove forest present in 1969 and 1996 in the departments of the Pacific coast of Colombia,

showing the number of hectares destroyed in this period (INVEMAR, 1998). YEAR

NARIÑO

VALLE

CHOCO

1969

44,660.48

154,617.52

42,306.45

64,852.18

1996

36,276.78

149,735.75

41,961.41

64,750.45

8,383.7

4,881.77

345.04

101.73

Difference

CAUCA

MANGROVES IN PROTECTED AREAS

The national natural parks on the Pacific coast in which mangrove forests are found are Sanquianga, Utría, and Uramba Bahía Málaga (Table 2). The delta of the Río San Juan and the delta of the Río Baudó also were designated recently to be included in the list of wetlands of international importance (Ministerio del Medio Ambiente, 2002). Table 2. Number of hectares of mangrove ecosystems in national natural parks on the Pacific coast of Colombia. PROTECTED AREA

TOTAL AREA

AREA OF MANGROVES

Ensenada de Utría

54,3

33,0

0.06

Uramba Bahía Málaga

47,1

4,6

9.93

Parque Sanquianga

80,0

58,5

73.18

The Parque Sanquianga (plate B: B) is the largest conservation unit that has protected mangroves (58,544 ha) on the entire Pacific coast of South America, which represent 73.18% of the total area of the park and 20% of all the mangrove forests of the Pacific coast of Colombia. In contrast, the inlet of Utría has only 33 ha, which is a very small mangrove forest. It should be noted, however, that Isla Gorgona, another important national natural park on the Pacific coast of Colombia, does not have mangrove forests, but it does have small groups of trees of Avicennia germinans, Conocarpus erectus, Laguncularia racemosa, and Rhizophora spp. Conocarpus erectus is the most abundant species. The recently designated national natural park Uramba Bahía Málaga has interesting river and border mangrove swamps, some of which grow on hard substrates.

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MANGROVE FORESTS OF ECUADOR

Xavier Cornejo and Carmen Bonifaz

Herbarium GUAY, Facultad de Ciencias Naturales, Universidad de Guayaquil, Ecuador.

xcornejoguay@gmail.com

INTRODUCTION

Ecuador is located on the Pacific coast of northwestern South America, bordered by Colombia to the north and Peru to the south and east. The coastline extends 950 km into both hemispheres, from 1°28’N 78°52’W to 3°23’S 80°18’W, and bears many fine sandy beaches, rocky cliffs, and mangroves. The mangrove forests are distributed in scattered patches along estuarine river mouth and deltas, forming huge extensions along the Gulf of Guayaquil, and in the northwestern most part of the country, between the Cayapas and Mataje rivers. The tidal amplitude often ranges from 2.5 to 3 m, reaching up to 5 m in the Gulf of Guayaquil. One hundred and seventy nine species of plants are estimated to occur in the mangroves of the country (Table 8). Rhizophora mangle, R. racemosa, R. x harrisonii, Laguncularia racemosa, Avicennia germinans and Conocarpus erectus are common shrubs and trees found in most mangrove forests. Other species as: Pelliciera rhizophorae, Tabebuia palustris, Mora oleifera and Pterocarpus officinalis are frequent trees restricted to the Chocoan mangroves, located at the northwestern part of the country. Mangroves dominated by Rhizophora are flooded daily (ca. 700 times per year), while those dominated by Laguncularia, Avicennia, and Conocarpus, are flooded half of the time (Bodero, 1993). Herbs commonly associated with mangroves are ferns of Acrostichum spp. Epiphytes such as orchids, bromeliads, araceae, and ferns can sometimes be found, depending on the availability of moisture being transported by air currents or birds/bats, and whether the mangrove is located in a moist to wet life zone. Associated fauna comprises dozens of species of birds, mammals, reptiles, fish, crustacean, molluscs, insects, etc…

TYPES OF MANGROVE VEGETATION

Three types of mangrove forests have been reported to occur in Ecuador (fide Lugo and Snedaker, 1974; modified by Cintrón et al., 1980), as follows:

Riverine forests

Riverine forests are present upstream along rivers following the intrusion of salty to brackish waters. In such environments, the freshwater and nutrient inputs are generally high, yielding forests of

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

well-developed structural characteristics. The water salinity changes seasonally, but the variability is generally less than that found in the remaining types of forests. The riverine forests encompass a total of 13,034 ha, and are mostly located in Guayas Province, along the Guayaquil Gulf and Guayas river.

Basin forests

Basin forests occur in landward areas near Salinas (salt pan areas). They can only be reached by the highest tides, and drain very slowly after remaining flooded for a certain period. The interstitial salinity is very high during the dry season, decreasing somewhat in the rainy season. Some physical parameters such as high salinity contribute to a lesser structural development of vegetation and limit the occurrence of some mangrove species. The trees and shrubs best adapted to these areas are Avicennia germinans and Conocarpus erectus. The basin forests encompass a total surface of 39, 028 ha. The largest areas of this type of forest are located in Esmeraldas Province.

Fringe forests

Fringe forests are subjected to daily flooding (over 700 times per year). Vegetation dominated by Rhizophora spp. has a high structural development due to large amounts of nutrients and sediments brought in by the tide. The fringe forests encompass a total surface of 125,388 ha. The largest stands of this type of forest occur in Guayas Province.

MAIN MANGROVE FORESTS IN THE COUNTRY

The largest areas of mangroves are located along river estuaries such as Mataje-SantiagoCayapas, Muisne, Cojimíes, Chone, Guayas, and Jubones-Santa Rosa-Arenillas. The Mataje-Santiago-Cayapas estuary is located on the northernmost coast of Esmeraldas province. It belongs to the wet to pluvial forests of Choco, and has an annual rainfall from 3,000 to 4,000 mm. In Ecuador, that area is the southern part of the Chocoan mangroves. Northwards, the mangroves of Mataje-Santiago-Cayapas estuary form a continuum to those of Nariño, in southwestern Colombia. The watershed encompasses 751,300 ha, and 21,400 ha are mangroves that belong to the Reserva Ecológica Manglares Cayapas Mataje (known as REMACAM; Clirsen, 2006). It is there that the most developed trees of the region can be found, with some red mangroves (Rhizophora) reaching 50 m tall and over 100 cm dbh, and some black mangroves (Avicennia) amazingly reaching 80 cm dbh (plates B: C; 5: E; 39: F). Floristically, these are the most diverse and luxuriant mangroves in the country, often exhibiting abundant epiphytes, and even containing new species, such as Sobralia

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rhizophorae, Platystele cornejoi, and Muellera chocoensis, the first two being discovered during the field work for this flora. San Lorenzo, Limones, and La Tola are the most important villages in the area.

The Cojimíes estuary is located along northwestern Manabí and southwestern Esmeraldas

Province; it encompasses 6,088 ha of a drainage basin of approximately 72,700 ha, which is shared with the Muisne estuary. Annual rainfall ranges from 2,000 to 3,000 mm. San José de Chamanga (simply called Chamanga) is the main village in the area.

The Chone river estuary is located at the center of the Manabí coastal line. The mangrove

encompasses 932,5 ha (Clirsen, 2006), and the annual rainfall ranges from 500 to 700 mm. It is surrounded by low hills (300 m max) and covered by a native vegetation of dry thorn scrub and very dry deciduous forests. Bahía de Caráquez and San Vicente, connected now by a bridge crossing the estuary, are the main towns.

The Guayas river estuary is located in the province of Guayas. The watershed encompasses

3’523,800 ha, feeding the Gulf of Guayas the largest gulf and estuarine system on the Pacific coast of South America. The Guayas mangrove encompasses 105,130.4 ha at low tide (Clirsen, 2006), subjected to a daily tidal range from 2.5 to 3 m, reaching up to 5 m. The tidal influence reaches up to 80 km north of Guayaquil, and nearly 40 km during the rainy season, through the Daule and Babahoyo rivers. The average annual rainfall ranges from 500 to 1,300(-1500) mm. Guayaquil, the main city in the area, is also the largest and most populated city in the country.

The Jubones-Santa Rosa-Arenillas estuary is located in the El Oro province in southwestern Ecuador.

It forms a continuum to the Tumbes mangroves in northwestern most Peru. The Jubones-Santa RosaArenillas mangrove encompasses 15,207.6 ha (Clirsen, 2006), and has an annual rainfall range between 400 to 600 mm.

Other mangroves of smaller surface area can be found at many small river mouths

located throughout the coast of Ecuador, such as Calope, Rio Verde, Montalvo, Atacames, Jama, Tonchigue, Pedernales, Olon. All these smaller mangroves encompass a total area of 488 ha.

FAUNA

The fauna of Ecuadorian mangroves is represented by 52 species of birds, 15 spp. reptiles, 19 spp.

mammals, approximately 100 spp. fish, 20 spp. crustaceans, and 70 spp. molluscs (Bodero, 1993; Salas, 2010).

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Mangrove soil is a unique habitat that is also a source of food for many crabs, snails, and bivalves. Among these, the most common resident species is the cangrejo rojo or red crab, Ucides ucidentalis (Ucididae; plate C: D), the largest and commercially most important crab that is sold locally for food. The population of well-developed and abundant individuals of Ucides ucidentalis is directly correlated to the presence of well-developed stands of red mangrove trees (Rhizophora spp.; Cornejo, 2005). The cangrejo azul or blue crab, Cardisoma crassum (Gecardinidae), builds burrows at the higher and drier areas at the back fringes of mangroves. The omnivorous crab is locally recognized by the conspicuous blue color of its cephalotorax. The fiddler crabs or violinistas, Uca spp. (Ocypodidae; plate C: C), frequently present at lower tides on channel banks and mud flats, and the spider crab, locally known as tasquero or cangrejo araĂąa, Goniopsis pulchra (Goniopsidae), which has a spider-like appearance while climbing trees, are crabs of smaller size that also build mud holes as burrows. These crabs are not captured for food by fishermen, but are hunted by several natural predators such as birds, e.g. the American white ibis, locally known as garza cangrejera, Eudocimus albus (Threskiornithidae), and the crab-eating raccoon, locally known as tejon or osito lavador, Procyon cancrivorous (Procyonidae). Hermit crabs as Clibanarius sp. (Diogenidae) are also occasionally present on mangrove soils. Cerithidea mazatlanica (Potamididae) is a common snail that is often easily visible, abundant as it is on the mud flats located in the intertidal zone at lower tide. Other representative mangrove snails are the lame platos, Natica unifasciata (Naticidae), an edible gastropod that has been observed on the mud of the Rio Chone estuary, and Littorina spp. (Littorinidae; plate C: B), which could be found on the ground, otherwise climbing red mangrove prop roots or on the branches of white and black mangroves. The concha prieta, Anadara tuberculosa (plate C: A), the concha macho or mica, Anadara similis, and the pata de mula, Anadara grandis (all Arcidae), are the most common mangrove cockles that are sold locally for food. These bentonic and often dominant bivalves at larval stage are carried out by the currents until they settle into the soft muddy soils under shallow brackish waters. Oysters such as Crassostrea columbiensis (Ostreidae), which grow adhered to Rhizophora prop roots, mangrove mussel, Mytella guyanensis (Mytilidae), and clams as Protothaca spp. and Chione subrugosa (both Veneridae), which inhabit beneath the muddy ground surface right under to nearby the trees (Cruz, 2003), are also intensively harvested intertidal bivalves of high nutritional value, and represent an important economical source for fishermen. Many other aquatic species, many of commercial importance, visit mangrove estuarine waters in order to obtain food, shelter, and breeding. As an example, it is worthy to mention the whiteleg shrimp, Litopenaeus vannamei (Penaeidae), one of the most common and delicious crustaceans produced in shrimp farms throughout tropical America. Since the 1970â&#x20AC;&#x2122;s, the whiteleg shrimp represents one of the

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main income sources for the country. Related crustaceans of commercial value are: The western white shrimp, L. occidentalis (Penaeidae); western blue shrimp, Litopennaeus stylirostris (plate C: E); and marine crabs of flattened bodies and paddle-like posterior legs, adapted to swim and dive as jaibas, Callinectes arcuatus and C. toxotes (Portunidae). Additionally, an endless number of fish, many of those regarded as the main source of income for thousands of fishermen, are visitors or residents of mangrove waters. Some of these are: The lisa, Mugil cephalus (Mugilidae), the corvina, Cynoscion albus (Sciaenidae), the robalo, Centropomus spp. (Centropomidae), the pargo, Lutjanus sp. (Lutjanidae), the catfish (Ariidae; plate C: F), etc.

Following the presence of countless invertebrates and fish, dozens of bird species are also found

in mangroves, as they find there abundant food sources. The most common mangrove birds are egrets and herons (Ardeidae), the same as are found in fresh water wetlands, such as the great white egret or garza blanca grande, Ardea alba; cocoi heron or garza real, Ardea cocoi; white egret or garza blanca pequeĂąa, Egretta thula; little blue heron or garza morena, Egretta caerulea; striated heron or chuque, Butorides striatus; and yellow-crowned night heron or guaque, Nyctanassa violacea, a bird of mostly nocturnal habits. Other common birds are: The neotropic cormorant or patocuervo, Phalacrocorax brasilianus (Phalacrocoracidae), a voracious shrimp predator regarded by shrimp farmers as enemy number one; American white ibis or garza cangrejera, Eudocimus albus (Threskiornithidae) can be often found on mud flats chasing Uca crabs; Roseate spoonbill or espatula rosada, Platalea ajaja (also Threskiornithidae) can be seen swinging side to side the elongate and dorso-ventrally flattened bill in shallow waters; common black hawk or gavilan cangrejero, Buteogallus anthracinus (Accipitridae), another crab predator that is frequently found standing on tree branches while gazing down at the muddy ground while hunting; osprey or aguila pescadora, Pandion haliaetus (Pandionidae), fish feeders; and the omnivorous greattailed grackle or negro manglero, Quiscalus mexicanus (Icteridae). Common sea birds that build nests in mangroves are: The magnificent frigatebird or fragata, Fregata magnificens (Fregatidae); and the pelecan or pelicano, Pelecanus occidentalis (Pelecanidae).

Most mammals found in mangroves are visitors searching for food or protection. The most

common of these are: The crab-eating raccoon, known as tejon or osito lavador, Procyon cancrivorous (Procyonidae), which feeds on crab and fish; deer, Odocoileous virginianus (Cervidae), which visits mangroves looking for refuge; felids, as the jaguarondi, Puma yagouaroundi, and tigrillo, Leopardus tigrinus (both Felidae), which often hunt the crab-eating raccoon and young deer; and opossums such as Didelphis masurpialis (Didelphidae). In the canopy, monkeys such as the mischievous Ecuadorian capuchins or micos, Cebus albifrons aequatorialis (Cebidae) can sometimes be found. They visit mangroves to feed on bird eggs and predate juveniles or even adults of red crab Ucides occidentalis that have been left tied up by fishermen (Fernando CedeĂąo and Byron PeĂąafiel, pers. comm.). Swimming mammals are Neotropical otter or nutrias, Lontra longicaudis (Mustelidae), and the common bottle-nose

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

dolphin, Tursiops truncatus (Delphinidae) that feed on fish. At night, many insectivorous and fishing bats can be found flying in the mangrove forests and along watercourses. The following bats are reported to occur in Ecuadorian mangroves: Artibeus lituratus, A. fraterculus, Desmodus rotundus, Diclidurus albus, Glossophaga soricina, Molossus molossus, Myotis nigricans, Noctilio leporinus subsp. mastivus, and Rhogeessa velilla (Moscoso and Tirira, 2009; Salas, 2010). It is very likely that many more bat species are waiting to be recorded for the Ecuadorian mangroves. Among reptiles, it is worthy to mention: The green iguana, Iguana iguana (Iguanidae), which feeds on buds and young leaves of Avicennia germinans; snakes such as the boa, Boa constrictor (Boidae), that can predate incubating females, eggs, and nestlings; and the American crocodile or Tulisio, Crocodylus acutus (Crocodylidae), which feeds mainly on fish. The loss of crocodile populations in the mangroves of Guayaquil and the role of the leaf-eating mangrove crabs have been the topics for local songs, as ¨El guardian del manglar¨ and ¨Cangrejo de manglar, available in this link: https://soundcloud.com/kareen-mendoza

PREHISPANIC CULTURES AND PRESENT COMMUNITIES

Correlating the dates of the earliest fossil pollen of red mangrove (Rhizophora spp.) in Ecuador that is recorded from the Early Holocene off the Peninsula de Santa Elena (Heusser and Shackleton, 1994) against the archaeological data, it is certain that the Las Vegas (between 10,840 and 6,600 RCYBP), one of the most ancient preceramic cultures in South America, flourished in the new Holocene environment right after the settlement of the early mangrove stands in the coast of Ecuador. During the Early Holocene the littoral of Ecuador was approximately 30 m lower than today. The sea level rose and fluctuated under the then oscillating environmental conditions allowing the creation and destruction of mangrove swamps due to continue marine transgression. Las Vegas people then ¨adapted successfully to the changing environments in the Late Pleistocene and Early Holocene periods¨ (Stothert, 2011). Consequently, the mangroves turned out to be an important and one of the main sources of food for them. Moreover, it is not only that some mangrove species became a significant element of their diet, some of those species became also a part of their belief, as it is the case of cockles of concha prieta (Anadara tuberculosa; Stothert, 2011; plate C: A). After Las Vegas, other prehispanic coastal cultures that in similar life style settled and lived for centuries in the lowlands of western Ecuador, along river banks and deltas, developed subsistence activities such as agriculture, hunting, and also fishing in mangroves and associated environments. Some of these are: Punáes, Huancavilcas, Chonos, and Valdivia in Guayas and Santa Elena Provinces; Caras, Machalilla, Manteño, Chorrera, Jama-Coaque, the señorío de Salangome, and Salango in Manabi; and Atacames and La Tolita in Esmeraldas (DIGMER, 1994).

Xavier Cornejo, editor - 2014

Regarding to prehispanic native American localities settled behind or near mangrove swamps,

it cannot be discarded -as it is hypothethyzed here- that the protector barrier effect that mangrove soils and vegetation provide sometimes may had influenced the choice for their settlement place. That ecosystem service may have been used to avoid unfriendly paleoindian or prehispanic visitors or pilmgrims that were arriving in groups sailing along the Pacific coast.

Nowadays, in the lowlands of western Ecuador, the Chachis is the main surviving Native American

ethnic group. The Chachis, commonly known as Cayapas, live along the Cayapas river downstream to the estuarine river mouth in Esmeraldas province. They subside on mangrove products, and are socially and economically close to Eperas and Afroecuadorians. The Epera is a small ethnic group with its own language, having migrated from southwestern Colombia into the coast of northwestern Ecuador, where they have been officially recognized since 1993.

Most populations that inhabit coastal areas in and near the mangroves of northwestern Ecuador

are Negros brought to the Americas from Africa for slavery. Accordingly to oral tradition, earliest settlement was carried out by groups of fugitive slaves searching for refuge and freedom during the XVI century. Many Afroecuadorian descendents still inhabit scattered villages along the coastal line or along riverbanks, and have developed subsistence activities such as hunting wild fauna and growing coconut and plantain trees, while also eating mangrove fish, bivalves, crab, and shrimp (plate D: A).

The remaining Pacific coastal areas in and near mangroves are inhabited by mestizos, who carry

out various different economic activities. Many have the traditional habit of visiting mangroves for fishing, by the use of small trammel nets, lines, hooks or hoops (plate D: B). In addition, they capture cockles, shrimp, and crabs for domestic consumption as well as to be sold or exchanged for other products.

SOME HISTORICAL DATA

Early references about mangrove ecosystems and vegetation deserve to be mentioned. During

the XVI century, the Spanish conquerors found an impressive and formidable mangrove forest along the Pacific coast of South America. However, the mangrove swamps represented such a nuisance to troops and horses that Cristóbal de Molina described them in 1552 as the “most difficult land of these kingdoms” (Alvarez-León, 1993).

In 1684, William Dampier, a British buccaneer and explorer, visited the coasts of Ecuador, and

reported that both sides of the Guayas river were thickly covered with strands of red mangrove (Rhizophora spp.; Dampier [1697] 1968; Madsen et al., 2001). This is the earliest reference of mangrove plants for the

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

country. The earliest botanical collection in Ecuadorian mangroves was gathered by the Czech explorer and botanist Thaddäus Haenke, in October of 1790. As part of the scientific staff of Alejandro Malaspina’s expedition around the world (1789–1794), Haenke visited Guayaquil on the corvette Descubierta, collecting the type of Laguncularia racemosa var. glabriflora among many other specimens found along the Daule river. In the late XIX century Danish botanist Baron Von Eggers described the mangroves of the Gulf of Guayaquil, recording that the old part of Guayaquil city was built up very largely of mangrove wood (Eggers, 1892), and reporting Rhizophora trees reaching 50 m high from Puna Island (Eggers, 1894).

In 1904, Henry Brougham Guppy (1854–1926), a British naturalist, visited the coasts of the

country guided by Eggers’s earlier references and the map of Ecuador elaborated by Dr. Wolf (1892). He sailed through the Gulf of Guayaquil, but did not record a detailed description of the mangrove formations, as these were “indeed made by Eggers”, focusing instead on describing the mangroves of El Oro province, detailing their appearance and floristic composition at a rather generic level. Additionally, he made some interesting observations about plant seeds, fruits and seedlings being transported seawards by currents, finding seedlings of Rhizophora, Avicennia, germinating fruits of Laguncularia, Salicornia, and seeds of Talipariti tiliaceum from ten to twenty miles south of the mouth of the Guayaquil Gulf estuary. The salt pan areas found in the Ecuatorial mangroves much impressed him, who apparently was the first to describe them as “arid saliniferous plains” (Guppy, 1906).

After World War II and mainly during the past four decades the studies of mangroves have become

more intense and many botanists and ecologists have visited the country for research.

USES OF MANGROVE VEGETATION

Mangrove wood of all species has been used for centuries, mainly for construction as columns, and

as firewood. During the post-Columbian period, the rise of the shipbuilding industry created a demand for hardwoods, which brought mangrove trees to prominence, causing the red mangroves (Rhizophora spp.) to be among the most heavily exploited species for local uses as well as for exportation to Lima. Since that time, the red mangrove timber has been used for house construction, walls of fortification, boards, boats, ribs of ship frames, and keels of vessels due to its incorruptibility and high resistant to marine water and because of the elongated and straight trunks that can reach more than 30 meters in length; the branches have been used to produce charcoal. Because of the presence of tannins used for tanning leather and dye, bark exploitation of red mangroves officially began in 1879, increasing in productivity throughout the years and reaching a peak during World War II, mainly in the Gulf of Guayaquil. Avicennia germinans and Laguncularia racemosa have traditionally been used for the construction of house columns, canoes, stakes, fences, nets, tool handles. Additionally, they have been used as charcoal (Madsen et al., 2001).

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PRINCIPAL HUMAN IMPACT ON THE MANGROVE VEGETATION OF THE COAST OF ECUADOR

The main human impacts on mangrove vegetation and ecosystems during the past century in Ecuador have been regarded as environmental and social (Bravo, 2010). The main environmental impacts are: The expansion of the agricultural frontier, cattle farming, the uncontrolled growth of cities, towns and villages, and the unsustainable massive expansion of industrial shrimp farming. The expansion of the agricultural frontier against mangroves date back to 1940, with the establishment of 10,000 ha of coconut palm (Cocos nucifera) over mangrove ecosystems in the province of Esmeraldas. The trees were felled and the soils suffered severe alteration during land preparation, introduction of cultivated coconut palms, and the construction of artificial drainage canals. The most affected areas were riverine mangroves and landward creeks (Bodero, 1993). Mainly since the 1960’s, over 40,000 ha of mangrove forest have been converted into cattle farms. The basin forests have been the most affected by this activity. In these areas, the vegetation has been felled and several grass species tolerant to brackish environments have been planted, causing the soils to suffer irreversible structural modifications. In the early 1970’s, Ecuador started intensive shrimp farming, locally called the shrimp aquaculture boom. Large surfaces of mangrove ecosystems were converted into shrimp ponds, causing a massive loss of habitat, producing short-term profits at the expense of long-term productivity. Shrimp farms were first built up on salt pan areas, then later over mangroves, and finally on slightly higher terra firme agricultural lands. The main shrimp species cultivated since that time are Litopenaeus stylirostris (plate C: E) and L. vannamei, and shrimp exportation became one of the main income sources for the country. Ecuador has been the largest shrimp supplier to the United States and one of the largest maricultural producers in the world. The golden age of shrimp exploitation in Ecuador was during the 1970’s and 1980’s. It has declined in productivity since the 1990’s to the present, because lack of an adequate integrated planning and the excessive and uncontrolled shrimp farm expansion without consideration of environmental cost. To the present, this has been the most devastating impact against mangrove ecosystems in the country (Table 3); the most affected area was the Gulf of Guayaquil, in the province of Guayas.

Table 3. Hectares of mangrove forests versus shrimp farms surface recorded between 1969 and 2006.

Mangroves

1969

1984

1987

1991

1995

1999

2001

2006

203,624

182,157

175,157

162,186

146,938

149,556

154,087

148,230

89,368

117,728

145,998

178,071

207,000

Shrimp farms

Source: CLIRSEN, 2006.

170,648

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

In addition, cities such as Guayaquil and Machala, as well as several coastal towns and villages have been partially expanding over mangrove areas while enlarging their boundaries, building up houses and neighbourhoods from above the high tide level to even into the intertidal zone, in what were riverine and basin mangrove forests. Thus, several to hundreds of rustic small houses whose walls are made of bamboo have been built up on wood columns raised over the water surface, interconnected by narrow elevated wooden walkways. The construction style reflects that of the typical fishermen’s houses that can be seen along the coast (plate D: A). In these areas, the woody vegetation has been felled, and the creeks and tide canals closed, blocking water flow and drainage. This has caused formerly muddy loose soils to be compacted, filled with rocky soil borrowed from hills and terra firme ecosystems inland. In this way, it has been estimated that a surface of 3,000 to 5,000 ha of mangroves have already been converted into urbanized areas (Bodero, 1993). The social impact refers to the growing unemployment and displacement of ancestral land owners that have managed mangroves during decades before the shrimp farm massive expansion, resulting in the increasing of poverty, multiplying conflicts among fishermen in regards to the diminishing resources and immigration (Bravo, 2010). Nowadays, mangrove trees and the whole mangrove swamp ecosystems are protected by Ecuadorian laws such as the Ley Forestal y de conservación de áreas naturales y vida silvestre, L. 74, Reg. Of. 64, since 1981; since 1981; Decreto Ejecutivo No. 3327, Reg. Of. 848, since 1995; and the Ley reformatoria a la ley forestal y de conservación de áreas naturales y vida silvestre, L. 91, Reg. Of. 495, since 1990. These laws declare mangroves and its goods (that means molluscs, crutaceans, fish, wood, etc…) to be a state property.

MANGROVES IN PROTECTED AREAS

The following protected areas that contain mangroves have been created in order to preserve their ecosystems. These belong to the Ecuadorian state, and are administrated by the Ministerio del Ambiente (MAE) through the Patrimonio de Áreas Naturales del Estado (PANE): 1. Reserva Ecológica Manglares Cayapas Mataje (REMACAM). It is located at the northwesternmost part of the country in the province of Esmeraldas, reaching the Ecuador-Colombia boundary at the Mataje River. The REMACAM encompasses 17,955.54 ha of mangroves. 2. Refugio de vida silvestre manglares del Estuario del río Esmeraldas. It is located at the estuary of the Esmeraldas River, in the province of Esmeraldas. This protected area encompasses 242.58 ha of mangroves and abandoned shrimp ponds that are in the process of rehabilitation.

Xavier Cornejo, editor - 2014

3. Refugio de vida silvestre manglares del Estuario del río Muisne. It is located at the estuary of Muisne River, in the province of Esmeraldas. This protected area encompasses 3,173 ha of mangroves. 4. Refugio de vida silvestre Islas Corazón y Fragatas. It is composed of two close islands located at the estuary of Chone River, in the province of Manabi. This protected area encompasses 2,811.67 ha of mangroves. 5. Refugio de vida silvestre manglares El Morro. It is located at the mouth of the Gulf of Guayaquil, approximately 15 km northwest of Puna Island, in the province of Guayas. This protected area encompasses 10,130.16 ha of mangroves. 6. Reserva de Producción Faunística Manglares El Salado. It is located at the interior of the Gulf of Guayaquil, in the city of Guayaquil, province of Guayas. This protected area encompasses 10,635.12 ha of mangroves. 7. Área Nacional de Recreación Isla Santay y Gallo. A RAMSAR site located in the province of Guayas, on the Guayas River, at the east of Guayaquil. The Santay Island encompasses a total of 2,179 ha, containing several ecosystems such as dry forest, fresh water wetlands, and mangroves. 8. Reserva Ecológica Manglares Churute (REMCH). It is located at the interior of the Gulf of Guayaquil, in the province of Guayas. The REMCH encompasses a total of 50,000 ha, containing several ecosystems such as terra firme dry and moist forests, seasonally fresh water wetlands, and approximately 35,000 ha of mangroves. 9. Reserva Ecológica Arenillas. It is located at the southwestern part of the province of El Oro, between Arenillas and Huaquillas, near the Ecuadorian-Peruvian frontier. This protected area encompasses a total of 17,000 ha, containing several ecosystems such as terra firme dry forest, dry thorn scrub, and 2,800 ha of mangroves.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

MANGROVE FORESTS OF PERU Reynaldo Linares-Palomino Herbario Forestal MOL, Universidad Nacional Agraria La Molina, Apartado 456, Lima 1, Peru. reynaldo.lp@gmail.com

Marisela Monzón-Ramos Universidad Nacional de Piura, Peru.

INTRODUCTION

The mangroves of Peru are located in the extreme northwestern tip of the country, and constitute the southernmost location of mangroves on the Pacific coast of America (REF). Most of the mangroves are concentrated in Tumbes, with a small and isolated fraction in Piura. Little information is known about this ecosystem, and what has been published refers mainly to the two conservation areas that have been setup within it: the Los Manglares de Tumbes National Sanctuary (SNLMT), in Tumbes, and the Municipal Conservation Area Manglares San Pedro de Vice in Piura. Much of the information for this chapter has been extracted from an extended study of the Tumbes mangroves (INRENA, 2007). The Peruvian mangroves grow in an arid climate, influenced by the oceanic system. Climatic condition are generally similar at both mangrove systems, with mean temperatures of 23 ºC, mean annual precipitation of 100–300 mm, and a mean monthly relative humidity of approximately 70%. Rains falls generally between January and March, the rest of the year being rainless except for extraordinary rainfall events such as those that occur during El Niño years (Ortlieb and Macharé, 1993). Soils are generally dark sandy, and in few places occasionally sandy-loamy and calcareous, salty and strongly acidic, or basic. These soils have been classified as badlands, with no potential for agriculture or cattle production. Aquacultural activities are developed in Tumbes, but not in Piura.

Both mangrove systems are heavily dependent on the confluence of the warm waters of the El Niño current, flowing north to south, and on the major rivers in the region. The Tumbes mangroves are influenced by the Zarumilla and Tumbes rivers in Peru, and the Jubones and Arenillas rivers in Ecuador. The Piura mangroves are influenced by the Piura river and the Sechura drainage. Additionally, sea water enters through the estuaries into both mangrove systems. The flux and reflux or hydro-period happens every six hours with low and high tides. Sea water enters at the beginning of the high tide through the estuaries, bringing nutrients with it and allowing the migration of species. When the low tide sets in,

Xavier Cornejo, editor - 2014

sediments, excess of salts, and leaf litter are redistributed into the open sea (INRENA, 2001). The San Pedro mangroves deserve a special mention, as the flow of the Piura river and the Sechura drainage is seasonal and restricted to the rainy season between January and March, and to exceptional El Niño years. The rest of the year these systems are practically dry. Thus, the San Pedro mangroves experienced an increase in area after the 1993 and 1998 El Niño years and have since shrunk slowly, but permanently, because the necessary continental water supply that initially set up the system is no longer available.

The flora of the mangrove ecosystem is composed of plants that are typical of these systems and also of plants belonging to contiguous vegetation formations, especially those of the halophytic and dry forest communities. The SNLMT reports 41 species (22 families and 36 genera) (INRENA, 2007). The San Pedro mangroves report 44 species (21 families and 36 genera) (Charcape-Ravelo and Moutarde, 2005), with the two mangrove species Laguncularia racemosa (mangle blanco) and Avicennia germinans (mangle prieto).

TYPES OF MANGROVES VEGETATION The types of mangroves vegetation that occur in Peru are similar to those that occur in Ecuador.

That means, riverine, basin, and fringe forests are present, but having the tendency of a lesser structural development of the forests.

DISTRIBUTION OF MANGROVE VEGETATION ON THE PACIFIC COAST OF PERU The Tumbes mangroves are located between 3º24’ (on the Ecuadorian border) and 3º35’ S

(ONERN, 1992) and cover between 4,000 and 6,000 ha., depending on the year and assessment method. The entire mangrove ecosystem, including associated vegetal formations, is estimated to cover between 25,000 and 28,000 ha. (ONERN, 1992). The Tumbes mangroves can be separated into four vegetation formations and three ecotones (INRENA, 2007). The typical mangrove vegetation in Tumbes is dominated by Rhizophora mangle (locally known as mangle rojo), the main structural component, with an important presence of Rhizophora racemosa and R. x harrisonii (mangle colorado). These species form on a thin mud belt along the continental-tide canal ecotone. Landwards, on wet sandy soils, follows the manglar-gramadal ecotone, which is usually dominated by Laguncularia racemosa (mangle blanco) and accompanied by Avicennia germinans (mangle salado), Batis maritima (vidrio), Sesuvium portulacastrum (verdolaga salada) and Salicornia fruticosa. Further inland, two additional formations appear; hypersaline flooded lands, barely vegetated, and areas covered by Distichlis spicata (grama salada), on higher grounds and locally called gramadal, sometimes associated with Sesuvium portulacastrum and Salicornia fruticosa.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

The San Pedro mangroves (plate A: D), located between 5°30’–5°34’S cover some 3,000 ha, including the adjacent Prosopis sp. forests (locally called Algarrobales). The reason mangroves are not found further south on the South American Pacific coast has been found to be the presence of the cold Peruvian current, flowing parallel to the coast, south to north until 6º, which creates cold climatic conditions that impede the development of mangroves (Clüsener, 1987; Clüsener and Breckle, 1987). The San Pedro mangroves in Piura are almost exclusively dominated by Avicennia germinans (mangle prieto) which inhabit muddy and humid substrates with high amount of detritus. It is interesting to note that Rhizophora is absent (Charcape-Ravelo and Moutarde, 2005). The gramadal, close to the mangroves, comprises species such as Prosopis pallida (algarrobo), and Acacia macracantha (faique) (Rodríguez et al., 1997).

FAUNA The fauna in the mangroves is very diverse, with some taxonomic groups especially important because of their socio-economic value. The only exception are amphibians, which are not representatives in this system. Faunal information for the mangroves as a whole ecosystem is not available, only species lists for both conservation areas. In the Manglares de Tumbes national sanctuary there are 148 species of birds, 6 reptiles, 5 mammals, 105 fish, 16 crustaceans, 74 pelecipods, 69 gastropods, 3 echinoderms, 4 annelids, and 21 insects (INRENA, 2007). In the San Pedro mangroves, has been reported 98 bird species, six species of mammals, eight species of reptiles, 23 species of fish, 11 crustaceans, and 60 protozoa (Barrionuevo and Marcial, 2006, Chávez, 2007).

PREHISPANIC CULTURES

Human presence along the northern Peruvian coast has been documented by Hocquenghem (1998). There is evidence that nomad hunters and gatherers used the mangrove ecosystem as a permanent source of food as early as 10,000 BC. It was also an important resource for the civilizations that later settled the region, most importantly those of the Tumpis culture. After the Spanish conquest, use of the mangrove resources by local people continued, basically for self-consumption (Takahashi, 2002; Moya, 2006).

USES OF MANGROVE VEGETATION

Both mangrove systems are locally very important in economic and social terms. Local people use them directly or indirectly to extract goods for sale, and take advantage of them for being a tourist attraction. However, the main economic activity is small scale fishing and extraction of hydro-biological resources.

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PRINCIPAL HUMAN IMPACT ON THE MANGROVE VEGETATION OF THE COAST OF PERU

The main threat of the Peruvian mangroves is deforestation, mainly caused by productive activities. Historical photographic studies show the shrinking of mangrove vegetation from 5,964 ha in 1982 to 4,541 ha in 1992 (ONERN, 1992). A more detailed study showed that of the 1,791 ha that disappeared between 1982 and 1992, 1,294 ha were caused by deforestation to build new shrimp farms (Caccha, 1999). Another threat, the magnitude of its impact not yet assessed, is the blocking of water flow in the estuaries by the construction of ponds for shrimp farms and pumping stations. These activities change the hydro-period in the estuaries, potentially affecting the ecology of the ecosystem (CDC-UNALM, 1986). Migrant populations can triple the permanent population of the small villages located close to the mangroves (Takahashi, 2002). These people, with little knowledge of the workings of the mangrove ecosystem, do not value it and usually perform destructive extraction of its resources. In the San Pedro mangroves, disturbances by recreational activities (access to the beaches, waste contamination) have also been identified as threats.

MANGROVES IN PROTECTED AREAS

Conservation areas have been set up in the Piura and Tumbes mangroves: Los Manglares de Tumbes is a National Sanctuary, included in the protected areas system of Peru and managed by the National Protected Areas Service (SERNANP). The Municipal Conservation Area Manglares San Pedro de Vice in Piura is managed locally by the council of the city of Vice. Additionally, both areas have been recognized as RAMSAR sites.

Plate A: Major mangrove stands and root adaptations, respectively: A–B. Rhizophora. C–D. Avicennia. E–F. Laguncularia.

CHAPTER II.

MANGROVE PLANTS by Xavier Cornejo

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

PLANT HABITS, DIVERSITY AND DISPERSAL METHODS

Plant species commonly exhibit defined habits. The habit of a plant refers to its growth form, whether

it be a tree, shrub, liana, epiphyte, parasite, or strangler. Mangrove and associated plant species are herein gathered and discussed in their respective habits and dispersal method of fruits and/or seeds (Figs. 2, 3).

Trees and shrubs The common definition of a tree is a woody perennial plant, taller than 2.5 m, characterized by a

more or less elongate stem that is unbranched in its lower part. In this work we expand the definition of tree to include species of arborescent appearance but with non woody stems, such as palms and a Zamia.

Shrubs are woody and subwoody perennial plants, usually less than 2.5 m tall, which are always

branched from or near the base.

The same species can sometimes present both tree and shrub habits, even in the same locality.

For example, plants of some species may have a shrubby appearance during their early years, even upon reaching sexual maturity (producing flowers and fruits), but with further development may become a well defined tree. This is the case for all major mangroves species in our area (Rhizophora spp., Avicennia germinans, and Laguncularia racemosa), and for some species of the remaining woody flora. Thus, for the purpose of analyzing the percentages of mangrove plants per habit, trees and shrubs have been merged into the same category.

In regards to the plant species of all groups, this category comprises 15 families (Table 4). Each

family in the category contains one to three species only. The dicotyledons are predominant among trees and shrubs, representing the 90 % (27 spp.) of the species of this habit. The only mangrove monocotyledon family that contains arborescent members is Arecaceae, represented by only two species of palms, of which only Euterpe oleracea is native. Gymnosperms is solely comprised of Zamia roezlii, a Zamiaceae No arborescent species of peridophyte is recorded to inhabit the mangrove interior.

In the present work, 30 species of trees and/or shrubs have been recorded, including the

introduced coconut palm, Cocos nucifera. From the 29 native species, only seven have an ecologically wide distributional range, inhabiting mangroves located in the rainforest, wet forest, and moist forest life zones, and also the dry, very dry, and thorn woodland life zones. Geographically, from the Choco Department in Colombia, and southwards to northwestern Peru, those species are: All major mangroves, Rhizophora mangle, R. racemosa, Avicennia germinans, and Laguncularia racemosa; and some facultative mangroves, Conocarpus erectus, Talipariti tiliaceum var. pernambucensis, and Annona glabra. The cited major mangroves are the common tree and/or shrub elements, widely distributed in almost all mangrove

MANGROVE PLANTS

ecosystems in our area. There are very few exceptions to this pattern, such as the absence of Rhizophora in the Manglares de San Pedro, Peru (Charcape-Ravelo and Moutarde, 2005), a small isolated mangrove ecosystem located right in the southernmost corner of the entire mangrove distribution range along the Pacific coast of the American continent. Whether the natural absence of Rhizophora in the Manglares de San Pedro is a latitudinal effect or a response to oceanic water currents is unknown, and deserves further research.

Seventeen native species of trees and shrubs (including the minor mangroves, Pelliciera

rhizophorae and Tabebuia palustris) inhabit only those mangrove ecosystems located in the wet forest and rainforest life zones, which range between the Choco Department, Colombia and northwestern Ecuador. Most species of this group also occur northwards along the Pacific coast (and sometimes also on the Atlantic coast) of Mesoamerica, in mangroves located in wet forest and rainforest life zones. None of those species extend their populations into the drier life zones located southwards on the Pacific coast, from the Province of Manabi in Ecuador to northwestern Peru. This pattern of distribution, restricted as it is to the wetter life zones (1500â&#x20AC;&#x201C;6000 mm) suggests that the drier life zones and consequent higher levels of soil salinity, which result in lesser precipitation (below 1000 mm) and water runoff, act as an ecological barrier. Another important point is the fact that the Peru (or Humboldt) current (which aids in forming the drier life zones), flowing north along the western coast of South America, does not allow southwards water dispersal for the northern woody species.

The most prevalent dispersal method of fruits and/or seeds among mangrove trees and shrubs is

via water (17 spp., Table 5). The second major diaspore dispersal syndrome is via birds and presumably bats (9 spp.). The drupaceous or baccate fruits of a few species have been recorded to be dispersed by other vertebrates and fish (e.g., Euterpe oleracea; Moegenburg, 2002; Pedersen 104001, AAU); however, the number of plant species that have fish dispersed fruits and/or seeds in mangrove swamps is still unknown.

Among trees and shrubs, Cosmibuena macrocarpa, Hillia maxonii, and Rustia occidentalis

are the only species adapted to wind dispersal in our area, all of which are comprised by the Rubiaceae family.

Herbs

Herbs are non woody plants of less than 4 m tall, e.g., the terrestrial ferns of Acrostichum

spp. This habit is represented by 29 spp. (Table 4). Monocots is the richest group in this habit (19 spp.), predominantly represented by Cyperaceae (10 spp.) and Poaceae (4 spp.). Both families comprise 73.7 % of all monocot terrestrial herbaceous diversity, and 48.2 % of all terrestrial mangrove herb species herein recorded; their floristic elements are rather infrequent in disturbed hyposaline ecosystems, however.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

The pteridophyte genus Acrostichum, comprised of two species, is the most common terrestrial

herb, inhabiting disturbed areas in the mangrove interior, landward fringes, and also the supratidal zone on adjacent river banks. Among dicots, salt tolerant species such as Sesuvium portulacastrum (Aizoaceae), Salicornia fruticosa (Chenopodiaceae), and Batis maritima (Bataceae) are frequently found as landward fringe elements and also in disturbed mangrove interiors located in drier life zones, mainly from the Province of Manabi in western Ecuador to northwestern Peru, and in the Galapagos islands. Most terrestrial herbs inhabiting the interior of mangrove ecosystems have a widespread pattern of distribution, and only Crenea patentinervis (Lythraceae) is a typical element semi-restricted to the mangroves located in the rainforest and wet forest life zones.

The most prevalent dispersal mode of fruits and/or seeds in this habit seems to be via birds. At

present, absence of information does not allow us to provide a reliable percentage of dispersal modes for some species of this habit.

Lianas and vines

Lianas are woody climbing plants, e.g., Phryganocydia phellosperma; vines are nonwoody

climbing plants, e.g., Chamissoa altissima. These climbers are represented by eight families, comprising 16 spp. All species of lianas and vines in our area are dicots. In this habit, Malpighiaceae is the most diverse family, containing 43.7 % of the species.

The most prevalent dispersal mode of fruits and/or seeds among mangrove liana and vine

species is via wind (10 spp., Table 5). The second major diaspore dispersal syndrome in this habit is via water and zoochoreal dispersal by birds and presumably bats (3 spp. each). Among mangrove lianas, it is worth mentioning that Phryganocydia phellosperma and Tetrapterys subaptera have a similar distribution pattern that is restricted to mangroves, and that both species are commonly characterized by the presence of seeds/fruits adapted for water dispersal. Such an uncommon feature in regards to their respective genera suggests independient adaptations to mangrove ecosystems following different evolutionary pathways.

Epiphytes

Epiphytes are plants that grow atop other plants, either for an entire life cycle (holoepiphytes,

e.g., mangrove orchids and bromeliads) or only a stage (hemiepiphytes, e.g., most of species of Philodendron, Araceae) (Font Quer, 1985). Epiphytic plants often present aerial roots, using the host plant only for support while obtaining nutrients from debris, microparticles carried out by wind and rain, and sometimes associated invertebrate fauna excrement.

MANGROVE PLANTS

In the mangroves of the Pacific coast of South America, epiphytes are the most diverse habit,

comprising at least 60.8 % of the total flora. Most of the diversity is concentrated in the Chocoan mangroves located in wet forest and rainforest life zones, from the Choco Department in Colombia to the REMACAM, in the Province of Esmeraldas, northwestern Ecuador. Epiphytic diversity drastically decreases and is even commonly absent in the southern mangroves located in drier life zones.

Among the total flora of all plant habits in our area, epiphytes represent the most diverse habit

among monocotyledons, dicotelydons, and pteridophytes, respectively. Twenty six plant seed families having as many as five or more epiphytic species have been recorded for epiphytes in the Neotropics (Gentry and Dodson, 1987), and this reseach indicates that half of those (13) are present in the mangroves of the South American Pacific coast.

In regards to the family diversity among epiphytes, the dicotelydons are comprised by a higher

number (10) than pteridophytes (5), and monocotyledons (4, Table 4). The pteridophytes are represented by Polypodiaceae, Aspleniaceae, Dryopteridaceae, Hymenophyllaceae, and Pteridaceae; the monocots by Orchidaceae, Bromeliaceae, Araceae, and Cyclanthaceae; and the dicots by Piperaceae, Ericaceae, Marcgraviaceae, Cactaceae, Clusiaceae, Gesneriaceae, Melastomataceae, Asteraceae, Rubiaceae, and Scrophulariaceae. These are the largest and most common families that are predominantly epiphytic or that comprise epiphytic species in wet forest and rainforest terra firme ecosystems on the Pacific side of northwestern South America, where there is usually more diverse epiphytic flora (e.g., Dodson and Gentry, 1978; Cornejo, pers. obs.).

On the subject of species composition, the results indicate that monocotyledons are the most

diverse group (88 spp.), followed by the dicotelydons (34 spp.) and ferns (13 spp., Table 4). These data at the family and species level match the previously reported common pattern for epiphytes in the Neotropics (Gentry and Dodson, 1987). Overall for plant habits, the most diverse families in our area are: Orchidaceae (38 spp.), Bromeliaceae (28 spp.), and Araceae (19 spp.).

These predominantly epiphytic monocot families have been also previously reported to be the

most diverse among epiphytes in the Neotropics (Gentry and Dodson, 1987). They are represented by epiphytic species only (with a few species that are tolerant to hyposaline soils occasionally observed as facultatively terrestrial, e.g., Philodendron sparreorum), comprising 62.9 % of the epiphytic flora and at least 38.6 % of the total floristic composition.

Among epiphytes, there are no taxa restricted to mangroves at the family or generic level. At the

specific level, so far only the recently discovered Platystele cornejoi, Scaphyglottis sp. nov., and Sobralia rhizophorae are regarded, as semi-restricted to mangroves. It is expected that further exploration will yield the discovery of new records and increase the percentage in this habit.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

The most prevalent dispersal mode of fruits and/or seeds among epiphytes is via wind (79 spp.,

Table 5), and the second is via birds and presumably bats (40 spp.). Exozoochory via sticky diaspores have been reported for some species of Peperomia (Piperaceae; Gentry and Dodson, 1987), but as I have yet no observations to confirm such a dispersal syndrome in our area, all the 12 spp. of Peperomia have been lumped into the unknown dispersal mode category. Remarkably none of the epiphytic species have been recorded as water dispersed. The common presence of pappus in the Asteraceae species is a wellknown adaptation for wind dispersal in the family; however, Tuberostylis rhizophorae (Asteraceae), a coastal species semi-restricted to mangroves, have seeds lacking pappus, which suggest adaptation for dispersal via birds or even by water. Field observations are needed for verification.

Stranglers

Stranglers are woody plants that initially grow upon a host, as an epiphyte does, but soon develop

elongate roots that grow several to many meters to the ground to obtain nutrients, while additional roots surround the host, strangling it to death. A strangler can become a free-standing tree after the host plant dies.

This habit is represented by the dicot genus Ficus (Moraceae), and comprises only 5 spp. Most

strangler species have a widespread distribution range in several life zones and ecosystems throughout the Neotropics, but Ficus jacobii subsp. mantana is known only in western Ecuador. In mangrove ecosystems, stranglers mostly occur as landward fringe species, predominantly inhabiting the hyposaline mangroves located in moist forest, wet forest and rainforest life zones. The seed dispersal method is via vertebrates, mainly by frugivorous bats feeding on figs, but also by stout-billed birds such as Psittacidae, and occasionally by monkeys.

Parasites (incl. hemiparasites)

Parasites (incl. hemiparasites) are those plants that grow upon another plant, obtaining nutrients

from the host. At first sight, parasites are hardly differentiable from epiphytes, yet the roots of parasites differ from the true epiphytes by displaying root adaptations for penetrating the host xylem, floem, or both, to connect the vascular system of the host, and obtain nutrients.

This habit is represented by two well-known dicot families of mistletoes in Northwestern

South America: Loranthaceae and Viscaceae, both comprise 7 spp. The mistletoes are rather infrequent elements in the mangrove interior. Most species of parasites and hemiparasites recorded in our area have a widespread distribution range. Only Psittacanthus divaricatus is restricted to the dry forests of Western Ecuador to northwestern Peru, but no parasitic species are known to be restricted to mangroves. All species in this group are dispersed via birds.

MANGROVE PLANTS

Plate B: Some mangrove samps in northwestern South America: A. Bahia Malaga, Colombia. B. Parque Nacional Sanquianga, Colombia.C. Majagual, Ecuador. D. San Pedro de Vice, Peru. Photos A, B courtesy of J. Cantera-Kintz; photo D courtesy of R. Linares-Palomino.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate C: Some mangrove mollusca, crustaceans and fish. A. Cockles of concha prieta (Anadara tuberculosa). B. Gastropods (Littorina sp.). C. Fiddler crab (Uca ecuadoriensis). D. Red crab (Ucides occidentalis). E. Western Blue shrimp (Litopenaeus stylirostris). F. Fish (mostly Ariidae).

MANGROVE PLANTS

Plate D: A. An afroecuadorian community settled along estuarine waters, El Viento, Reserva Ecologica Manglares CayapasMataje (REMACAM), northwestern Ecuador. B. Fishermen, Refugio de vida silvestre manglares El Morro, western Ecuador.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate E. Some flower visitors to major mangroves: A. The red mangrove flower mite (Hattena rhizophorae). B. Syrphid fly (Palpada vinetorum). C. Lady beetle (Paraneda sp., to the left) and honey bee (Apis mellifera, to the right). D. Syrphid fly (Palpada albifrons). E. Flesh fly (indet., Sarcophagidae). F. Ant (indet., the upper) and lady beetle (Olla roatanensis, the lower). A, collected on Rhizophora racemosa. B, observed on Laguncularia racemosa. C-F, observed on Avicennia germinans.

MANGROVE PLANTS

Table 4. Number of families and species per habit of vascular plant groups from mangrove ecosystems on the Pacific coast of South America.

VASCULAR PLANT GROUPS TOTAL

HABIT MONOCOTYLEDONS

DICOTYLEDONS

Nº. spp.

PTERIDOPHYTE

GYMNOSPERMS

Fam

spp.

Fam

spp.

Fam

spp.

Fam

spp.

Trees/shrubs

–

Lianas/vines

–

Herbs

–

Epiphytes

–

135

Parasites

–

Stranglers

–

TOTAL

109

222

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Figure 1. Plant diversity by groups and taxonomic rank in mangrove ecosystems on the Pacific coast of South America.

Figure 2. Percentage of plant habits in mangrove ecosystems on the Pacific coast of South America.

MANGROVE PLANTS

Table 5. Dispersal method of plant species per habit from mangrove ecosystems on the Pacific coast of South America. Wind

Birds/bats

Water

Unknown

Trees/shrubs

Lianas/vines

–

Herbs

Epiphytes

Parasites

–

Stranglers

–

TOTAL

Figure 3. Percentage of fruits and/or seeds dispersal method in mangrove ecosystems on the Pacific coast of South America.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

MAJOR, MINOR, FACULTATIVE MANGROVES, AND OTHER FLORISTIC ELEMENTS Excepting the well-known true or major mangroves (Rhizophora, Avicennia and Laguncularia), considerable debate has taken place concerning which species should be regarded as mangroves. The key for determining which species are mangroves, and for understanding their types and sucessional patterns landwards from the sea, lies in the physical parameters of the substrates (soil, water, salinity), that are in direct correlation to the morphological and physiological adaptations of the species inhabiting such ecosystems. In this work, the term mangrove is applied to tree species or species having the potential to become trees. They are herein categorized as major, minor, and facultative. The MAJOR MANGROVES (Table 6), are those which commonly grow from the outermost seaward fringe, from the very soft-saline-muddy-water saturated- and poorly aired loose soils. They have morphological and physiological adaptations, mainly in the radicular system, as prop roots (Rhizophora), pneumatophores (Avicennia) or subsuperficially net-like roots (Laguncularia), suited to non-consolidated substrates (plate A). The hypocotyls (in Rhizophora), or fruits and seeds (Avicennia, Laguncularia) are salt-tolerant and water-dispersed. In our area, the pioneer species recognized as primary major mangroves are Rhizophora mangle, R. racemosa, and R. x harrisonii, which grow from extremely soft, non-consolidated soils. The primary major mangroves are followed by the secondary major mangroves Avicennia germinans and Laguncularia racemosa, in no particular order. Both species grow from somewhat more consolidated but still loose, poorly aired, and periodically flooded soils, in salty to brackish environments. This definition of major mangroves and all of the cited species fit well into Chapman’s characterization (1975), which states: “strictly only the saline facies can be regarded as true mangroves”, and into Tomlinson’s statement (1986), who considered the most significant criterion for major mangrove elements to be: “complete fidelity to the mangrove environment; that is, they occur only in mangrove ecosystems and do not extend into terrestrial communities”. Nevertheless, all species herein cited as major mangroves occasionally also occur on the adjacent or nearby sandy beaches; in the latter case, the major mangrove species are often secondary floristic elements. Therefore, in regards to the ecological distribution pattern, all species considered as major mangroves are semi-restricted to mangrove ecosystems. The MINOR MANGROVES (Table 6), are distributed landwards, behind the outer seaward fringe, and are restricted to somewhat more compact but still softly muddy, water-saturated, periodically flooded, and poorly aired soils, usually in hypo-, but occasionally in hyper-saline environments. Their fruits and seeds are salt-tolerant and water-dispersed. The minor mangroves sometimes form pure communities at peripheral habitats, as do back-fringe mangrove species. In our area, these species are Pelliciera rhizophorae and Tabebuia palustris. In regards to the ecological distribution pattern, the minor mangroves are semi-restricted to mangrove ecosystems.

MANGROVE PLANTS

The FACULTATIVE MANGROVES (Table 6), is a new category here presented to highlight the noteworthy mangrove-like role that is played by some species. It is composed of riparian and/or fresh water swamp trees that inhabit upstream river sides and adjacent freshwater swamps, that can extend their populations downstream to the mangrove ecosystems, co-existing among the major and minor mangrove communities or even forming pure stands in the intertidal zones, in the interior of the true mangrove swamps. These species are morphologically and physiologically adapted to inhabit poorly aired, muddy soils that are periodically flooded and saturated with fresh to saline water. They always have water dispersed fruits/seeds. In our area, the facultative mangroves are: Mora oleifera, Pterocarpus officinalis, Conocarpus erectus, Talipariti tiliaceum var. pernambucence, Amphitecna latifolia, and Annona glabra. It is noteworthy that Mora oleifera, Pterocarpus officinalis, and Annona glabra often form pure stands in fresh water swamps, and commonly share the feature of peculiar buttresses (a variable characteristic only in A. glabra), which play an important ecological role on the muddy soil and/or under water in semi-aquatic environments. On the other hand, the riparian and salt tolerant Conocarpus erectus and Talipariti tiliaceum var. pernambucence that occur in mangroves and at times on sandy or even volcanic beaches do not develop buttresses and apparently have not been recorded as inhabiting fresh water swamps. Regarding the geographic distribution pattern, the facultative mangroves fit within the widespread associate species (see Other floristic elements). It is needed to highlight that the facultative mangroves are the main floristic link between the terra firme and mangrove ecosystems. This original statement is based on the noteworthy mangrove-like role that they play in the soil-water-plant interaction carried out between fresh water swamps, river banks, and mangrove ecosystems.

Table 6. Categorization of mangrove types for species on the Pacific coast of South America. Family Rhizophoraceae Rhizophoraceae Rhizophoraceae Acanthaceae Combretaceae Tetrameristaceae Bignoniaceae Fabaceae Fabaceae Combretaceae Malvaceae Bignoniaceae Annonaceae

Scientific name Rhizophora mangle Rhizophora racemosa Rhizophora x harrisonii Avicennia germinans Laguncularia racemosa Pelliciera rhizophorae Tabebuia palustris Mora oleifera Pterocarpus officinalis Conocarpus erectus Talipariti tiliaceum var. pernambucence Amphitecna latifolia Annona glabra

Mangrove type major major major major major minor minor facultative facultative facultative facultative facultative facultative

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

OTHER FLORISTIC ELEMENTS: The remaining and often conspicuous floristic elements,

mostly composed of herbs to (sub)shrubs, lianas and epiphytes, are grouped according to their fidelity to mangrove environments. These are classified into three groups: restricted, semi-restricted, and widespread. The restricted species (Table 7), are those that occur in mangrove ecosystems only, forming healthy communities in brackish water saturated soils. The species comprised in this group are: (sub) shrubs, Pavonia rhizophorae, P. kearneyi; and lianas, Phryganocydia phellosperma, and Tetrapterys subaptera. These floristic elements are regarded as truly mangrove swamp species and not merely an associate flora. As in the case of lianas, restricted as they are to the Chocoan mangroves and having water dispersed corky seeds or wingless fruits, which are atypical for the predominantly wind dispersed taxonomical groups that they belong, Phryganocydia phellosperma, and Tetrapterys subaptera may well be typically called mangrove-lianas. The semi-restricted species (Table 7), are those which most commonly inhabit the back fringe or the interior of mangrove ecosystems, and extend their populations to the adjacent riparian or coastal terra firme forests, or vice-versa. They may also occur as transitional floristic elements. The species comprising this group are: herbs/(sub)shrubs, Acrostichum aureum (Pteridaceae), Crenea patentinervis (Lythraceae); lianas, Hiraea brachyptera (Malpighiaceae); shrub/trees, Hirtella carbonaria (Chrysobalanaceae), Muellera chocoensis (Fabaceae), Tuberostylis axillaris (Asteraceae); and epiphytics, Tuberostylis rhizophorae (Asteraceae). Epiphytics known by the type or as most two collections as Platystele cornejoi, Scaphyglottis sp. nov., and Sobralia rhizophorae are included here based on the fact that the remaining epiphytic plants in mangrove swamps of northwestern South America have a broader pattern of distribution due to the wind-dispersal syndrome. With the exception of the geographically widespread Acrostichum aureum, the species of this group occur only in the Chocoan mangroves and are absent from the EquatorialPacific mangroves. It is interesting to note that some semi-restricted species of terrestrial habits known to inhabit brackish waters do not have morphological adaptations (e.g., Crenea patentinervis). The widespread species are mangrove associates. These occur from the terra firme forests or fresh water wetlands inland to the interior of mangrove ecosystems, usually having a broad latitudinal and often wide altitudinal range of distribution. This group is comprised of the remaining species that do not belong in the restricted and semi-restricted groups. The predominant habit of these species is epiphytic, and their main substrate/host in the mangrove ecosystems are trees, e.g.: Guzmania monostachia, Tillandsia usneoides, Vittaria lineata (Pteridaceae), etc... This group also comprises terrestrial herbs such as Acrostichum danaeifolium (Pteridaceae); treelets, such as Chrysobalanus icaco (Chrysobalanaceae); lianas, such as Stigmaphyllon ellipticum (Malpighiaceae); and infrequently, parasites such as Phthirusa stelis (Loranthaceae).

MANGROVE PLANTS

Table 7. Other floristic elements restricted and semi-restricted to the mangroves on the Pacific coast of South America. Family

Scientific name

Bignoniaceae Malpighiaceae Malvaceae Malvaceae Acrostichaceae Asteraceae Asteraceae Chrysobalanaceae Fabaceae Lythraceae Malpighiaceae Orchidaceae Orchidaceae Orchidaceae

Phryganocydia phellosperma Tetrapterys subaptera Pavonia rhizophorae Pavonia kearneyi Acrostichum aureum Tuberostylis axillaris Tuberostylis rhizophorae Hirtella carbonaria Muellera chocoensis Crenea patentinervis Hiraea brachyptera Platystele cornejoi Scaphyglottis sp. nov. Sobralia rhizophorae

Distribution Restricted Restricted Restricted Restricted Semi-restricted Semi-restricted Semi-restricted Semi-restricted Semi-restricted Semi-restricted Semi-restricted Semi-restricted? Semi-restricted? Semi-restricted?

THE CHOCOAN AND EQUATORIAL-PACIFIC MANGROVE FORESTS Based on a particular composition, structure and different patterns of diversity that is related to different climates (life zones), and regardless of the widespread major mangroves, the present results suggest that two main floristic groups are present subdividing the mangrove forests on the Pacific coast of South America. Those are proposed here as: i) The Chocoan mangroves, and ii) the Equatorial-Pacific mangroves. The Chocoan mangroves are geographically centered at the Pacific coast of Colombia, discontinuously extending towards northwest along the adjacent coast of southwestern Mesoamerica to the Peninsula de Osa in southwestern Costa Rica, reaching to the south to the estuary of Cojimies (approximately to 0° 06´ N), in the north of Manabi Province, northwestern Ecuador. Those mangroves are distributed in the rainforests, wet forest and to the moist forest life zones, and are characterized by the presence of trees as: Pelliciera rhizophorae, Tabebuia palustris, Mora oleifera, Pterocarpus officinalis, Amphitecna latifolia, Hirtella carbonaria and Muellera chocoensis; (sub)shrubs as: Crenea patentinervis, Pavonia kearneyi, and P. rhizophorae; lianas as: Phryganocydia phellosperma, Tetrapteris subaptera and Hiraea brachyptera; and epiphytes as: Platystele cornejoi, Scaphyglottis sp. nov., and Sobralia rhizophorae; 91 % of the reported species (=202 spp.) are estimated to occur in the Chocoan mangroves. The Equatorial-Pacific mangroves, located southwards from the Chocoan mangroves, that is from near latitude 0º, nearby the town of Pedernales in the Manabi Province, western Ecuador, and southwards reaching the coast of northwestern Peru in the isolated Manglares San Pedro de Vice in Piura (5° 34’ S); and in the Galapagos islands. Those mangroves are distributed at the lower precipitation life

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

zones, in the dry and very dry forests, thorn woodland and desert scrub. The flora of Equatorial-Pacific mangroves is a depauperate version of that of the Chocoan mangroves, that is, containing the common major mangroves but characterized by the total absence of the previously cited Chocoan mangrove species. Interesting taxa as a variety of white mangrove: Laguncularia racemosa var. glabriflora; herbs as: Sesuvium portulacastrum, Batis maritima and Salicornia fruticosa; and lianas as: Cryptocarpus pyriformis, are present in this area; 20.3 % of the reported species (=45 spp.) are estimated to occur in the Equatorial-Pacific mangroves.

CONCLUSIONS AND RECOMMENDATIONS Despite multiple projects in mangrove conservation and management, it has been a surprise to find out that of the 222 species herein recorded as occurring in the mangrove interior on the Pacific coast of South America, only 14 species, which fortunately comprise the most common woody elements in our area, have been previously classified according to the IUCN criteria (2001). A categorization following the IUCN criteria (version 3.1, 2001) is here provided for all 222 species (Table 9, p. 251). From the floristic perspective, the present results indicate that specific and particular strategies must be addressed for the proteccion and management of the Chocoan and Equatorial-Pacific mangrove forests, respectively. The herein recorded as restricted and semi-restricted species that mostly occur in the Chocoan mangrove forests definitively have priority in terms of conservation due to their narrow range of distribution, with exception of Acrostichum aureum. As the facultative mangrove species constitute the main floristic link between the terra firme and mangrove ecosystems, the ecosystems and areas adjacent or connected to mangroves where the populations or most of the populations of facultative mangrove species occur must be also included for mangrove conservation and management plans. Study of the populations involving phenology, pollination, viability of seeds, growth rate, heavy metal concentrations and molecular studies are recommended for the white mangrove Laguncularia racemosa var. glabriflora, that occur in the Equatorial-Pacific mangroves. The preservation of structurally well-developed Rhizophora trees having roots at least 6 m in diameter is highly valuable for a long term sustainability of healthy mature populations of the Red crab Ucides occidentalis of commercial sizes (Cornejo, 2005). Including this parameter in the management and conservation projects of mangroves is highly recommended. Finally, as one of the greatest mangrove ďŹ sh species richness in the whole tropical eastern Pacific occurred at the central and southern coast of Colombia, that is related to the most extensive mangrove areas in the region (Castellanos-Galindo et al., 2012), the conservation and restoration of the Chocoan mangrove forests is a priority for a long-term sustainability of that valuable resource.

CHAPTER III.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

by Xavier Cornejo

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

FORMAT OF THE FLORA Vascular plants are arranged in the main groups of Pteridophytes, Gymnosperms, and Seed Plants (Monocotyledons and Dicotyledons). Families and species are alphabetically arranged in each group. For Pteridophytes, the classification for extant ferns (Smith et al., 2006) has been followed. For the remaining family names, the Angiosperm Phylogeny Website (APG) system (Stevens, 2001 onwards) has been followed. Author abbreviations for species adhere to Brummitt and Powell (1992). The botanical terminology used is Font Quer (1985), Jackson (1991), and Stearn (1995). The treated species are cited with basyonyms and/or synonyms, descriptions, distribution, ecology, IUCN Red list category, and, when available, vernacular names, biological interactions, phenology, etymology, uses, notes and dicussions. The description of species is mainly based on dry herbarium specimens. As the reproductive organs of plants usually have the tendency to shrink once dry, the measurements provided in the descriptions could be of slightly lesser length than fresh measurements would be. Several descriptions have been borrowed from those previously published in taxonomic revisions or monographs, such as Flora of Ecuador and Flora Neotropica, although the descriptions have often been improved by the authorâ&#x20AC;&#x2122;s field observations. Photos of 141 species are provided on plates 1 to 42. Line drawing of leaves arrangement (phyllotaxy), types and outlines of leaves are provided on plate 43. Species data such as distribution and phenology is mainly based on information provided on the label of herbarium specimens. For each species, the occurrence in mangroves and the general range of distribution is given. The Departments (Colombia and Peru) and Provinces (Ecuador) in which the species occur, were collected (e.g., Killip 5222 [NY]) or seen in the field (S), or if expected to occur (E), are provided; they are abbreviated after the respective country, as follows:

COLOMBIA. Choco: CH. Valle del Cauca: V. Cauca: C. NariĂąo: N. ECUADOR. Esmeraldas: E.

Manabi: M. Guayas: G. Santa Elena: SE. El Oro: O. Galapagos: GAL. PERU. Tumbes: T. Piura: P. To strictly reflect the species occurrence in mangroves of the cited departments or provinces, it has not been cited when the species occur in ecosystems other than mangroves. The presence of the species is supported by one herbarium collection per each department or province, if studied. For each herbarium collection reported, the collector, the number of collection and herbarium acronym are cited: e.g., Killip 5222 (NY). The herbarium acronyms follow Holmgren et al. (1990). The herbarium collections are reported to support species occurrence in mangroves. However, it is neccesary to remark that the herbarium records do not constitute the full distributional data. In some cases, the apparent absence of records in the Departments (Colombia and Peru) and Provinces (Ecuador) does not mean that the species does not occur there. Only species collected or seen in the field are numbered throughout the text.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

Key to the families 1. Epiphytes, stranglers or parasites 2. Plants bearing spores on the abaxial surface of lamina (ferns) or palm-like with conspicuous strobiliform cones (gymnosperms); epiphytic herbs only 3. Typical epiphytic herbs, bearing spores on the abaxial surface of lamina 4. Laminas monomorphic, all of similar sizes; sori rounded or linear 5. Sori arranged on elongate submarginal grooves along lamina; lamina ca. 2 mm wide Pteridaceae 5. Sori rounded or linear, if linear then arranged in parallel rows, oblique to midrib; lamina 1â&#x20AC;&#x201C;50 cm wide 6. Sori rounded 7. Leaves to 100 cm long, loosely covering the substrate; sori arranged along the lamina or at the upper third Polypodiaceae 7. Leaves to 14 cm long, densely covering the substrate or the leaves appressed; sori marginal, arranged near or at apex of pinnae lobes, otherwise stalked Hymenophyllaceae 6. Sori shortly linear Aspleniaceae 4. Laminas dimorphic; sori covering the whole abaxial surface of lamina 8. Laminas with margins Âą entire, the fertile laminas distinctively narrower Dryopteridaceae 8. Laminas with margins conspicuously dentate or serrate, all laminas of similar sizes Polypodiaceae 3. Plants palm-like with strobiliform cones, never bearing spores on the abaxial surface of lamina Zamiaceae 2. Plants bearing flowers and/or fruits (angiosperms); epiphytic herbs, lianas and shrubs, stranglers and parasites 9. Epiphytic herbs 10. Leaves absent Cactaceae 10. Leaves present 11. Leaves sessile 12. Leaves rosulate, forming a cup-like; stems absent or nearly so; flowers with all petals monomorphic; fruit berry or capsular, if capsular then with plumose or caudate winged seeds Bromeliaceae 12. Leaves basal, apical or alternate; stems cane-like or transformed in pseudobulbs (absent only in Ornitocephalus); flowers with one petal transformed (lip); fruit capsular with minute dustlike seeds Orchidaceae 11. Leaves petiolate 13. Lamina bifid, all nerves longitudinally arranged Cyclanthaceae

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

13. Lamina entire, the secondary nerves patent to ascendant, otherwise inconspicuous 14. Leaves alternate, spiral or subwhorled; the flowers densely packet in cylindric or oblong spadices or linear-oblong spikes 15. Stems usually bearing cataphylls or fibrous remains of cataphylls; spadices cylindric or oblong, at least 5 mm diam., subtended or surrounded by a spathe Araceae 15. Stems without cataphylls; spikes linear-oblong, less than 4 mm diam., the spathes absent Piperaceae 14. Leaves opposite or decussate; the flowers solitary or in heads 16. Inflorescences axillary, the flowers solitary, 2â&#x20AC;&#x201C;5 cm long; fruit a berry or fleshy capsule Gesneriaceae 16. Inflorescences terminal, the flowers in heads, ca. 3.5 mm long; fruit an achene Asteraceae 9. Epiphytic vines, lianas and shrubs, stranglers and parasites 17. Leaves absent; stems flattened or trigonous Cactaceae 17. Leaves present; stems more or less terete or tetragonous 18. Epiphytic vines or lianas 19. Leaves alternate or spiral 20. Pedicellary or rachial nectaries present; corolla of free petals Marcgraviaceae 20. Pedicellary or rachial nectaries absent; corolla of fused petals Ericaceae 19. Leaves opposite or decussate 21. Leaves with glandular fields near base of blade beneath Scrophulariaceae 21. Leaves without glandular fields near base of blade beneath 22. Interpetiolar stipules absent; ovary superior Gesneriaceae 22. Interpetiolar stipules present; ovary inferior Rubiaceae 18. Epiphytic shrubs, stranglers and parasites 23. Latex present 24. Leaves opposite or decussate; fruits capsular, dehiscent at maturity Clusiaceae 24. Leaves alternate or spiral; fruits figs, indehiscent at maturity Moraceae 23. Latex absent 25. Leaves alternate or spiral 26. Pedicellary or rachial nectaries present; corolla of free petals Marcgraviaceae 26. Pedicellary or rachial nectaries absent; corolla of fused petals Ericaceae 25. Leaves opposite 27. Leaf blades with conspicuously parallel tertiary nerves Melastomataceae 27. Leaf blades with tertiary nerves absent or if present, then reticulate, inconspicuous 28. Epiphytes; corolla gamopetalous, conspicuous Gesneriaceae 28. Parasites; corolla dialipetalous, often inconspicuous 29. Flowers with calyculus Loranthaceae 29. Flowers without calyculus Viscaceae

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

1. Terrestrials or aquatics 30. Plants bearing spores on the abaxial surface of lamina (ferns) or palm-like with conspicuous strobiliform cones (gymnosperms); herbs only 31. Typical herbs, bearing spores on the abaxial surface of lamina 32. Plants to 0.8 m tall; pinnae serrulate or crenate-dentate at margin; sori linear arranged on both sides of midvein or (sub)circular and submarginal on pinnae beneath 33. Pinnae serrulate at margin, the lateral veins parallel, conspicuous; sori linear, longitudinally arranged along both sides of midvein on pinnae Blechnaceae 33. Pinnae crenate-dentate at margin; the lateral veins inconspicuous; sori (sub)circular, submarginally arranged on pinnae Lomariopsidaceae 32. Plants taller than 0.8 m tall; pinnae entire at margin; sporangia covering the whole pinna beneath Pteridaceae 31. Plants palm-like with strobiliform cones Zamiaceae 30. Plants bearing flowers and/or fruits; herbs, lianas, shrubs or trees (angiosperms) 34. Trees and/or shrubs 35. Unbranched, leaves longer than 1 m Arecaceae 35. Branched, leaves shorter than 50 cm 36. Arching prop roots present; developed individuals with adventitious roots from branches, these pendulous, rope-like; leaves abundantly black-dotted beneath; fruits producing hypocotyls linearobclavate, 20â&#x20AC;&#x201C;50 cm long Rhizophoraceae 36. Arching prop roots absent; adventitious roots absent; leaves without black dots beneath; fruits do not producing hypocotyls linear-obclavate, 20â&#x20AC;&#x201C;50 cm long 37. Petioles with a distal pair of lateral glands; blades with scattered pits or domatia beneath Combretaceae 37. Petioles without a distal pair of lateral glands; blades without pits or domatia beneath (blades with glands only in Malpighiaceae) 38. Leaves opposite or decussate 39. Secondary superficial roots with many pneumatophores, these narrow, pencil-thick, erect Acanthaceae 39. Secondary superficial roots without pneumatophores 40. Leaves 3-foliolate Bignoniaceae 40. Leaves simple 41. Leaves plinerved 42. Tertiary nerves parallel; fruit berry Melastomataceae 42. Tertiary nerves inconspicuous; fruit achene Asteraceae 41. Leaves pinnatinerved 43. Interpetiolar stipules present, soon deciduous; ovary inferior; fruits linearoblong, dehiscent at maturity; seeds winged Rubiaceae

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

43. Interpetiolar stipules absent; ovary superior; fruits globose to elliptic, indehiscent at maturity; seeds not winged 44. Leaf blades with “T” shape trichomes and scattered glands beneath; corolla yellow; sepals bearing 1(–2) large abaxial glands Malpighiaceae 44. Leaf blades glabrous or with simple trichomes and eglandular beneath; corolla greenish to white; sepals abaxially eglandular 45. Corolla gamopetalous, zygomorphic, 3.8–6.2 cm long; stamens 4; fruits large, 6–9 cm wide Bignoniaceae 45. Corolla dialipetalous, actinomorphic, less than 2 cm long; stamens 8 to ca. 24; fruits small, less than 2 cm wide 46. (Sub)shrubs; stem tetragonous; inflorescences of solitary flowers or cymes; petals entire; stamens 8 Lythraceae 46. Trees; stem more or less terete; inflorescences fasciculate; petals fimbriate; stamens numerous Rhizophoraceae 38. Leaves alternate or spiral 47. Leaves paripinnate or imparipinnate Fabaceae 47. Leaves simple or unifoliolate 48. Buttresses narrow, irregularly conical, to 1.5 m tall; leaves asymmetric, sessile; flowers with filaments of stamens arranged for most of their length closely appressed in and along longitudinal grooves of narrowly conical ovaries; fruits napiform, with a long acuminaterostrate at apex Tetrameristaceae 48. Buttresses absent; leaves symmetric, usually petiolate; flowers with filaments of stamens never appressed on ovaries; fruits of variously shape, but never napiform with an acuminate-rostrate apex 49. Terminal branches, pedicels and blades beneath covered by stellate or lepidote trichomes; flowers with stamens arranged in an elongate central column (monadelphous); fruits dehiscent at maturity Malvaceae 49. Stellate or lepidote trichomes absent; flowers with stamens not arranged in an elongate central column; fruits indehiscent at maturity 50. Leaves unifoliolate; fruits strongly complanate in cross section Fabaceae 50. Leaves simple; fruits globose to more or less elliptic in cross section 51. Latex white to cream present; fruits figs; stranglers when juveniles Moraceae 51. Latex absent; fruits of variously types, but never figs; never stranglers 52. Leaf blades beneath, calyx, corolla, and fruits glandular-punctate Myrsinaceae 52. Leaf blades beneath, calyx, corolla, and fruits not glandular-punctate 53. Leaves spiral, grouped at the end of branches; flowers gamopetalous Bignoniaceae

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

53. Leaves alternate, distichous, arranged along terminal branches, flowers dialipetalous 54. Corolla yellow, fleshy, arranged in two whorls of 3 petals each; stamens inserted; fruits syncarpic, subglobose, 5–12 cm long, light-green to yellowish at maturity, many-seeded Annonaceae 54. Corolla white, thin, arranged in one whorl of 5 petals; stamens exserted; fruits fleshy drupes, ovate to obovate, 2–3 cm long, red to dark-purple or black at maturity, one-seeded Chrysobalanaceae 34. Herbs, vines and lianas 55. Herbs 56. Plants floating or the whole plants permanently submersed 57. Plants floating on water surface or stranded in the mud; petioles swollen, spongy and aerenchymatous within; leaves deltoid, suborbicular, reniform, ovate or elliptic; flowers showy, the petals and sepals present Pontederiaceae 57. Plants permanently submersed in stagnant waters; petioles not swollen, not spongy and not aerenchymatous within; leaves linear; flowers inconspicuous, the petals and sepals absent Potamogetonaceae 56. Plants not floating, the whole plants never permanently submersed 58. Leaves rudimentary, reduced to minute scales or subulate (to 10 x 2 mm), otherwise absent 59. Plants solitary; stems branched Amaranthaceae 59. Plants caespitose; culms unbranched Cyperaceae 58. Leaves developed, flattened or succulent, always present 60. Leaves succulent 61. Branches prostrate, trailing and often rooting at nodes; petioles clasping at base; inflorescences solitary flowers Aizoaceae 61. Branches (sub)erect, not rooting at nodes; petioles absent, not clasping at base; inflorescences strobiliform Bataceae 60. Leaves flattened, thin 62. Leaves (sub)whorled or rosulate, arranged at base or at base and apex of stem (if stem is present) 63. Leaves less than 2 cm wide; inflorescences in spikelets, the flowers inconspicuous, the petals absent Cyperaceae 63. Leaves 2–5(–8) cm wide; inflorescences in umbels, the flowers showy, the petals present Amaryllidaceae 62. Leaves loosely alternate, spiral, opposite or decussate, arranged along stem 64. Leaves alternate or spiral, sessile; inflorescences in spikelets, the petals absent 65. Culms terete Poaceae 65. Culms 3- or 5-angulate Cyperaceae

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

64. Leaves opposite or decussate, petiolate; inflorescences a panicle, solitary or usually in cymes, the petals present 66. Branches subterete; interpetiolar stipules present; inflorescences axillary heads; ovary inferior Rubiaceae 66. Branches 4-angled; interpetiolar stipules absent; inflorescences solitary flowers or usually cymes; ovary superior Lythraceae 55. Vines and lianas 67. Leaves alternate or spiral 68. Leaves unifoliolate; flowers ca. 1 cm long; fruits strongly flattened, 2–5 cm long Fabaceae 68. Leaves simple; flowers 2–4 mm long; fruits subglobose, ca. 3–4 mm long 69. Subwoody lianas; lateral veins 2–4 per side; stamens exserted Nyctaginaceae 69. Herbaceous vines; lateral veins 5–10 per side; stamens inserted Amaranthaceae 67. Leaves opposite or decussate 70. Leaves 2–3-foliolate; tendrils present Bignoniaceae 70. Leaves simple; tendrils absent 71. Interpetiolar stipules present; ovary inferior Rubiaceae 71. Interpetiolar stipules absent; ovary superior 72. Leaf blades plinerved; inflorescences in sessile heads; fruits achenes Asteraceae 72. Leaf blades pinnatinerved; inflorescences in pedunculate panicles; fruits capsular or samaroid 73. Latex (white) present; corolla gamopetalous, infundibuliform Apocynaceae 73. Latex absent; corolla dialipetalous 74. Leaves with “T” shape trichomes beneath; petals clawed; sepals bearing 1(–2) large abaxial glands; infructescences samaroid, indehiscent Malpighiaceae 74. Leaves glabrous beneath; petals sessile; sepals abaxially eglandulate; infructescences schizocarps, dehiscent Hippocrateaceae

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

Ferns FILICOPSIDA (FERNS) Herbs or vines, often epiphytic, rarely aquatic; usually rhizomatous. Leaves variously divided or entire, uncurling apically in openning. Flowers wanting; instead spore producing sporangia spread on the leafunderside or on distinct fertile leaves. ASPLENIACEAE Terrestrial or epiphytic herbs, perennial, less than 1 m tall. Rhizomes erect, decumbent or creeping, bearing clathrate scales, these often iridiscent with prominent, dark brown or black lateral walls and hyaline exterior walls, glabrous. Fronds circinate in bud, monomorphic, fasciculate to spaced on a usually short-creeping rhizome; stipe not articulate; lamina unlobed to more commonly pinnate to decompound; rachis often ridged, slighltly grooved adaxially; pinnae often inequilateral, excavate on proximal side, often truncate, angulate, or auriculate on distal side. Sori commonly borne only along acroscopic side of veins on abaxial surface of lamina, oblong to linear. Sporangia glabrous, commonly long-stalked, the stalk usually 1-rowed; paraphyses lacking; indusia well developed, infrequently absent. Spores ellipsoid, monolete, surface often sharply winged or cristate. Gametophytes epigeal, chlorophyllous, ± cordate. Key to the species 1. Blades simple, 20–70 cm long 1. Blades pinnate-pinnatisect, 13–20 cm long

Asplenium serratum Asplenium sp.

1. Asplenium serratum L., Sp. Pl. 1079 (1753). Syn.: Asplenium subsessile Cav. Asplenium nidus Raddi Asplenium crenulatum Presl Asplenium schomburgkianum Kl. Holoepiphytic herbs, caespitose. Rhizome stout to massive, thickly provided with browntomentose roots, the clathrate scales linear to linear-lanceolate, 4–12 mm long, attenuate at apex, iridiscent, light gray-brown to blackish. Leaves erect; stipe 0–4 cm long; blade simple and unlobed, chartaceous, narrowly-oblanceolate to elliptic, 20–70 x 4–15 cm, long attenuate at base, acute to abruptly caudate at apex, entire to crenulate or blunty serrate at margins, the surface and margins glabrous, but commonly with a few dark-brown to blackish clathrate

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

scales scattered along midrib, especially abaxially. Veins simple to once forked, crowded, rather obscure. Sori crowded, greatly elongate, extending from midrib more than halfway to margin. Indusium narrow, elongate, delicate, pale green, scarious, extending the length of sorus. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 2726 (AAU, GUAY, QCA). Distribution: Southeastern United States (Florida), West Indies, Mexico to northern Argentina and Paraguay. Ecology: In moist forests, wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that refers to the serrate blade margins. 2. Asplenium sp. Holoepiphytic herbs, caespitose. Rhizome stout, the clathrate scales lanceolate to ovate, acute or acuminate at apex, dark-brown. Leaves erect; stipe (3–)5–10 cm long, dark-brown to blackish; blade pinnate-pinnatisect, lanceolate, 13–20 x 3–6 cm, gradually terminating in a pinnatifid to serrate apex; rachis with scattered scales; pinnae several per side, 1–3 cm long, short-stalked, patent or ascending, oblong or elliptic and obtuse, scarcely auriculate at base, the veins forked. Indusium linear. ECUADOR. E: (S). Ecology: In wet forests. IUCN Red list category: Deficient Data (DD). BLECHNACEAE Terrestrial herbs, perennial, erect or scandent. Rhizomes creeping, suberect or erect, occasionally climbing trees, sometimes bearing stolons, scaly, dictyostelic. Fronds circinate in bud, monomorphic or dimorphic, the fertile ones often longer, more erect, and with much narrower segments, the young ones often reddish due to presence of anthocyanins; stipes non articulate; lamina commonly pinnatifid to 1-pinnate pinnatifid; rachis adaxially grooved. Sori linear to oblong, borne on an outer arc of an areole or on a vein that closely parallels midrib of ultimate divisions, or on submarginal veins that join vein tips; paraphysis lacking; indusia with same shape as sorus or absent, opening toward veins or axis. Sporangia oblong; annulus vertical; stalk 2- or 3-rowed; spores 64 per sporangium, commonly beige or light-brown, ellipsoid to subglobose, monolete, the surface variously ornamented. Gametophytes epigeal, chlorophyllous, cordate to elongate, the antheridia 3-celled.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

3. Blechnum serrulatum Rich., Actes Soc. Hist. Nat. Paris 1: 114 (1792). Plate 1: A. Syn.: Blechnopsis serrulata (Rich.) C. Presl Spicanta serrulata (Rich.) Kuntze Terrestrial herbs, 0.7 m tall. Rizomes underground, elongate, ca. 7 mm diam., with tomentulous roots, the hairs light-brown to yellowish, 0.5–1.5 mm long, the apical portion of rhizome densely covered by scales, the scales narrowly-lanceolate, 2–4 mm long, darkbrown to blackish, lustrous. Leaves suberect; stipe ca. 13–18 cm long, glabrous; blade imparipinnate, 16–50 cm long; pinnae ca. 25–31, subopposite, sessile, oblong, 4.5–7 x 1–1.5 cm, auriculate at base, acute at apex, serrulate at margin, with many forked veins from base and parallel to each side of midvein, above lustrous and glabrous, beneath with scales ca. 1 mm long on midvein. Sori linear, on both sides of midvein on pinnae. COLOMBIA. N: Mora 4185 (COL). ECUADOR. E: Cornejo et al. 7230 (AAU, GUAY, MO, QCA). Distribution: Southeastern United States (Florida), and southern Mexico to northeastern Argentina. Ecology: Blechnum serrulatum grows in fresh water swampy areas. Occasionally it is a backfringe species in the hyposaline mangroves, growing in dry, moist and wet forest life zones. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that refers to the serrulate blade margins. DRYOPTERIDACEAE Terrestrial or epiphytic plants, perennial. Rhizomes creeping to erect, rarely arborescent, sometimes climbing, dictyostelic, bearing scales. Fronds circinate in bud; axes and lamina usually with glands, hairs and/or scales abaxially. Stipes at base usually with persistent scales, lacking spines. Lamina undivided to commonly 1–5-pinnate. Sori borne abaxially on veins, or sporangia achrostichoid and covering abaxial surface of lamina, if sporangia in sori then sori variously shaped (round, oblong or elongate); paraphyses generally absent, indusia variously linear, sickle-shaped, reniform, or peltate. Sporangia with 2- or 3-rowed stalk; annulus vertical, interrupted by stalk; spores all alike, usually lacking chlorophyll, oblong or reniform, monolete, ornamented, 64 per sporangium. Gametophytes epigeal, chlorophyllous, obcordate; archegonia and antheridia borne on abaxial surface, antheridia 3-celled. Key to the species 1. Sterile blades 30–40 x 4–8 cm, covered by dark-brown scales beneath; fertile blades 15–27 x 1.5–3 cm, the stipes 15–20 cm long Elaphoglossum latifolium 1. Sterile blades 12–20 x 1.5–2.5 cm, glabrous beneath; fertile blades 7–12 x 0.6–1 cm, the stipes 5–15 cm long Elaphoglossum productum

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

4. Elaphoglossum latifolium (Sw.) J. Sm., London J. Bot. 1: 197 (1842). Plate 1: B. Bas.: Acrostichum latifolium Sw. Syn.: Acrostichum sartorii Liebm.

Elaphoglossum sartorii (Liebm.) Mickel Acrostichum scapellum Kunze ex Fée Elaphoglossum scapellum (Kunze ex Fée) T. Moore Holoepiphytic herbs. Rhizome with villose roots, densely covered by scales linear-lanceolate, 3–4 mm long, dark-brown, the hairs light-brown, ca. 1 mm long. Leaves erect, simple, regular; blades narrowly-elliptic, dimorphic: The sterile ones 30–40 x 4–8 cm, narrowly-attenuate at base, acuminate at apex, beneath covered by scales more or less fusiform, dark-brown on both sides of midvein; stipe 7–10 cm long. The fertile smaller, 15–27 x 1.5–3 cm, narrowlyattenuate at base, acute at apex, entirely covered by sori beneath; stipe longer, 15–20 cm long, with scales scattered clathrate, linear-lanceolate, 3–4 mm long, the margin pilose.

COLOMBIA. N: (S). ECUADOR. E: Cornejo & Bonifaz 6942 (AAU, GUAY, QCA). Distribution: Mexico to Bolivia. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet is formed by the Latin words lata (wide) and folia (leaves) that refers to the wide leaves. 5. Elaphoglossum productum Rosenst., Repert. Spec. Nov. Regni Veg. 6: 315 (1909). Plate 1: C. Holoepiphytic herbs. Rhizome densely covered by narrowly-lanceolate scales, 3–4 mm long, brown. Leaves erect, simple, regular; blades very narrowly-elliptic or narrowlyoblong-elliptic, dimorphic: The sterile ones 12–20 x 1.5–2.5 cm, narrowly-attenuate at base, narrowly-acute to acuminate at apex, with scattered stellate-lepidote scales beneath; stipe 4–8 cm long. The fertile narrowly-oblong-ellliptic, 7–12 x 0.6–1 cm, narrowly-attenuate at base, acute at apex, entirely covered by sori beneath; stipe 5–15 cm long, mostly glabrous. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8197 (GUAY, NY). Distribution: Costa Rica to Bolivia. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means produced, extended, elongated, or brought forward. However, it is not clear what it refers to in this species.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

HYMENOPHYLLACEAE (Filmy fern Family) Plants epiphytic or terrestrial. Rhizomes protostelic, pubescent (not scaly), usually creeping and bearing distichous leaves, less commonly erect and radially symmetrical. Leaves generally small, 1–40 cm long, monomorphic or (less commonly) dimorphic. Petioles with a single vascular bundle. Laminae almost always one cell thick between the veins, without stomata. Veins free. Sori marginal, borne at the vein tips. Indusia formed by the green laminar tissue, either urceolate, tubular or bivalvate. Sporangia borne on all sides of the receptacle, short-stalked (the stalk about 6 rows), with annulus oblique, not interrupted at the stalk. Spores green, trilete, tetrahedral-globose. Gametophytes filamentous or thallose or both, often gemmiferous; antheridia with 5 to many cells. Key to the species 1. Leaves flatty appressed to tree trunks; pinnae crenate-lobed to serrate at margins; sori stalked Trichomanes ankersii

1. Leaves erect to pendulous, not appressed to tree trunks; pinnae entire at margins; sori sessile 2. Midvein flexuose; involucre 2-valvate

Hymenophyllum polyanthos

2. Midvein straight; involucre not 2-valvate

Trichomanes crinitum

6. Hymenophyllum polyanthos (Sw.) Sw., J. Bot. 1800(2): 102 (1801). Plate 1: D. Bas.: Trichomanes polyanthos Sw. Holoepiphytic creeping short herbs, forming dense wet masses. Rhizome creeping, slender, branched, with numerous short, brown trichomes. Leaves suberect to subpendulous; stipe 1–2 cm long; blades simple, deeply and irregularly pinnatifid, lanceolate in outline, ca. 4–6 x 2–3.5 cm, the midvein flexuose, the ultimate lobes usually emarginate, the veins dichotomously branching several times in lobes, glabrous. Sori near apex of blade, the indusia 2-valvate. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8282 (GUAY, NY). Distribution: Southern Mexico to Bolivia, and in the West Indies. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC).

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Etymology: The epithet is formed by the Greek prefix poly (many, numerous) and the Greek word anthos (flower), which most likely refers to the sori that the blade bears on its underside, near the apex. The describing author probably had in mind that the sori are the reproductive parts and thus there is a vague analogy to flowers (R. Moran, pers. comm.). Note: The provided identification is provisional; a monograph of the genus Hymenophyllum is needed (R. Moran, pers. comm.). Trichomanes ankersii C. Parker ex Hook. & Grev., Icon. Filic. 2(11): t. 201 (1831). Hemiepiphytic creeping herbs, rhizome creeping, slender, blackish. Leaves flatly appressed to trunks, spreading; stipe not obvious; blades simple, deeply pinnatifid, lanceolate in outline, 5–13 x 3–8 cm, cordiform to cuneate at base, acute to acuminate at apex; veins and rachis appressed pilose; pinnae oblong to lanceolate, 8–18 pairs, 1.5–4 x 0.5–1 cm, the basal and medial pinnae longer than distal pinnae, all crenate-lobed to serrate at margins. Sori 1–11 per pinnae; involucre 3–4 mm long, free, stalked. COLOMBIA. CH: (E). V: (E). C: (E). Distribution: Nicaragua to Bolivia, and in the West Indies. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). 7. Trichomanes crinitum Sw., Prodr. 136 (1788). Syn.: Ragatelus crinitus (Sw.) C. Presl Hemiepiphytic creeping herbs, rhizome creeping, slender, ferrugineous. Leaves erect to pendulous, spreading; stipe 0.8–5 cm long, pilose; blades 2-pinnatifid to 1-pinnate-pinnatifid, lanceolate in outline, 3–14 x 1.2–2.5 cm, truncate at base, acute at apex; sparsely pilose; pinnae oblong-obovate, 11–25 pairs, 0.8–2 x 0.5–0.8 cm, the basal pinnae sometimes slightly reduced, entire at margins. Sori 1–3, at pinnae apex; involucre ca. 2.8 mm long. COLOMBIA. V: Gentry & Juncosa 40727 (MO). Distribution: Costa Rica to Peru, and in the West Indies. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet means having tufts of long, weak hairs, referring to the pubescence of this species.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

LOMARIOPSIDACEAE Plants terrestrial, epilithic, hemiepiphytic, or epiphytic. Leaves monomorphic or dimorphic, hairs generally absent. Petioles with 3 or more vascular bundles, the two adaxial ones largest. Laminae 1-pinnate, rarely simple; stomata polocytic, occasionally anomocytic. Pinnae articulate to the rachis. Rachises and costae grooved or terete. Veins free. Sori round and indusiate or acrostichoid. Sporangial stalk 3-seriate. Spores monolete. Gametophytes epigeal or on the base of tree trunks, green, either cordate or elongate and irregularly lobed.

A pantropical family comprising four genera, only one of which is present in our area.

8. Nephrolepis biserrata (Sw.) Schott, Gen. Fil. pl. 3 (1834). Plate 2: A. Bas.: Aspidium biserratum Sw. Terrestrial herbs, 0.7 m tall. Rizomes underground, elongate. Leaves suberect; stipe ca. 10–20 cm long, with pluriseriate whitish-hyaline hairs; blade imparipinnate, 40– 80 cm long; rachis with abundant pluriseriate whitish-hyaline hairs; pinnae ca. 25–60, alternate, subsessile, oblong-lanceolate, 5–8 x 1.5–2 cm, broadly-obtuse to truncate and asymmetric at base, narrowly-acute to acuminate at apex, crenate-dentate at margin, with inconspicuous lateral veins, above mostly glabrous, beneath abundantly pilose with pluriseriate whitish-hyaline hairs, 0.3–1 mm long, those mainly arranged along and near midvein. Sori (sub)circular, submarginally arranged in one row along pinnae. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Nazareno 8234 (GUAY, NY). Distribution: Southern Mexico to Bolivia, also in the West Indies, and Africa. Ecology: In wet forests. Use: It may be cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet means having leaflets with margins double serrate. POLYPODIACEAE Terrestrial or frequently epiphytic herbs, perennial. Rhizomes long- to short-creeping, bearing scales, usually dictyostelic. Fronds monomorphic to dimorphic; stipes usually articulate at base; rachis grooved or not adaxially. Sori round to oblong; sporangial stalk 2- or 3- rowed; indusium absent. Spores bilateral, monolete, usually hyaline or yellowish, surface most often smooth, verrucose, or tuberculate, occasionally spinose. Gametophytes epigeal, chlorophyllous, cordate.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Key to the species 1. Leaf blades entire 2. Rhizome short; leaf blades 35–60 x 4–9 cm; sori pluriseriate Niphidium crassifolium 2. Rhizome elongate, to several meters long; leaf blades to 14 x 2 cm; sori continuously covering, or rounded along blade beneath 3. Leaf blades with margins dentate or serrate; sori continuously covering most of the blade beneath Cochlidium serrulatum 3. Leaf blades with margins entire; sori rounded arranged along blade beneath Microgramma percussa 4. Leaves monomorphic, the blades 10–14 cm long, 1.3–2 cm wide 4. Leaves dimorphic, the blades 2–8 cm long, the fertile blade 2.5–4 mm wide Microgramma reptans 1. Leaf blades 1-pinnate 5. Blade ovate in outline, the lateral segments 3–7 pairs, 8–30 cm long; sori 1.5–2 mm diam., in 2 or 3 rows beside midrib Serpocaulon triseriale 5. Blade oblong-oblanceolate in outline, the lateral segments 35 to 60 pairs, 1–3 cm long; sori ca. 1 mm diam., in one row beside midrib Polypodium bombycinum

9. Cochlidium serrulatum (Sw.) L.E. Bishop, Amer. Fern J. 68(3): 80 (1978). Plate 2: B. Bas.: Syn.:

Acrostichum serrulatum Sw. Asplenium serrulatum (Sw.) Sw. Grammitis serrulata (Sw.) Sw. Gymnopteris serrulata (Sw.) Bernh. Xiphopteris serrulata (Sw.) Kaulf. Micropteris serrulata (Sw.) Desv. Polypodium serrulatum (Sw.) Mett. Micropteris orientalis Desv.

Holoepiphytic small herbs, caespitose. Rhizome creeping, densely covered by thin narrowly-lanceolate scales, 1.5–2 mm long, brown. Stipe 1–10 mm long, slender; blades simple, linear-oblong, 2–5 cm x 1–2.5 mm, attenuate at base, dentate at apex, the sterile fronds and at least the sterile portions of the fertile fronds distinctly serrate or acutely lobed at margins; fertile fronds covered by sori beneath. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8399 (NY). Distribution: Southern Mexico through Central America to Bolivia, the West Indies, and in the wetter parts of tropical Africa. Ecology: In wet forests and rainforests.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that refers to the characteristic serrulate leaf blade margins. 10. Microgramma percussa (Cav.) de la Sota, Physis (Buenos Aires), Secc. C, 44: 28 (1986). Plate 2: C. Bas.: Polypodium percussum Cav. Syn.: Pleopeltis percussa (Cav.) Hook. & Grev. Holoepiphytic climber herbs. Rhizome creeping, clothed with thin linear-lanceolate scales, 2–3 mm long. Stipe 2.5–4 cm long, slender, scaly; fertile blades simple, narrowly-lanceolate, 10–14 x 1.3–2 cm, attenuate at base, acute at apex, covered by lepidote scales on abaxial side. Sori uniseriate, rounded, prominent, ca. 2 mm diam. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Bonifaz 2723 (GUAY). M: Neill et al. 11422 (MO). Distribution: Southern Mexico through Central America to Bolivia. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). 11. Microgramma reptans (Cav.) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40(8): 230 (1975). Plate 2: D. Bas.: Acrostichum reptans Cav. Holoepiphytic climber herbs. Rhizome creeping, clothed with thin linear-lanceolate scales, 1–2 mm long, light-brown. Leaf blades dimorphic; sterile ones with stipe 0.3–0.7 cm long, scaly, the blades simple, lanceolate, ca. 2–7 cm x 8–13 mm, obtuse at base, acute at apex, covered by linearlanceolate scales on adaxial side; fertile ones with stipe 0.5–2 cm long, scaly, the blades simple, narrowly-oblong, 2–8 cm x 2.5–4 mm, attenuate at base, acute to acuminate at apex, covered by linear-lanceolate scales on adaxial side. Sori biseriate, rounded, prominent, ca. 1–1.5 mm diam. COLOMBIA. CH: (E). ECUADOR. E: (E). M: Neill et al. 11415 (MO) Distribution: Southern Mexico through Central America to Bolivia. Ecology: In moist forests, wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means creeping, referring to the habit of this species. 12. Niphidium crassifolium (L.) Lellinger, Amer. Fern. J. 62: 106 (1972 [1973]). Plate 3: A. Bas.: Syn.:

Polypodium crassifolium L. Anaxetum crassifolium (L.) Schott. Dipteris crassifolia (L.) J. Sm. Drinaria crassifolia (L.) J. Sm. Pessopteris crassifolia (L.) Underw. & Maxon Pleuridium crassifolium (L.) Fée

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Holoepiphytic herbs. Rizome short, creeping, woody, 5–15 mm thick, enveloped in a dense mass of rootlets, clothed at apex with scales, the scales imbricate, finely clathrate, lanceolate, 8–12 mm long, bicolours. Leaves few, close, stiffly erect to 60 cm long; stipe stout, 3–10 cm long; blade ligulate or narrowly-oblanceolate, 35–60 x 4–9 cm, attenuate at base, truncate to rounded at apex; primary veins oblique, parallel. Sori pluriseriate, rounded, prominent, 1.5–3 mm diam., mostly confined to distal half of blade. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Bonifaz 2673 (AAU, GUAY). Distribution: Southern Mexico through Central America to Bolivia. Ecology: In moist forests, wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Latin prefix crassi (thick) and the Latin word folia (leaves), referring to the thick blades of this species, a non-distinctive feature. 13. Pleopeltis bombycina (Maxon) A.R. Sm., Candollea 60(1): 281 (2005). Plate 3: B. Bas.: Polypodium bombycinum Maxon Holoepiphytic herbs. Rizome short creeping, with numerous branches, these 3–5 mm diam., densely and divaricately paleaceus, the scales lanceolate, 3–4 mm long, linear–attenuate at apex, castaneous, concolor. Leaves several, stiffly-erect, subpinnatisect, ± narrowly oblongoblanceolate in outline, 12–45 x 2–5 cm, very densely appressed-paleaceous throughout; stipe 0.3–2 cm long; segments 35 to 60 pairs, patent, oblong in the distal 2/3 of blade, mostly 1–3 cm x 2.5–5 mm, entire at margins, acute at apex. Sori ca. 10 pairs, rounded, ca. 1 mm diam., in one row beside midrib. COLOMBIA. V: (E). ECUADOR. G: (S). Distribution: Panama to Paraguay. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means silken, referring to the silky appearance of the blade’s underside. 14. Serpocaulon triseriale (Sw.) A.R. Sm., Taxon 55(4): 929 (2006). Plate 3: C. Bas.: Polypodium triseriale Sw. Syn.: Polypodium brasiliense Poir. Holoepiphytic climber. Rizome short creeping, woody, 5–15 mm thick, with many fine, densely scaly rootlets, the scales ovate-lanceolate, 8–12 mm long, often caudate-acuminate at apex, dull-brown. Leaves few, (sub)erect, 1-pinnate, ovate in outline, ca. 30–70 cm long, glabrous; stipe 8–50 cm long; segments 3–7 pairs, lorate, 8–40 x 1.2–3 cm, entire to crenate at margins, obtuse to cuneate at base, usually acute to acuminate at apex; veins areolate. Sori rounded, 1.5–2 mm diam., in 2 or 3 rows beside midrib.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

COLOMBIA. V: Killip 5238 (NY). N: (S). ECUADOR. E: Cornejo & Cangá 8144 (GUAY). Distribution: Southern Mexico to Bolivia. Ecology: In moist forests, wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: It is most likely that the epithet refers to the 3-seriate arrangement of sori beneath the pinnae, a variable feature of this species. PTERIDACEAE (Maidenhair Family) Plants terrestrial, epipetric, or epiphytic. Rhizomes erect or creeping, scaly or hairy; dictyostelic. Leaves monomorphic or dimorphic. Petioles with one vascular bundle or several. Blades 1- to 6-pinnate. Veins free or anastomosing. Sori borne on the lower surface of the lamina, usually spreading the veins. Indusia present or absent, when present formed by the revolute margin of the lamina (i.e., a false indusium; a true inner indusium is absent). Spores trilete, globose or globose-tetrahedral. Gametophytes epigeal, green. A cosmopolitan family represented in our area by Acrostichum and Vittaria. Acrostichum is comprised of three or more pantropical species (Arbeláez, 1996). Two Acrostichum species have been recorded growing in swamps and disturbed mangroves, often near sea level (Chapman, 1975; Arbeláez, 1996), both of which are known as “ranconcha” (Spanish), which is one of their most common vernacular names. They usually form dense clumps known as “ranconchales”, a derivation of “ranconcha”. Vittaria is a Neotropical genus, widely distributed in wet forest and rainforests. Key to the species 1. Terrestrial plants; leaf blades pinnate, 60–350 x 20–60 cm; sori covering all surface beneath pinnae 2. Fertile leaf blades usually less than 2 m long; distal pinnae fertile; pinnae rather distant and fewer than 30 Acrostichum aureum 2. Fertile leaf blades 1.5–3.5 m long; all or most of pinnae fertile; pinnae rather closely set and usually 40–60 Acrostichum danaeifolium 1. Epiphytic plants; leaf blades entire, 10–50 cm x 2–3 mm; sori linear-elongate, on submarginal grooves along blade Vittaria lineata

15. Acrostichum aureum L., Sp. Pl. 2: 1069 (1753). Plate 4: A, B. Syn.:

Acrostichum quineense Gaudich. Acrostichum inaequale Willd. Chrysodium aureum (L.) Mett. Chrysodium inaequale Fée Chrysodium vulgare Fée

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Vernacular names: chicharrón, chiguamacho, chascarra, lengua de vaca, racancha, ranconcha. Terrestrial herbs, caespitose, 0.7–2.5 m tall. Rhizomes forming tussocks in more or less saline swamps, clothed at apex with scales linear to ovate, rigid, dark-brown to 1.5 cm long. Stipe much shorter than blade, subterete, bearing several alternate indurated spurs; blade 60–200 x 20–40 cm, obtuse or somewhat acute in outline at apex; fronds monomorphic, the fertile blades with 1–4 pairs of distal pinnae bearing sporangia beneath; pinnae usually spaced apart, 10–14 pairs and a similar terminal one, oblong to oblong-lanceolate, 10–25 x 3–6 cm, cuneate at base, rounded-obtuse and emarginate or short-apiculate at apex; tissue coriaceous, light-green, glabrous. COLOMBIA. CH: Fuchs 21873 (COL, F, US). V: Gentry & Juncosa 40546 (COL, MO). C: Cabrera 3819 (CUVC). N: Mora 4184 (COL). ECUADOR. E: Cornejo & Bonifaz 2613 (GUAY [3], QCNE [3]). M: Neill et al. 11434 (QCNE). G: Bonifaz & Cornejo 3915 (GUAY [4]). PERU. T: (S). Distribution: Widespread in tropical and subtropical America, and in the Old World. Ecology: This species often forms pure stands in clayish, periodically flooded soils at river mouths and deltas (Arbeláez, 1996). It occurs in very dry, dry, moist, wet forest and rainforest life zones. Its abundant presence in some localities is probably related to clear-cutting that allows this fern to colonize extensive areas. It only occurs in disturbed sites, and is considered an agressive weed. Biological interactions: The populations of Acrostichum aureum provide shelter for the edible blue crab (Cardisoma crassum). The blue crab builds burrows underground, around, and in between the caespitose individuals of A. aureum, in the slightly higher and more consolidated soils from the mangrove backfringe to the adjacent terra firme inland ecosystems. IUCN Red list category: Characteristics of Acrostichum aureum, including a widespread distribution range, a common presence of abundantly healthy populations, a caespitose invasive habit, and an abundance of juvenile plants, lead me to deem the categorization of this species as of Least Concern (LC) (Polidoro et al., 2010). Etymology: The generic name is derived from the Greek words akros (summit), and stichos (row), referring to the fact that the sori are arranged only on the terminal pinnae in Acrostichum aureum, the type species of the genus. The specific epithet is a Latin word that means gold or golden, referring to the golden-yellow color of the underside of the terminal fertile pinnae. It was taken from Plumier (1703), who described the species as Filix palustris aurea, foliis linguae cervinae (cited in Linnaeus, 1753), that means golden fern from flooded areas, with leaves in the shape of a deer tongue. Notes: Acrostichum aureum has been called “mangrove fern” (Duke, 1992). This species is easy to recognize by its fronds of few distal fertile pinnae.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

16. Acrostichum danaeifolium Langsd. & Fish., Icon. fil. 5, t. 1 (1810). Plate 4: C, D. Syn.:

Achrostichum excelsum Maxon Achrostichum lomarioides (Jenman) Jenman Chrysodium lomarioides Jenman Chrysodium lomarioides var. hastatum H. Christ

Vernacular names: chicharrón, chiguamacho, chascarra, lengua de vaca, racancha, ranconcha. Terrestrial herbs, caespitose, to 4 m tall. Rhizomes massive, clothed at apex with scales, rigid, spreading, linear or narrowly-lanceolate, up to 2 cm long, light- to dark-brown. Stipe very stout and subwoody, grooved on adaxial side, dark-brown (at least on fertile fronds), clothed at base with broadly linear scales up to 2.5 cm x 3 mm, these bicolours, having a dark-brown longitudinal band in middle and paler, retrorsely fibrillose margins; blade 150– 350 x 20–60 cm, obtuse or somewhat acute in outline at apex; fronds dimorphic, the sterile ones 1.5–3.5 m long, the fertile ones taller and more rigidly erect; fertile blades usually with all pinnae bearing sporangia beneath; pinnae oblong, close, often somewhat imbricate, very numerous, usually 20–40 pairs or more and a similar terminal one; tissue chartaceous, often finely pubescent beneath. COLOMBIA. CH: (E). V: Gentry et al. 40427 (COL, MO). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 2646 (GUAY [3], QCNE). M: (E). Distribution: Widespread in tropical and subtropical America. Ecology: This species inhabit mangroves, riversides and fresh water flooded inland areas. It occurs in dry, moist, wet forest and rainforest life zones. IUCN Red list category: Characteristics of Acrostichum danaeifolium, including a widespread distribution range, a common presence of healthy populations, and a caespitose invasive habit, lead me to deem the categorization of this species as of Least Concern (LC) (Polidoro et al., 2010). Etymology: The epithet is formed by the fern genus Danaea (Marattiaceae) and the Latin word folium (a leaf blade), that refers to the resemblance of the leaves to those of Danaea. Notes: Acrostichum danaeifolium is the largest fern of the mangroves in our area. The species is easily recognizable by the large fertile blades, with most or all pinnae fertile on the underside. 17. Vittaria lineata (L.) Sm., Mem. Acad. Roy. Sci. (Turin) 5: 421, pl. 9 (1793). Plate 3: D. Bas.: Pteris lineata L. Syn.: Vittaria filiformis Cav. Holoepiphytic herbs, caespitose. Rhizome with tomentous roots, densely covered by scales linear to linear-lanceolate, 3–6 mm long, dark-brown to blackish, and hairs, ca. 1 mm long,

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

yellowish-brown. Leaves suberect; stipe 1.5–3 cm long; blade simple, linear-lorate, 10–50 cm x 2–3 mm, acute at apex, entire at margins, inconspicuously veined, glabrous. Sori elongate, on submarginal grooves along blade. COLOMBIA. CH: Fuchs et al. 22248 (COL). V: Juncosa 2092 (MO). N: (E). ECUADOR. E: Cornejo & Bonifaz 6949 (AAU, GUAY). Distribution: Mexico to Paraguay. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the linear-lorate leaf blades. Note: This genus has been previously placed in the family Vittariaceae.

Gymnosperms ZAMIACEAE Stem subterranean to arborescent, when arborescent smooth or clothed in persistent leaf bases. Ageotropie eoralloid roots. Leaves with or without small stipules, paripinnately compound, in a crown of few (1–15) to many (30); petiole and lower third of rachis smooth or with small prickles or with reduced spinose pinnae; pinnae 10–120 in subopposite to opposite pairs, articulated with rachis, densely pubescent when young, glabrous when mature, venation dichotomous, with or without a midrib. Plants dioecious. Strobili one to several, usually emerging from center of crown; sporophylls flat to peltate. Pollen strobili of numerous microsporophylls with sporangia located on the abaxial surface and rarely on the adaxial surface; microsporangia many, in groups of 2–5 and dehiscing by longitudinal slits. Ovulate strobili of numerous ovule-bearing, stalked megasporophylls, each with two ovules. Seedcoat when mature consisting of a red or orange/red or white outer fleshy layer and an inner stony layer; megagametophyte farinaceous; embryo with two cotyledons and a coiled suspensor. 18. Zamia roezlii Linden, Cat. Général 9: 10 (1873). Syn.: Aulacophyllum roezlii (Linden) Regel Vernacular names: chigua, piña de monte, zamia de los manglares. Arborescent plants to 7 m tall, sometimes subepiphytic. Leaves 5–10, 1–3 m long; petiole 0.5–1 m long, sparsely to densely prickled; rachis with 10–20 pairs of leaflets, occasionally with a few prickles in the lower third. Leaflets linear-lanceolate, grooved between veins on adaxial surface, cuneate at base, acute at apex, entire, 30–50 cm long, 12–15 cm wide. Pollen strobili cream to tan,

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

cylindrical to elongate-cylindrical, 20–30 x 4–6 cm. Ovulate strobili brown, short pedunculate, cylindrical to ovoid cylindrical, 30–50 x 10–20 cm. Seeds red, ovoid, 1.5–2.5 cm diam. COLOMBIA. V: Gentry & Juncosa 40710 (MO, NY, US). Distribution: Coastal Colombia and contiguous coastal Ecuador. Ecology: This is a salt-tolerant species. It occurs near sea level, from the terra firme forests to the mangroves, in the wet forest and rainforest life zones. IUCN Red list category: This species is often locally abundant. “It produces very large seed cones and thus numerous seeds and seedlings. However, seed cones are produced by arborescent plants and under continual disturbance seed production would be severely diminished, resulting in a high vulnerability” (Stevenson, 2004). This species has been categorized as Vulnerable (VU B1abiii) (Galeano et al., 2005). Etymology: The epithet honors Benedikt Roezl (1824–1885), a Czech botanist, the collector of the type. Note: Zamia roezlii is the largest South American Zamia.

Seed Plants ACANTHACEAE (Acanthus Family) Herbs, shrubs, trees or vines. Leaves simple, usually decussate or opposite, simple; exstipulate. Inflorescences axillary or terminal, solitary, spicate, racemose, paniculate or cymose. Flowers bisexual, usually zygomorphic, bracteate. Calyx deeply lobed. Corolla tubular, bilabiate. Stamens 4 and didynamous or 2, attached to corolla tube. Ovary superior. Fruit capsular, dehiscent by 2 valves.

This family is represented in mangroves by Avicennia, a pantropical genus comprised of eight

species. In our area two species have been reported, A. bicolor and A. germinans, but only A. germinans has been found. Avicennia has been traditionally placed in Verbenaceae or even in its own family, Avicenniaceae. However, molecular work demonstrates that Avicennia is nested in Acanthaceae (McDade et al., 2008). Key to the species 1. Flowers opposite, loosely arranged in separate pairs along rachis 1. Flowers polystichous, densely arranged along rachis

Avicennia bicolor

Avicennia germinans

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Avicennia bicolor Standl., J. Wash. Acad. Sci. 13(15): 354 (1923). Vernacular names: mangle negro, palo de sal. Trees or shrubs 1–20 m tall, to 30 cm dbh. Secondary superficial roots with many pneumatophores, these narrow, pencil-thick, erect to 30 cm high, lenticellate. Terminal branchlets with bark which can be pealed out as long narrow strips. Leaves arranged along branches; blades broadly-elliptic to ovate, 6–15 x 3–8 cm, obtuse to cuneate at base, rounded or obtuse at apex, olive-green and glabrous above, dull and with a minute withish indument beneath, the lateral veins 6–12 per side, eucamptodromous; petioles 0.4–2 cm long. Inflorescences terminal panicles, 3–17 cm long, these bearing to 4th order branches, the spikes 1.5–10 cm long, the primary peduncle to 4 cm long; flowers sessile, opposite or decussate, loosely arranged in pairs, each pair separate 4–10 mm long. Calyx dialisepalous, sepals 5, peristent in fruit, green, covered by a dense appressed pubescence. Corolla tubular with spreading lobes, white to creamish-white, sericeous without, the lobes 3–5 mm long, the throat yellow within. Stamens 4, alternipetalous, 3–4 mm long, the filaments adnate at corolla tube base. Ovary superior, ovate, ca. 2.5 mm long, cream, pubescent; style ca. 1.5 mm long, white; stigma bifid. Fruit barely asymmetric, lateraly compressed, ca. 2 x 1.3 cm, dehiscent in 2 valves (after having fallen in the water), greenish-yellow without; seed 1, without testa, cotyledons 2.

This is a Mesoamerican species that inhabits the Pacific side of the American continent, from Nicaragua to Panama. It has been reported to occur in the mangroves at the mouth of the Buenaventura River on the Pacific coast of Colombia (Tomlinson, 1986). However, herbarium collections that would support the presence of Avicennia bicolor in western Colombia have not been found. Erroneous reports of its presence there could have originated from misidentified collections of Avicennia germinans from the same area (e.g., Gentry & Juncosa 40586 [MO, NY], det. as A. bicolor by Gentry, 1983). 19. Avicennia germinans (L.) L., Sp. Pl. ed. 3: 891 (1764). Plate 5: A–E. Bas.: Bontia germinans L. Syn.: Avicennia nitida Jacq. Avicennia tomentosa Jacq. Avicennia tomentosa var. guayaquilensis Moldenke Avicennia germinans var. guayaquilensis (Kunth) Moldenke Avicennia germinans var. cumanensis (Kunth) Moldenke Avicennia tonduzii Moldenke Avicennia elliptica Holm in Thunb. Avicennia floridana Raf. Avicennia lamarckiana C. Presl Avicennia meyeri Miq. Avicennia oblongifolia Nutt. ex A. W. Chapm.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

Vernacular names: comedero, jelí salado, mangle de humo, mangle llorón, iguanero, mangle iguanero, mangle negro, mangle prieto, mangle salado, palo salado, salado. Trees or shrubs 0.5–20(–25) m tall, to 120 cm dbh. Bark black to grey or dark-brown. Secondary superficial roots with many pneumatophores, these narrow, pencil-thick, erect to 40 cm high, lenticellate. Terminal branchlets with bark which can be pealed out as long narrow strips. Leaves arranged along branches; blades lanceolate to narrow-elliptic or oblong-elliptic, 7–18 x 1.5–6 cm, cuneate at base, acute to obtuse at apex, olive-green and usually shiny above, dull beneath, the lateral veins 10–14 per side, brochidodromous, glabrous on both sides; petioles 1–2 cm long. Inflorescences terminal and axillary, in dense panicles or spikes 1–5 cm long, the spikes 1–2.5 cm long, sessile or on peduncles to 2.5 cm long; flowers sessile, polystichous, densely arranged. Calyx dialisepalous, sepals 5, peristent in fruit, green, covered by a short appressed pubescence. Corolla campanulate, white to creamish-white, pubescent without, the lobes 3–5 mm long, the throat yellow within. Stamens 4, alternipetalous, ca. 4 mm long, the filaments adnate at corolla tube base. Ovary superior, ovate, ca. 1.2 mm long, cream, pubescent; style ca. 2 mm long, white; stigma bifid. Fruit asymmetric, lateraly compressed, 1.5–2.5 x 1–1.7 cm, dehiscent in 2 valves (after having fallen in the water), rugose, greenish-yellow without; seed 1, without testa, cotyledons 2, the embryo with radicle white, densely-pilose. COLOMBIA. CH: Gentry & Juncosa 40941 (COL, MO). V: Gentry & Juncosa 40586 (MO, NY). C: (S). N: Romero-Castañeda 5309 (COL, MO). ECUADOR. E: Cornejo et al. 7226 (GUAY, QCNE). M: Bonifaz & Cornejo 3994 (AAU, GUAY, QCNE). SE: (S). G: Madsen 63975 (AAU [wood collection available], MO, NY, QCA, QCNE). O: Cornejo & Bonifaz 5984 (GUAY, QCNE). GAL: Darwin s.n. (CGE). PERU. T: Vargas 3945 (NY). P: (S). Distribution: Widely distributed along the tropical shores of West Africa and the Caribbean region, and from southeastern United States (southern Florida and southeastern Texas) to Brazil, as well as along the west coast of the Americas from Mexico to northwestern Peru, also in the Galapagos islands. Ecology: Semirestricted to mangroves and occasionally also growing on sandy beaches, in desert scrub, dry thorn scrub, very dry, dry, moist, wet forests, and rainforests. Phenology: This species has been collected flowering throughout the year, and fruiting generally from June to February. The flowers remain open 3 to 5 days and emit fragrance both day and night. When the flowers are just opening, the anthers are a pure white color, but darken to a pure black over the ensuing hours. The pollen has been observed to be released ca. 20 hours after the flower’s openning. Biological interactions: At least 25 species of insects have been observed visiting the flowers of black mangrove during the day (see Plate E). The flowers of Avicennia germinans are mainly pollinated by honey bees, Apis mellifera (Apidae). Other common visitors and most likely secondary pollinators that feed on the nectar of black mangrove flowers are diurnal Syrphid flies as Palpada vinetorum, P. albifrons, P. aemula, P. sp. 1 and P. sp. 2, several unidentified species of flesh flies (Sarcophagidae), green bottle flies (Calliphoridae), lady beetles as Olla roatanensis and Paraneda sp. (Coccinellidae), and butterflies

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

as Melanis leucophlegma (Riodinidae). The flowers are also often visited by wasps (Vespidae), small bees (Halictidae), several species of ants and tiny flies, and occasionally by small beetles as Hyperaspis onerata, H. esmeraldas (Coccinellidae) and black scavenger flies (Sepsidae). Jumping spiders (Salticidae) could be often found hidden among the spikes or flowers or even feeding on floral nectar. A video about some of the mentioned species visiting flowers of black mangrove is available in this link: https://www. youtube.com/watch?v=dad_ILLB8lo In addition, many species of birds and insects have been observed resting on the branches and leaves of A. germinans; among those some insects (e.g., Hemiptera) lay eggs on the underside of leaves. The terminal branchlets and leaves are reportedly eaten by goats (Madsen et al., 2001). One of the vernacular names, “iguanero”, a word derived from the common iguana (Iguana iguana), is said to refer to the reptile occasionally feeding on the young leaves and buds of this species. Snails of Littorina varia (Littorinidae) occasionally feed on the leaves of A. germinans. Dispersal: Avicennia germinans is a water-dispersed species. According to Moldenke (1980), it usually drifts as seedlings, not as seeds or fruits. The pericarp of the fruit acts as a surrogate seed coat, since the actual seed coat is absent. The embryo germinates while the fruit is still attached to the parent tree. Afterwards, the fruit falls from the tree, and the embryo absorbs moisture from the water or mud environment in which it lands, causing the fruit coat to tear. One to two days after the fruit coat has been shed, the radicle grows out from the cotyledons, revealing a curvature with positive geotropism, regardless of whether the seedling is floating on water or stranded in mud. Uses: The wood is hard and incorruptible, but considered of lesser quality than that of Rhizophora spp. (Acosta-Solís, 1944). It is used to construct canoes and is made into charcoal. When the wood is cut, an exudate is produced that is reported to be an eye irritant (Romero-Castañeda 5309, MO). Avicennia germinans can concentrate heavy metals as: Cadmium (Cd), cupper (Cu), lead (Pb), and zinc (Zn) (Campos and Gallo, 1997; Villamil, 2010). IUCN Red list category: According to the Registro Oficial No. 148, issued on March 16 1993, Avicennia germinans has been categorized as an endangered species in Ecuador (INEFAN/GEF, 1998). However, due to factors such as a widespread range of distribution, a common presence of healthy populations usually comprised of abundant individuals, a high rate of seed viability, and an abundance of observed seedlings and juveniles, it is deemed the categorization of this species as of Least Concern (LC) (Polidoro et al., 2010). Etymology: The genus is named in honor of Abū Alī al-Husain ibn Abd Allāh ibn Sīnā (Abu Ali Alhosian Ben Sina, 980–1037), also known as “Ibn Sina”, or more commonly by his latinized name “Avicenna”, a famous Persian physician, mathematician, philosoper, and naturalist (Moldenke, 1960). The specific epithet is a Latin word that refers to the fact that the seeds of this species germinate while still attached to the mother plant. Linnaeus chooses the epithet based on the field observations made by Patrick Browne in Jamaica (1756). Discussions: There has been disagreement over the correct name of this species. Little (1961) argued that it should be A. nitida. However, the nomenclatural decision, the correct one, ended up being A. germinans (Compère, 1963). This is a vegetatively variable species. The several taxa proposed by Moldenke in

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

Avicennia seem to be of no taxonomical value; therefore they have not been recognized here and are placed in the synonymy. A. tonduzii has been cited as a synonym of A. bicolor (Tomlinson, 1986). However, the type collection of A. tonduzii (Tonduz 6776, US) matches A. germinans rather than A. bicolor. Notes: Occasionally, mature individuals of Avicennia germinans may have a very different habit in the same locality, varying from small shrubs 0.5 m to trees 6 m high. The upper surface of the leaf blades has been often observed to be wet during the day, most likely caused by salt excretion. AIZOACEAE Annual or perennial herbaceous or suffruticose plants, prostrate or upright, often succulent. Leaves usually simple, alternate, opposite or pseudoverticillate, often fleshy, entire; stipules scarious or absent. Inflorescences axillary or terminal, modified cymes or flowers solitary. Flowers bisexual or unisexual, actinomorphic. Perianth monochlamydeous, usually 4- or 5-parted, free or connate, often persistent in fruit. Stamens 4–5 or many, the filaments free or connate. Pistil 1, the ovary superior to inferior. Fruit capsule or indehiscent and either baccate or nut-like. 20. Sesuvium portulacastrum (L.) L., Syst. Nat. ed. 10: 1058 (1759). Plate 6: A. Vernacular name: verdolaga. Terrestrial herbs, perennial, succulent, prostrate, glabrous throughout. Branches trailing and often rooting at nodes. Leaves linear-elliptic or narrowly-obovate, 1–5 x 0.3–1 cm, gradually tapered to base, subacute at apex; petioles 1–5 mm long, dilated at base, clasping stem. Inflorescences axillary, the flowers solitary, pink; pedicels 2–11 mm. Perianth tube 1.5–3 mm long, the lobes ovate, 4–7 mm long, persistent and somewhat reflexed in fruit. Stamens numerous, inserted at mouth of perianth tube, the filaments 1.5–3.5 mm long. Ovary ovoid to subglobose, 3–3.5 mm long; styles 3 or 4. Capsule ovoid to obovoid, ca. 6.5 mm long; seeds 8–30, 1–1.5 mm diam., black. ECUADOR. E: Løjtnant & Molau 15229 (AAU). M: (S). SE: (S). G: Iltis et al. E–619 (WIS). O: HolmNielsen 22823 (AAU). PERU: T: (S). P: (E). Distribution: Circumtropical. Ecology: In Ecuador and adjacent northwestern Peru, Sesuvium portulacastrum is distributed along coastal dry forests, often as a common, prostrate and succulent halophyte that inhabits open, sunny places just above the uppermost tidal level, usually behind the mangroves, although at times in the interior when the mangroves are disturbed. It is relatively common at the edge of shrimp pools, where it often forms pure beds. It is also present in salt marshes and on sandy beaches. This species mostly inhabits dry thorn scrub, very dry, and dry forests. Phenology: The flowers are fully open early in the morning, but are half-closed at the full sun of midday and afternoon.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Biological interactions: Domestic bees have been observed visiting the flowers. Use: This species is cultivated as an ornamental in the Galapagos islands. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the the genus Portulaca (Portulacaceae) and the Latin suffix astrum (related), referring to the resemblance of this species to those of Portulaca. AMARANTHACEAE (Amaranth Family) Herbs, shrubs, small trees or lianas. Stems prostrate to erect or scandent. Leaves oposite or alternate; blade generally entire. Inflorescences terminal and/or axillary, generally cylindrical to head-like spikes or cymes or cyme-like clusters, these simple or arranged in open to compact spiciform, paniculate, or head-like compound inflorescences. Flowers bisexual or unisexual; each flower subtended by a ventral bract and two lateral bracteoles. Tepals mostly 3–5. Stamens mostly as many as tepals and opposite these. Ovary ovoid to cylindrical or subglobose, unilocular; styles mostly 1–2; stigma entire, bilabiate or capitate. Fruit mostly a dry utricle, sometimes a berry-like capsule. Key to the species 1. Vines; leaf blades elliptical, ovate to lanceolate, 7–15(–20) x 3.5–7(–8) cm

Chamissoa altissima

1. Herbs; leaf blades inconspicuous, lorate, 5–10 x 1–2 mm, or rudimentary, reduced to minute scales 2. Stem segmented-articulated, glabrous; leaves rudimentary, reduced to minute scales 2. Stem not segmented, pubescent; leaves lorate

Salicornia fruticosa Suaeda sp.

21. Chamissoa altissima (Jacq.) Kunth, Nov. Gen. Sp. Pl. 2(7): 197, t. 125 (1818). Bas.: Achyranthes altissima Jacq. Syn.: Celosia paniculata L.

Chamissoa macrocarpa Kunth

Celosia tomentosa Willd. ex Roem. & Schult.

Chamissoa altissima (Jacq.) var. rubella Suess.

Chamissoa frondosa Hashim. Vines. Leaves alternate; blades elliptical, ovate to lanceolate, 7–15(–20) x 3.5–7(–8) cm, rounded to acute at base, obtuse to long-acuminate at apex, membranous to subcoriaceus, mostly glabrous on both sides, the lateral veins 5–10 per side; petioles mostly 1–5 cm long. Inflorescences terminal and axillary, flowers mostly functionally unisexual, greenish-white;

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

bracts scarious, subdeltoid to ovate, up to ca. 2.2 mm long; bracteoles shorter than bracts, scarious, subdeltoid to round-triangular, slightly to prominently curved at apex. Tepals subelliptic to ovate-lanceolate, whitish-green to reddish, 2.5–3.5 mm long, with 1–3(–5) sometimes very faintly marked veins. Functional anthers up to 1 mm long, the filaments united at base into a low cup ca. 0.2 mm deep. Ovary barrel-shaped, narrowed at base and occasionally surrounded by a spongy collar, narrowed or truncate at apex; style and stigmas ca. 0.5 mm long. Fruit capsule, 3–4 mm long; seeds shiny, black, mostly totally enclosed in a bivalved whitish aril. ECUADOR. E: Cornejo & Bonifaz 1900 (GUAY). Distribution: Mexico to Argentina, and in the West Indies. Ecology: Chamissoa altissima is mostly a terra firme colonizing species, and a rather infrequent element within the hyposaline mangroves. It grows from sea level to about 2,000 m (Eliasson, 1987). It occurs in dry, moist, wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means very high. It is most likely refers to the high places that the vine is able to reach in its growth. 22. Salicornia fruticosa L., Sp. Pl. ed. 2, 5 (1762). Plate 6: B. Syn.:

Salicornia europaea var. fruticosa L. Salicornia peruviana Kunth Salicornia biloba Kunze ex Fenzl. Salicornia equisetifolia Willd. ex Moq. Salicornia gaudichaudiana Moq. Salicornia neei Lag. Arthrocnemun fruticosum (L.) Moq.

Vernacular names: parachique, vidrillo, vidrillo salado. Terrestrial herbs, perennial, suffrutescent, 0.3–1.5(–2) m tall. Stems and branches lightgreen, glabrous, fleshy; profusely branched in a candelabrum-like fashion, with ascending or erect articulate-jointed branches. Leaves rudimentary, reduced to minute scales. Sterile joins 7–15 mm long; sheats with very obtuse-angled lobes. Flowering joints 2–4 mm long, forming terminal, cylindrical spikes. Flowers perfect or polygamous, in groups of 3–7, sunken in joints of stems. Perianth obpyramidal, outer part rhomboid or rounded, ca. 2 mm across. Stamens 1–2, the anthers 0.8–1 mm long. Stigmas ca. 0.5 mm. Seeds 1.3–1.5 x 1 mm, hispidulous. ECUADOR. G: Cornejo & Castro 287 (GUAY, QCNE). M: (S). PERU. T: Smith 10077 (MO). P: (E). Distribution: Widely distributed on seashores throughout the world. Ecology: A pioneer in disturbed mangroves, it also occasionally grows on beaches. It occurs in dry thorn scrub, very dry, and dry forest life zones.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Biological interactions: The plants are eaten by donkeys (Madsen et al., 2001). IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means shrubby. Note: This species has been traditionally placed in the Chenopodiaceae family. 23. Saueda sp. Plate 6: C. Herbs, to 30 cm tall, forming broad patches. Stem succulent, pubescent, woody at base. Leaves simple, alternate; blades lorate, 5–10 x 1–2 mm, inconspicuous. Filaments yellow. Fruit 4–5 parted, hemispherical, 4 mm wide. ECUADOR. G: Madsen 104085 (AAU). Ecology: A rare halophytic weed, growing on destroyed mangrove terrain (Madsen, pers. comm.). It occurs in very dry and dry forests. IUCN Red list category: Data Deficient (DD). Notes: This genus has been traditionally placed in the Chenopodiaceae family. The description of the species has been taken from the publication Flora of Puná Island (Madsen et al., 2001). AMARYLLIDACEAE (Daffodil Family) Bulbous herbs. Leaves sessile and linear or lorate, or petiolate and lanceolate to widely elliptic, sometimes basally sheating and forming an aerial pseudostem. Inflorescence scapose, umbellate; scape terminated by two, seldom one, or more, spathaceous, valvate-imbricate or obvolute, usually marcescent bracts that enclose flowers in bud; inner bracteoles sucessively shorter and narrower. Flowers 1 to many, perfect, frequently large and showy, each usually subtended by a bracteole, actinomorphic or zygomorfic. Perianth consisting of 6 usually subequal petaloid segments in two series, connate below into a tube or rarely free almost to base. Stamens 6, free or variously connate. Ovary inferior; style filiform; stigma capitate, 3-lobed or deeply trifid, usually papillate. Fruit a dehiscent capsule, rarely baccate. 24. Crinum kunthianum M. Roem., Syn. Ensat. 80 (1847). Plate 6: D. Syn.: Crinum erubescens Kunth (non L. f. ex Dryander in Aiton). Terrestrial herbs. Bulb ellipsoid, with a short neck. Leaves rosulate, spreading; blades sessile, ligulate, 60–80 x 2–5(–8) cm, slightly undulate, the margin inconspicuosly and loosely denticulate. Scape 50–80(–100) x 1–2 cm; bracts 2, lanceolate-deltoid, 8–11 x 1.5–2.5 cm, greenish-white; bracteoles linear, whitish. Flowers 4–5(–10), fragrant, sessile to shortly pedicellate. Tube 10–15 x ca. 0.5 cm, abruptly dilated near throat to 10 cm, yellowish-green; tepals strongly reflexed, 7.5–9(–10) x 0.5–1 cm, apiculate at apex, cream. Filaments 5–7 cm

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

long, spreading, deep-pink to dark-purple distally, the anthers linear, 5–7(–10) mm, purple. Ovary oblong, 1.5–2 cm long. Fruit capsule, ellipsoid, succulent, 3-loculicidal. COLOMBIA. V: Killip & Cuatrecasas 38827 (MO). N: (E). ECUADOR. E: Cornejo & Bonifaz 7260 (GUAY). G: Bonifaz & Cornejo 3911 (GUAY [2]). Distribution: Colombia and Ecuador. Ecology: This colonizer is often a gregarious species, growing in carpets on swampy soils, mudflats and muddy river banks, just above the uppermost tidal level, occasionally inhabiting the interior of hyposaline mangroves. It occurs in dry, moist, wet forest and rainforest life zones. Phenology: The flowers of this species are very fragrant in the evening. IUCN Red list category: Least Concern (LC). Etymology: The epithet honors Karl Sigismund Kunth (1788–1850), a German botanist who described many of the new species gathered from northern South America by Humboldt and Bonpland in Nova Genera et Species Plantarum (1815–1825). Notes: Crinum kunthianum can be recognized by the terrestrial herbaceous habit in combination with the rosulate arrangement of the sessile ligulate leaves, and the inflorescences arising from a well-developed central scape, which bear a bouquet of showy white flowers. In the treatment of Amaryllidaceae of Flora of Ecuador, Crinum kunthianum has been described as having leaf blades with smooth margins, rarely bearing more than 6 (to 7) flowers (Meerow, 1990). However, the cited specimens differ in that they have loosely-denticulate margins; furthermore, the specimen Bonifaz & Cornejo 3911 has 10 flowers per inflorescence. ANNONACEAE (Clustard-Apple Family) Trees, shrubs, and rarely lianas. Leaves simple, alternate, usually distichous, pinnately veined, the margins entire; exstipulate. Inflorescence leaf-opposed, internodal or axillary, sometimes cauliflory and flagelliflory. Flowers solitary, in fascicles or cymose, bisexual or frequently unisexual. Sepals 3; petals usually 6, arranged in two whorls of 3. Stamens numerous and spirally arranged, the filaments usually short. Carpels numerous, rarely 1. Fruits usually of separate, stipitate, fleshy, indehiscent monocarps, or carpels coalescent to form an aggregate fruit.

A family represented by a tree species in mangroves.

25. Annona glabra L., Sp. Pl. 1: 537 (1753). Plate 7: A. Syn.: Annona palustris L. Vernacular names: guanábana de pozo, guanabanillo.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Trees to 10 m tall. Buttresses narrow, occasionally present. Leaf blades (sub)coriaceous, elliptic, 6–20 x 3–8 cm, acuminate at apex, glabrous, the lateral veins 12–17 per side; petioles 0.6–1.5 cm long. Flowers arranged along branches, extra-axillary. Sepals green, widelyovate, 3–5 mm, glabrous. Corolla of 6 fleshy petals, yellowish or yellow, glabrous outside, the outer petals ovate, 2.5–4 cm long, carinate, the inner petals somewhat smaller, purple at inner base. Fruit subglobose, fused carpels, to 12 x 10 cm, externally smooth, light green, turning yellow at maturity, glabrous. COLOMBIA. CH: (E). V: Cuatrecasas 16193 (F). C: (E). N: Cabrera 4107 (CUVC). ECUADOR. E: Cornejo et al. 7240 (GUAY). G: Cornejo & Bonifaz 5154 (GUAY). O: (S). PERU. T: (E). Distribution: Southeastern United States (Florida) and Mexico, to Ecuador and Brazil, and in the West Indies; also native in tropical West Africa. Ecology: Annona glabra occurs in secondary forests, from the intertidal to supratidal levels, on swampy soils of the hyposaline mangroves, in dry, moist, wet forests and rainforests. Biological interactions and dispersal: The saline-tolerant water-dispersed fruits and seeds are eaten by animals as fruit-bats and fish. Use: The fleshy mesocarp from mature fruit is mixed with water and made into a refreshing beverage. It is said that has a flavor similar but simpler to that of cherimoya (Annona cherimola), or sweet apple (A. squamosa). IUCN Red list category: Due to the widespread range of distribution and the common presence of healthy populations, this species is herein categorized as of Least Concern (LC). Etymology: The epithet is a Latin word that refers to the entire absence of pubescence in the fruits. Note: Taking into account such factors as a wide range of distribution throughout semiaquatic environments, a tolerance to salinity that allows successful settlement and germination in hyposaline environments, and a water dispersal system for fruits and seeds, this species is herein regarded as a facultative mangrove. APOCYNACEAE (Dogbane Family) Trees, shrubs, lianas, and herbs. Milky latex usually present. Leaves simple, entire, often opposite, less common alternate or verticillate. Flowers perfect, regular. Calyx lobes 5, distinct to base or nearly so. Corolla cylindrical-campanulate, tubular-salverform, or funnelform; lobes 5, convolute in bud. Stamens 5, inserted on corolla, the anthers often connivent and somewhat adherent to stigma. Pistils 2, separate; ovary superior or nearly so; stigmas often united in flower. Fruit a pair of erect or divergent follicles or drupe; seeds 1 to many.

A family represented by a species of liana in mangroves.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

26. Rhabdadenia biflora (Jacq.) Müll. Arg. in Mart., Fl. Bras. 6(1): 175 (1860). Plate 7: B. Bas.: Echites biflora Jacq. Syn.: Echites paludosa Vahl

Rhabdadenia paludosa (Vahl) Miers

Rhabdadenia macrantha Donn. Sm.

Vernacular names: bejuco de cerca, garza, lechera, lecho de mar. Lianas. Latex white. Stems many-lenticellate, glabrous. Leaves simple, opposite; blades firmly chartaceous to coriaceous, oblong or narrowly-lanceolate to elliptic, 3.5–12 x 1.5–6 cm, obtuse to cuneate at base, apiculate at apex, glabrous, the lateral veins 10–20 per side, brochidrodomous, the tertiary venation reticulate; petioles 8–15 mm long. Inflorescences terminal and axillary, 1–2-flowered, conspicuous; pedicels 1–2 cm long. Calyx dialisepalous; sepals 5, linear-lanceolate, 5–9 x ca. 1.5 mm, glabrous. Corolla infundibuliform, 6–8 cm long, white to pale pink, glabrous; petals 5, asymmetric, the lobes 1-toothed; corolla tube 2–2.5 cm long, yellow within. Stamens inserted at apex of corolla tube. Fruit follicles, linear, to 14 cm long, terete; seeds numerous, brown, wind-dispersed. COLOMBIA. CH: Haught 5416 (COL, MO). V: Killip & Cuatrecasas 38622 (COL, MO). C: (E). N: Madrigal & Mesa 661 (COL). ECUADOR. E: Cornejo & Bonifaz 6972 (AAU, GB, GUAY, QCA, QCNE). M: Cornejo & Bonifaz 7174 (AAU, GB, GUAY). Distribution: Southeastern United States (Florida) and Mexico to Ecuador and Brazil. Ecology: Rhabdadenia biflora grows in open places, most commonly on river banks. In our area, it is mostly distributed in moist, wet forest, and rainforest life zones. Phenology: This species has been collected in flower from January to August. The flowers are ephemeral. Dispersal: The seeds are wind dispersed. Uses: In some areas of the Province of Esmeraldas, Ecuador, the entire plant with its flowers is boiled as a beverage for women whose milk has not yet come in after the primogenial birth. It is said that three days after drinking the beverage the breasts will begin producing plenty of milk (Perea, pers. com.). The species is called “lechera” because its white latex resembles milk. In the province of Manabi the stem is used to tie fences, explaining its vernacular name “bejuco de cerca”. IUCN Red list category: This widespread species is often represented in mangroves by healthy populations of numerous individuals, and is therefore categorized as of Least Concern (LC). Etymology: The epithet is formed by the Latin prefix bi (two) and the Latin word flora (flower) that refers to the two-flowered inflorescences. Notes: Among the mangrove lianas, Rhabdadenia biflora is easily recognized by its beautiful, showy white flowers, as well as its vegetative characteristics such as opposite leaves and white latex. It is considered

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

a terrible weed, because upon being cut by a machete, it bleeds profuse latex which permanently stains clothing (Cangá, pers. com.). The cited specimens represent the first record of this species for Ecuador (Cornejo, 2003). ARACEAE (Arum Family)

Terrestrial, epiphytic, or aquatic herbs, perennial. Milky or acrid sap sometimes present. Stem

subterraneous and rhizomatous or cormous, or else the stem aerial and erect, creeping or climbing, often with aerial roots. Cataphylls deciduous or persistent. Leaves alternate, rarely solitary, basal or cauline, sometimes distichous, simple or rarely compound; blades entire, lobed or dissected, sometimes perforated. Inflorescence an unbranched spadix subtended by a single, herbaceous spathe. Flowers small sessile, ebracteate, bisexual or unisexual, when unisexual the lower flowers pistillate and the upper ones staminate; perianth absent or of usually 4–6. Stamens generally 3–6. Ovary usually superior. Fruits baccate.

This family is represented in mangroves by at least eigthteen epiphytic species, all of which have

great ornamental potential. None of the species is restricted to mangrove ecosystems; rather they are mostly distributed in the adjacent terra firme forests, in moist, wet, and rainforest life zones.

The fleshy, juicy berries of this family’s species form a congested arrangement at the distal portion

of usually elongate peduncles, which acts to tempt potential dispersers. They are reportedly eaten and the seeds endozoochorously dispersed by bats, birds and monkeys (Croat, 1978; Lobova et al., 2009). As far as it is known there are no studies of those interactions in the mangroves of our area.

Key to the species 1. Leaves subrosulate, suberect-spreading, usually forming a cup-like; the petioles less than one fifth of the blade length 2. Leaf blades (60–)90–150 cm long, not punctate beneath; infructescences pendulous, brown

Anthurium asplundii

2. Leaf blades 25–81 cm long, punctate beneath; infructescences spreading to suberect, red Anthurium hacumense 1. Leaves alternate or spiral, not forming a cup-like, the petioles one fifth of the blade length or usually longer 3. Leaf blades simple 4. Leaf blades deeply pinnatifid into narrow lobes

Monstera pinnatipartita

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

4. Leaf blades entire 5. Stems longer than 5 meters; lateral veins closely parallel and strongly ascendant 6. Peduncles of inflorescences 6–10 cm long; spadix 2–5 cm long Stenospermation angustifolium 6. Peduncles of inflorescences 24–58 cm long; spadix 7–13.5 cm long Stenospermation robustum 5. Stems of usually less than 5 m long; lateral veins never closely parallel and strongly ascendant 7. Pistillate flowers arranged together at base of spadix, the staminate flowers arranged together at apex 8. Leaf blades ovate-cordate to ovate-triangular, 25–57 cm wide; cataphylls unribbed; peduncle of inflorescence 8–21 cm long

Philodendron sparreorum

8. Leaf blades elliptic or oblong, 5–12 cm wide; cataphylls sharply 2-ribbed; peduncle of inflorescence 2–5 cm long 9. Leaf blades elliptic, 23.5–30 cm long, attenuate at base; spadix 8–15.5 cm long Philodendron chrysocarpum 9. Leaf blades oblong, 35–50 cm long, cordate at base; spadix 16–19 cm long Philodendron musifolium 7. Flowers homogeneously arranged along spadix 10. Leaf blades punctate beneath 11. Cataphylls non fibrous 12. Cataphylls soon deciduous Anthurium littorale 12. Cataphylls persistent

Anthurium paludosum

11. Cataphylls fibrous 13. Leaf blades profusely verrucose (with 10 x magnification) above Anthurium verrucosum 13. Leaf blades not verrucose above 14. Peduncle of inflorescence 2–8 cm long; spadix 2–7 cm long; spathe 2–4 cm long; berries white at maturity

Anthurium obtusum

14. Peduncle of inflorescence 22–43 cm long; spadix 9.5–30 cm long; spathe 4.5–8 cm long; berries orange to bright red at maturity 15. Petioles 3.3–15(–19) cm long, D-shaped in cross section; inflorescence pendulous; spadix 9.5–21 cm long

Anthurium cordobense

15. Petioles 15–28(–33) cm long, sharply triangular in cross-section; inflorescence erect; spadix 22–30 cm long

Anthurium oxyanthum

10. Leaf blades not punctate beneath

16. Leaf blades suberect, narrowly obovate or oblanceolate, elliptic, ovate-elliptic to oblong-elliptic, 4.5–18 cm wide; inflorescences erect

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

17. Leaf blades with midvein reddish to red beneath; spathe 1.5–2.5 cm wide Anthurium incomptum

17. Leaf blades with midvein greenish to yellowish beneath; spathe 0.7–1.5 cm wide Anthurium guayaquilense

16. Leaf blades pendulous, lorate, 0.8–4 cm wide; inflorescences pendulous Anthurium friedrichsthalii

3. Leaf blades 3-parted or deeply 3-lobed 18. Leaf blades 15–25 cm long, the central lobe without a submarginal vein distally; spadix 6–11 cm long, surrounded by spathe Philodendron tripartitum 18. Leaf blades [21–]40–83 cm long, the central lobe with a submarginal vein distally; spadix 14–32 cm long, not surrounded by spathe

Anthurium trilobum

27. Anthurium asplundii Croat, Ann. Miss. Bot. Gard. 78(3): 601, f. 8, 10, 35–37, 43 (1991). Plate 7: C. Holoepiphytic herbs, sometimes terrestrial. Stem to 60 cm long. Cataphylls lanceolate, straight or hooked to 14 cm long. Leaves suberect-spreading, forming a cup-like; blades coriaceous to subcoriaceous, oblanceolate to broadly oblanceolate-elliptic, (60–)90–150 x 20–40 cm, acute to broadly rounded or shallowly cordate at base, acute to abruptly and shortly-acuminate at apex, moderately undulate at margins, not punctuate beneath; petioles 7–15(–25) cm long. Inflorescences erect; peduncle (20–)33–60 cm long. Spathe reflexed, 9–30 x 1–2 cm, green-purplish. Spadix 12–23 cm long, purple to maroon. Infructescences pendulous, brown. COLOMBIA. N: (S). ECUADOR. E: Cornejo 1902 (GUAY, MO). Distribution: Southwestern Colombia to Ecuador (Peru?). Ecology: This species inhabits wet forests. It is one of the largest mangrove holoepiphytes in our area. Use: It is cultivated as an ornamental. IUCN Red list category: This species is persistent in secondary forests and clearings, and is categorized as of Least Concern (LC). Etymology: The generic epithet is formed by the Greek word anthos (flower) and oura (tail), which refers to the characteristic tail-like appearance of the inflorescences. The specific epithet was chosen to honor Erick Asplund (1888–1974), a Swedish botanist who apparently was the first to collect this species in 1955 (Croat, 1991). However, A. asplundii was previously documented from the REMACAM with a black and white photograph and cited as a “Giant epiphyte Anthurium” by Acosta-Solís (1944: 285). It is unknown if Acosta-Solís actually collected the species. Notes: A. asplundii can be recognized by its cup-like arrangement of large suberect-spreading leaves with

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

very short petioles, which mimic the rosulate large-leaved Bromeliads. This species has been reported in the REMACAM of northwestern Ecuador as Anthurium obtusum (Ecociencia, 2006: 50, fig. 9). 28. Anthurium cordobense Croat & Bay, Novon 16(1): 30 (2006). Epiphytic; stem short; internodes to 3 cm diam.; roots highly branching, forming a dense mass. Cataphylls to 8 cm long, persisting as weathered fibers. Leaves erect-spreading; blades moderately coriaceous, narrowly obovate, 26–47.5 x 7–15 cm, (2.7–)3.1–4.4 times longer than wide, (2.9–3.6)5.4–11 times longer than petiole, broadest beyond middle; short-acuminate to abruptly short-acuminate at apex, usually obtuse, sometimes attenuate at base, obscurely punctate above, minutely dark-punctate beneath; petioles 3.3–15(–19) cm long, D-shaped in cross section. Inflorescences pendulous; peduncle 23–40 cm, shorter than petiole. Spathe spreading to reflexed, 4.5–8 cm long, usually green, sometimes reddish. Spadix 9.5–21 cm long, golden-yellow to reddish-orange. Infructescences pendulous, to 22 cm long; berries orange. COLOMBIA. V: Killip 5232 (G, NY). N: (E). Distribution: Lowlands of western Colombia. Ecology: In rainforests. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the city of Córdoba, in the Department of Valle del Cauca, Colombia, the locality where the earliest collection of this species was gathered (Croat et al., 2006). 29. Anthurium friedrichsthalii Schott, Oesterr. Bot. Wochenbl. 5(9): 65 (1855). Syn: Anthurium linearifolium Engl.

Anthurium gracile var. friedrichsthalii (Schott) Engl. Holoepiphytic, usually pendulous, often loosely attached by only a few roots; stem usually less than 15 cm long, 1–1.5 cm diam. Cataphylls weathering to coarse linearfibers. Leaves pendulous; blades lorate, 12–56 x 0.8–4 cm, acuminate at apex, subacute at base; conspicuously punctate beneath; petioles 1.5–10(–14) cm long. Inflorescences pendulous; peduncle 19–30 cm long. Spathe lorate, 1–5 cm x 3–5 mm, pale green, soon deciduous. Spadix 3–15 cm x 4–5 mm, pale green, sometimes tinged purplish, turning brownish. Infructescences pendulous, to 25 x 2 cm; berries pale yellow-orange.

COLOMBIA. V: Killip 5224 (PH). Distribution: Guatemala to northwestern Ecuador. Ecology: In wet forests and rainforests.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

IUCN Red list category: Least Concern (LC). Etymology: The epithet honors Emanuel Ritter von Friedrichsthal (1809–1842), an Austrian collector of plants, daguerreotypist, and amateur archaeologist, who gathered the type species in Central America. 30. Anthurium guayaquilense Engl., Bot. Jahrb. Syst. 25(3): 373 (1898). Syn.: Anthurium gracillimum Engl. Holoepiphytic herbs. Stem short. Cataphylls coriaceous, 5.5–13 cm long, deciduous. Leaves suberect; blades coriaceous to subcoriaceous, narrowly obovate or oblanceolate, sometimes elliptic, 18.5–40 x 4.5–12 cm, cuneate at base, acuminate at apex, with numerous short white raphides (visible under 10 x magnification) above, not punctate beneath; petioles 6–20.5 cm long. Inflorescences erect to spreading, peduncle 15.5–35 cm long. Spathe spreading to subpendulous, 4–13 x 0.7–1.5 cm, green, often with maroon streaks. Spadix 7.5–18 x 0.3–0.5 cm, purple-violet. Infructescences subpendulous; berries orange to red-orange at maturity. COLOMBIA. CH: Fuchs et al. 21970 (U). V: (E). Distribution: Western Colombia to western Ecuador. Ecology: In wet forests and rainforests. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the city of Guayaquil, Ecuador. The type was collected in 1890 by Luigi Sodiro (1836–1909) in the region between Guayaquil and Balao. 31. Anthurium hacumense Engl., Bot. Jahrb. Syst. 25(3): 363 (1898). Holoepiphytic herbs. Stem short. Cataphylls coriaceous, ca. 9 cm long, weathering to coarse linear fibers. Leaves suberect-spreading, forming a cup-like; blades coriaceous, oblanceolate, 25–81 x 7.5–29 cm, attenuate at base, acuminate at apex, conspicuously punctate beneath; petioles 2–12 cm long. Inflorescences erect, peduncle 10–27 cm long. Spathe patent, 5.5–11 x 2.2–3.5 cm, yellow-green tinged with violet-purple, withering and early deciduous. Spadix 10–20 x 1–2.5 cm, purple-violet. Infructescences spreading to suberect, red; berries brightred at maturity. COLOMBIA. CH: Fuchs et al. 22033(COL, MO). Distribution: Costa Rica to western Colombia. Ecology: In wet forests and rainforests.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the Hacum River, San Jose Province, Costa Rica, where the type collection was gathered in 1892 by the Swiss botanist Adolphe Tonduz (1862–1921). 32. Anthurium incomptum Madison, Selbyana 2: 286 (1978). Plate 7: D. Holoepiphytic herbs. Stem to 50 cm long or more. Cataphylls brown, deciduous. Leaves arranged along distal portion of stem; blades coriaceous, ovate-elliptic to oblong-elliptic, 25– 55 x 8–18 cm, cuneate at base, obtuse to somewhat acute at apex, not punctate, the midvein reddish to red beneath; petioles to two-thirds as long as blade, 15–27 cm long, subterete. Inflorescences axillary from upper axils, solitary, erect; peduncle 20–50 cm long. Spathe patent, 7–12 x 1.5–2.5 cm. Spadix cylindric, 7–15 cm long, yellow. Infructescences not seen. COLOMBIA. V: Gentry & Juncosa 40704 (COL, MO). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8196 (GUAY). Distribution: Panama to western Ecuador. Ecology: In moist, wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means graceless. Note: Cornejo & Cangá 8196 differs from the general description of this species by the color of spadices (yellow vs. dark violet-purple). 33. Anthurium littorale Engl., Bot. Jahrb. Syst. 25(3): 405 (1898). Plate 8: A. Holoepiphytic herbs. Stem short, erect, cataphylls brown, nonfibrous, soon deciduous. Leaves arranged along stem; blades coriaceous, oblong, 15–35 x 5–12 cm, cuneate to obtuse at base, obtuse to retuse at apex, sparsely to moderately punctate and the midvein yellowish on both surfaces; petioles to two-thirds as long as blade, 5–20 cm long, semiterete. Inflorescences axillary from upper axils, solitary, erect; peduncle 10–25 cm long. Spathe pendulous, 5–10 x 0.7–1.2 cm. Spadix cylindric, 10–22 cm long, dark-purple. Infructescences pendulous, orange; berries orange at maturity. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8124 (GUAY). Distribution: Costa Rica to western Ecuador. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet means coast, referring to where the species was initially known to occur, along the Costa Rican Pacific coast.

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

34. Anthurium obtusum (Engl.) Grayum, Phytologia 82(1): 35 (1997). Plate 8: B. Syn.: Anthurium trinerve Miq. Anthurium trinerve var. obtusum Engl. Holoepiphytic climber herbs. Stem elongate, subpendulous, covered by fibrous remains of cataphylls. Leaves arranged along stem; blades coriaceous, elliptic to obovate, 7–20 x 3.5–8 cm, cuneate to obtuse at base, obtuse to acute at apex, densely punctate beneath; petioles to two-thirds as long as blade, 1–5 cm long, semiterete. Inflorescences axillary, solitary, erect; peduncle 2–8 cm long. Spathe (sub)erect, 2–4 x 0.4–1.3 cm, green. Spadix cylindric, 2–7 cm long, white. Infructescences (sub)erect, green; berries white at maturity. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6951 (AAU, GB, GUAY, MO). G: Cornejo 11 (GUAY). Distribution: Mexico to Bolivia. Ecology: In dry, moist, wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin Word that refers to the obtuse base and apex of the leaf blades (a variable feature). Note: At the present, “Anthurium obtusum is the best name to use for this species, but the leaves are much more ovate than the component from the Amazonian basin. This could be a new subspecies of A. obtusum or a distinct species” (Croat, pers. comm.). 35. Anthurium oxyanthum Croat & Bay, Novon 16(1): 42 (2006). Epiphytic; stem short; roots branched. Cataphylls 5–7 cm long, weathering to a reddish brown net-like reticulum, persisting somewhat intact. Leaves erect to erect-spreading; blades coriaceous, elliptic, 27–35(–38) x 8–15(–21) cm, (1.8–)2.2–3.5(–3.7) times longer than wide, 1–1.5(1.7) times longer than petiole, dark brown-punctate beneath; acuminate to obtuse apiculate at apex, acute to obtuse at base; petioles 15–28(–33) cm, sharply triangular in cross-section. Inflorescences erect; peduncle 22–43 cm long, usually longer than petiole. Spathe stiffly spreading, 5–8 cm long, coriaceous, medium green, sometimes slightly reddish, inrolled. Spadix 22–30 cm x 8–11 mm, green, rarely yellow-green to cream. Infructescences 22–24 cm long, usually maroon, rarely reddish-orange; berries bright red, usually sparsely scattered in basal 1/3 of spadix. COLOMBIA. V: Gentry & Juncosa 40703 (COL, JAUM, MO). Distribution: Lowlands of western Colombia. Ecology: In rainforests. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat it is herein categorized as Near Threatened (NT).

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

Etymology: The epithet is formed by the Greek prefix oxy (pointed), and the Greek word anthos (flower), referring to prickly appearance of the sharply exserted pistil, tepals, and anthers (Croat et al., 2006). 36. Anthurium paludosum Engl., Bot. Jarhb. Syst. 25(3): 401 (1898). Epiphytic; stems to 30 cm long. Cataphylls coriaceous, 5–6 cm long, persisting ± intact. Leaves spreading-pendulous; blades subcoriaceous, narrowly elliptic to oblanceolate, 10– 38 x 3–13.5 cm, short-acuminate at apex, acute to attenuate (rarely obtuse to subrounded) at base; sparsely punctate above and beneath; petioles 3–14 cm long. Inflorescences erectspreading, shorter than leaves; peduncle 7.5–15.5 cm. Spathe inserted at ca. 60° angle on peduncle, oblong-lanceolate, 5.5–7 x 0.4–1.4 cm, whitish to green or purplish, withering soon after anthesis. Spadix 8–14.5 cm x 6–7 mm diam. (at middle, scarcely tapered at apex), green, tinged orange, becoming violet-purple. Infructescences 6–25 cm long; berries orange, red-orange, or white tinged with orange. COLOMBIA. CH: (E). V: Killip 5225 (GH, MO, NY, PH). Distribution: Costa Rica to northwestern Ecuador. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that refers to the swampy habitat of the species, found in mangroves near Buenaventura, Department of Valle del Cauca, Colombia, where the type was collected in 1884 by the German botanist Karl Friedrich Lehmann (1850–1903). 37. Anthurium trilobum André, Ill. Hort. 24: 108, t. 283 (1877). Syn.: Anthurium insigne Mast. Anthurium tripartitum Engl. Anthurium stenoglossum Sodiro, Anthurium garagaranum Standl. Anthurium cineraceum Croat Vernacular names: corrientoso, deshinchador, guaca de mata, guaco, mano, tres dedos. Epiphytic, stem appressed-climbing; internodes up to 5 cm long. Cataphylls reddish-brown, [5–]8–14(–19) cm long, decomposing into fibers, persistent. Leaves erect–spreading; blades coriaceous to subcoriaceous, deeply trilobed, [21–]40–83 x [15–]30–65 cm, upper surface densely covered with raphides, lower surface covered with red blotches, base hastate; central lobe oblong-elliptic to narrowly obovate; collective veins 3–7 mm from margin; lateral lobes falcate, 50–60˚[–70˚] from central lobe, [10–]30–62 x [3.5–]8–21 cm, narrowly rounded at apex; petioles [26–]47–89 cm x [3–]6–12 mm. Inflorescences erect-spreading; peduncle 15–40(–47) cm x 3–7 mm, shorter than petiole. Spathe

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

lanceolate, 12–30 cm long, medium green. Spadix 14–32 cm x 5–12 mm, bright yellow to orange-yellow. Infructescences up to 36 x 1.7 cm; berries purple at apex, white at base. COLOMBIA. CH: Fuchs et al. 21963 (COL, MO). ECUADOR. E: (E). Distribution: Southeastern Panama to western Ecuador. Ecology: In wet forests and rainforests. Use: Deshinchador (Spanish), one of its vernacular names, suggests that this species might be used to reduce swelling. However, no such uses are recorded for this species. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Latin words tri (three) and lobum (lobes), referring to the trilobed shape of the leaf blades. 38. Anthurium verrucosum Croat & Bay, Novon 16(1): 46 (2006). Epiphytic; stem short. Cataphylls 4.5–15 cm long, persisting as weathered fibers, often in a net-like reticulum with apical epidermis intact. Leaves erect to erect-spreading; blades coriaceous, oblong-elliptic to elliptic, 24–57 x 5–19 cm, (2.7–)3.3–4.5(–5.4) times longer than wide, 1–2.4(–2.9) times longer than petiole, broadest near middle, profusely verrucose (with 10 x magnification) and sometimes with sparse pustules above, minutely and sparsely dark-punctate, sometimes with clustered pustules beneath, long acuminate at apex, usually obtuse or sometimes cuneate at base; petioles 9–38 cm x 3–6 mm. Inflorescences erect to spreading; peduncle 19–54 cm x 3–6 mm, longer than petiole. Spathe spreading to reflexed, lanceolate, 5–15 cm long, glossy, green, occasionally tinged red or yellow. Spadix 9.2–26.5 cm x 3–7 mm, weakly glossy to matte, gray, grayish white, gray-green, yellow, creamy yellow, yellowish green, or tan. Infructescences erect, 22–30.5 cm long; berries red to red-orange. COLOMBIA. V: Killip 3498 (US). Distribution: Lowlands of western Colombia. Ecology: In rainforests. Etymology: The epithet is a Latin word that refers to the presence of inconspicuous warts above the leaf blades. IUCN Red list category: Least Concern (LC). 39. Monstera pinnatipartita Schott, Oesterr. Bot. Wochenbl. 7(25): 197–198 (1857). Hemiepiphytic climber herbs. Stem 1–2 m long, appressed. Leaves arranged along distal stem; juvenile leaves not shingle-forming; mature blades deeply pinnatifid, imperforate or rarely with a few perforations, subcoriaceous to (rarely) coriaceous, ovate to elliptical, 25–50

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

x 20–35 cm, broadly cuneate to rounded or truncate at base; lateral veins ca. 6–19 per side; petioles 20–50 cm long, smooth, sheathed to within ca. 3–7 cm of geniculum, the sheath ± involute, persistent. Inflorescences erect; peduncle 9–23(–30) cm. Spathe pure white within. Spadix uniform, 5.5–11(–15) x 1.1–1.7(–2.5) cm, creamish. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 6980 (GUAY, MO). Distribution: Costa Rica to Bolivia. Ecology: In moist, wet forests, and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that refers to the deeply pinnatifid aspect of the leaf blades. 40. Philodendron chrysocarpum Croat & Bay, Novon 18(4): 432 (2008). Usually hemiepiphytic, occasionally terrestrial; stem appressed-climbing or scandent. Cataphylls 14–15.5 cm, sharply 2-ribbed, long-tapered and acute at apex, medium green, tinged purple, drying reddish brown, deciduous while intact. Leaves erect to spreading; blades subcoriaceous, elliptic, 23.5–30 x 5–9 cm, 3.4–4.8(–5.2) times longer than wide, 2.6– 3.8 times longer than petiole, acuminate at apex, attenuate at base; petioles (5.3–)6.5–9.5 cm long, sheathed in lower 1/3–2/3. Inflorescences erect, 1 per axil; peduncle 2–4 cm x 5–7 mm, shorter than petiole. Spathe long-tapered, 12–19 cm long, outside dark yellow-green, striate toward base, inside greenish-white with orange resin canals to within 2 cm of apex, only slightly constricted where the pistillate portion of the spadix ends; spathe blade 6–9 cm long; spathe tube 6.5–10 cm long. Spadix cylindrical, 8–15.5 cm long. Infructescences with pistillate spadix ca. 9 x 2.5–3 cm; berries bright golden yellow-orange. COLOMBIA. V: Killip & Cuatrecasas 39215 (US). Distribution: Lowlands of western Colombia. Ecology: In rainforests. IUCN Red list category: Least Concern (LC) (Croat et al., 2008). Etymology: The generic epithet is formed by the Greek words philo (love) and dendron (tree), referring to the usually epiphytic habit of the species in this genus. The specific epithet is formed by the Greek prefix chryso (gold) and the Greek word carpo (fruit), referring to the bright golden yellow-orange color of the berries (Croat et al. 2008).

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

41. Philodendron musifolium Engl., Bot. Jahrb. Syst. 37: 126 (1905). Plate 8: C. Hemiepiphytic climber herbs. Stem appressed, the internodes dark green, semiglossy, 4–6 x 1.7–2.5 cm. Cataphylls sharply 2-ribbed, pinkish to purplish-red, deciduous. Leaves arranged along distal stem; blade (sub)coriaceous, oblong, 35–50 x 8–12 cm, semiglossy, cordate at base, secondary veins numerous; petioles terete, 6–10 cm long. Inflorescences axillary, erect; peduncle 2–5 cm long, shorter than petiole. Spathe covering 2/3 of spadix, greenish to yellowgreenish, red at dorsal side of lower half. Spadix 16–19 cm long, reddish-cream. Staminate flowers arranged at upper half of spadix. Pistillate flowers arranged at lower half of spadix. COLOMBIA. N: (S). E: Cornejo & Bonifaz 1901 (AAU, GUAY, MO). Distribution: Southwestern Colombia to western Ecuador. Ecology: In wet forests. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat it is herein categorized as Near Threatened (NT). Etymology: The epithet is formed by the genus Musa (Musaceae, the banana family) and the Latin word folium (leaf blade), referring to the resemblance of the leaf blades to those of Musa. 42. Philodendron sparreorum Croat, Aroideana 27: 53 (2004). Plate 8: D. Hemiepiphytic climber herbs. Stem 1.5–3 m long, appressed. Roots elongated to several meters long. Cataphylls unribbed, 25–28 cm long. Leaves arranged along distal stem; blades subcoriaceous, ovate-cordate to ovate-triangular, 40–67 x 25–57 cm; prominently lobed at base, abruptly acuminate at apex; petioles subterete, 45– 100 cm long, sheated 10–23 cm long. Inflorescences 2–7 per axil, erect; peduncles 8–21 cm long. Spathe 7.5–10 cm long, orange, covering the spadix. Spadix 6–9.5 cm long. Staminate flowers arranged at upper half of spadix, creamish-brown. Pistillate flowers arranged at basal half of spadix. Berries pale green to orange at maturity. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6943 (AAU, GUAY, MO). Distribution: Western Colombia and western Ecuador. Ecology: In wet forests and rainforests. Uses: This species is occasionally cultivated as an ornamental. The unbranched, elongated fibrous roots produced from the stem, known in the REMACAM as Chandé, are locally used as ropes. Caution: The sap of this species causes permanent blindness and severe dermatitis (the skin feels like it’s burning on contact). Its chemical compounds are unknown.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat it is herein categorized as Near Threatened (NT). Etymology: Philodendron sparreorum was named in honor of Baron Benkt Sparre (1918–1986), a Swedish botanist and taxonomist, and the first co-editor of the Flora of Ecuador Project, who collected many interesting new species in Ecuador. 43. Philodendron tripartitum (Jacq.) Schott, Wiener Z. Kunst 3: 780 (1829). Bas.: Syn.:

Arum tripartitum Jacq. Caladium tripartitum (Jacq.) Willd. Philodendron tripartitum var. tricuspidatum Engl. Philodendron fenzlii Engl.

Vernacular name: guaral. Hemiepiphytic climber herbs. Stems 1–2 m long. Roots elongated. Cataphylls elongatelanceolate, caducous. Leaves arranged along distal stem; blades subcoriaceous, 3-parted, the segments elliptic-lanceolate, ca. 15–25 x 4–7 cm, acuminate at apex, the lateral ones conspicuously oblique; petioles terete, 20–50 x 1–1.5 cm. Inflorescences 2–5 per axil, erect; peduncles 3–8 cm long. Spathe 8–12 cm long, covering spadix, greenish. Spadix 6–11 cm long. Staminate flowers arranged at upper half of spadix, creamishwhite. Pistillate flowers arranged at lower half of spadix. Berries red at maturity. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6952 (AAU, GUAY, MO). Distribution: Mexico to Ecuador. Ecology: In wet forests and rainforests. Use: The latex and ashes from the leaves and stems are applied to cure a skin condition called “guaral”. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Latin words tri (three) and partitum (parted), referring to the 3-parted leaf blades of this species. 44. Stenospermation angustifolium Hemsl., Biol. Cent. -Amer., Bot. 3(18): 425 (1885). Epiphytic vine to at least 30 m long; stems 4–8 mm diam. Leaf blades lanceolate to narrowly elliptic, 4–20 x 1–7 cm, cuneate to rounded and often unequal at base, sharply acuminate at apex, the lower leaves deciduous; petioles 1.5–10 cm long, the sheath thick, extending almost to blade, the upper edge free. Peduncles 6–10 cm long, frequently recurved near apex before anthesis. Spathe 4–6 cm long, acuminate, white, soon deciduous. Spadix 2–5 cm x 3–5 mm diam., white to grayish or greenish, the stipe obsolete or to 3.3 mm long. Infructescences suberect; berries white at maturity.

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COLOMBIA. CH: (E). V: Gentry & Juncosa 40722 (B, COL, MO). Distribution: Nicaragua to southwestern Ecuador. Ecology: In wet forests and rainforests. Uses: Cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Latin words angusti (narrow) and folium (leaf blade), referring to the narrow leaf blades. Note: The species is distinguished by its small size, slender elongate stems, and small subsessile inflorescences. 45. Stenospermation robustum Engl., Bot. Jahrb. Syst. 37(1): 113 (1915). Epiphytic vine to at least 20 meters long. Leaf blades narrowly or ± broadly elliptical to lanceolate-oblong or oblanceolate, 15–30 × 5–10 cm, cuneate at base, acuminate at apex, the lower leaves deciduous; petioles 7.5–25 cm long, sheathed 2/5–9/10 to leaf-base. Peduncles 24–58 cm long. Spathe cream-white to yellow. Spadix 7–13.5 × 0.6–1 cm, white to yellowish orange or brownish, stipitate by 0.5–1.5 cm long. Infructescences suberect; berries pale yellow, orange at summit at maturity. COLOMBIA. V: Gentry & Juncosa 40721 (COL, MO). Distribution: Costa Rica to western Colombia. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that refers to the stout appearance of this species. ARECACEAE (Palm Family) Unbranched trees, shrubs, lianas or herbs. Leaves alternate, basal or clustered at apex of stem, petiolate; blades usually pinnately or palmately compound; exstipulate. Inflorescences of spicate or branched interfoliar or intrafoliar spadices. Flowers bisexual or unisexual (monoecious or dioecious), actinomorphic, often immersed in a cavity, subtended by 2 envelopes. Sepals 3, imbricate; petals 3 or tepals 6. Stamens usually 6, free, in two whorls of 3 each. Ovary superior. Fruit drupes; seeds 1.

The family is represented by only two species, one of which, the introduced coconut palm Cocos

nucifera is included in this work because has been planted in mangrove areas. Key to the species 1. Solitary palms, stem 20–40 cm dbh; fruits 15–25 cm diam., green, yellow, or brown Cocos nucifera 1. Caespitose palms, stems 7–18 cm dbh; fruits 1–2 cm diam., purple-black

Euterpe oleracea

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46. Cocos nucifera L., Sp. Pl. 2: 1188 (1753). Plate 9: A. Vernacular names: cocotero, palma, palma de coco. Solitary palms to 15 m tall and 40 cm dbh. Leaves 2.5–5 m long, pinnae to 100 on each side, regularly inserted in one plane, straight, the central ones to 1 m long. Inflorescences erect, one branched racemes, 0.5–1 m long, the basal part of each inflorescence bearing pistillate flowers; the distal part bearing staminate flowers. Fruits globose to subtriangular, green, yellow, or brown, 15–25 cm diam., with persistent, accrescent calyx; mesocarp thick, fibrous. COLOMBIA. N: (S). ECUADOR. E: (S). Distribution: Widespread along tropical shores. Ecology: Cocos nucifera is an introduced species in lowland areas (Jørgensen and León, 1999). Since 1940, some mangroves in western Colombia and northwestern Ecuador have been felled and planted with “pipa” trees. In some places the coconut palm remains mixed with the mangrove vegetation. It inhabits dry, moist, wet forests, and rainforests. Biological interactions: The flowers are visited by honey bees, and the fruit pulp is palatable to several species of shrimp. Dispersal: The fruits are water-dispersed. Uses: The immature fruit has a liquid endosperm commonly known as “agua de coco”, “agua de pipa” in Spanish, a refreshing drink with diuretic properties. There are several varieties, which can be distinguished in the field by differences in the taste of the liquid endosperm, which turns to be fleshy with age. The fleshy endosperm is used by the African descendents to cook a diverse number of delicious dishes named “encocados” and tasty desserts as the “cocadas”. The seed testa, known as “concha de coco”, is burned when dry to produce a strong flame. It is considered an excellent firewood as it cooks food faster. It is also used to manufacture handycrafts, earrings, and purses. The thick, fibrous mesocarp, called “estopa”, produces a dense smoke when burned, which is used to smoke fish and as a mosquito repelent. This “estopa” is also very useful for caulking a leaking canoe, boat or ship; the grooves are filled with the fibers and covered with a dense wax-like substance called “brea de abeja”, produced by a bee species “that lives in the forest inland and sucks the trunk-latex, not the flowers” (Cangá, field obs.). These uses of “estopa” explain the fact that the coconut palm was abundantly cultivated in Guayaquil, the main ship factory in the region during the colonial period, as it is recorded in a painting by José Cardero, the artist of the Alejandro Malaspina expedition that visited the city in October of 1790. IUCN Red list category: Least Concern (LC).

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47. Euterpe oleracea Mart., Hist. Nat. Palm. 2(2): 29, fig. 28–30 (1824). Plate 9: B. Syn.: Euterpe badiocarpa Barb. Rodr. Euterpe beardii L. H. Bailey Euterpe cuatrecasana Dugand Vernacular names: bambil, murrapo, naidí, palma naidí, palmicha, palmiche. Caespitose palms, up to 25 stems per clump, the stems 3–20 m tall, 7–18 cm dbh. Basal adventitious roots present, to ca. 70 cm tall. Leaves 1.5–2.5 m long, arching, pinnae 40–80 on each side, regularly inserted in one plane, subpendulous, the central ones 0.6–1 m long. Inflorescences erect, one branched racemes, ca. 0.5 m long. Fruits globose or depressed-globose, purple-black, 1–2 cm diam., the stigmatic remains lateral; mesocarp with radial sclereids. COLOMBIA. CH: Fuchs et al. 22053 (COL). V: Bernal et al. 1776 (COL). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6983 (AAU, GUAY). Distribution: Panama to Brazil. Ecology: Euterpe oleracea inhabits poorly aired and periodically flooded soils of mangroves, swamps, and along adjacent water courses, in the wet forest and rainforest life zones. It can be an aggressive colonizer of disturbed swampy areas (Henderson and Galeano, 1996), where it occasionally forms pure stands called “naidizales”. Biological interactions and dispersal: The fruits are eaten by birds, mammals and fish (Moegenburg, 2002). Commercial fish such as Sábalo (probably a Brycon sp., Characidae) feed on the fruits (Pedersen 104001, AAU), and the seeds are stomatochorously dispersed by bats (Lobova et al., 2009). Uses: The edible raw palm heart is canned for exportation. The fruits are edible as well (Henderson and Galeano, 1996); the fleshy mesocarp is mixed with water and made into a stimulant beverage called “jugo de Palmicha” (Palmicha juice), which is sold in western Colombia and northwestern Ecuador (e.g. Barfod et al. 60185, AAU, NY, QCA). The fruits are also locally used as an ice cream flavoring (Pedersen 104001, AAU). The fruits and discarded seeds are used as feed for domestic animals (Henderson and Galeano, 1996). The stems are used for building walls, fences, and columns to raise TV antennas, and the leaves are used to build roofs (Cornejo & Bonifaz 6983, AAU, GUAY; Bernal & Galeano 670, AAU). The young leaves are mashed and the sappy matter applied to stop bleeding; they can be taken orally to stop internal hemorrhaging. This is a graceful palm tree species that may be planted as an ornamental (Henderson and Galeano, 1996). IUCN Red list category: Least Concern (LC). Etymology: The genus is named after the muse Euterpe of Greek mythology. The specific epithet may refer to an unknown similarity of the palm heart to cabbage (Brassica oleracea, Brassicaceae).

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ASTERACEAE (Aster Family) Herbs, shrubs, trees, vines or sometimes lianas. Leaves opposite to alternate or whorled, simple to compound or dissected; exstipulate. Flowers sessile in close head on common receptacle, flowering in centripetal sequence within head, sometimes individually subtended by a small bract, and nearly always collectively surrounded by an involucre of few to many bracts. Heads arranged in various sorts of secondary inflorescences. Individual flowers epigynous, perfect or unisexual, gamopetalous, commonly 4(–5)-merous, without definite calyx. Stamens as many as corolla lobes and alternate with them, epipetalous. Ovary inferior. Fruit an achene.

In our area, this family is represented by Tuberostylis, a Chocoan coastal genus that comprises two

salt-tolerant species, both inhabiting mangroves, swamps and the terra firme forests nearby. The absence of pappus in the seeds of this genus, an uncommon feature for the family, is a particular adaptation that may suggest dispersal via birds or even by water. Field observations are needed to verify this hypothesis. Key to the species 1. Leaf blades ovate to ovate-elliptic or shortly lanceolate; inflorescences axillary, subsessile Tuberostylis axillaris 1. Leaf blades spatulate-obovate or obovate to rhomboidal; inflorescences terminal on lateral branches, on erect primary peduncles, the primary peduncles 2–10 cm long Tuberostylis rhizophorae

48. Tuberostylis axillaris S. F. Blake, J. Wash. Acad. Sci. 33: 265 (1943). Suffrutescent shrub, liana or treelet, often climbing on vegetation, sometimes over water. Leaves opposite; blades coriaceous, ovate to ovate-elliptic or shortly lanceolate, 4–8 x 2–4 cm, cuneate at base, usually acuminate at apex, entire to slightly undulate at margins, glabrous, 3-nerved from base; petioles ca. 1 cm long. Inflorescences axillary, the primary peduncle 1–3 mm long; heads sessile in clusters of 3 to 4, 7–9 mm long, with ca. 4 florets; involucral bracts green. Corolla cylindric without distinguishable tube, ca. 4.2 mm long, white. Achenes narrowly-oblong, blunty 3–5-angled or ribbed, 2.5–3 mm long; pappus lacking. COLOMBIA. CH: Killip 35515 (COL). V: Gentry & Juncosa 40724 (COL, MO). Distribution: Coastal Colombia. Ecology: This is a fringe species, commonly found near and at the edge of mangroves. It occurs in the wet forest and rainforest life zones. Etymology: The epithet is a Latin word, referring to the distinctive axillary subsessile inflorescences.

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IUCN Red list category: Due to factors such as a narrow habitat range of less than 2,000 km2, a loss of habitat, and a subwoody habit, the undercollected species Tuberostylis axillaris is herein categorized as Vulnerable (VU B2abiii). 49. Tuberostylis rhizophorae Steetz in Seem., Bot. Voy. Herald 142, t. 29 (1854). Plate 9: C. Hemiepiphytic or holoepiphytic vine, much branched. Stems slender, trailing, glabrous with adventitious roots along. Leaves opposite; blades slightly succulent, spatulate-obovate or obovate to rhomboidal, to 2.5 x 1.8 cm, cuneate and abruptly narrowly-decurrent at base, rounded to obtuse at apex, slightly crenate in distal half or two-thirds, glabrous, obscurely 3-nerved near base; petioles to 2.5 cm long. Inflorescences terminal on lateral branches, slightly corymbose on erect primary peduncles, the primary peduncles 2–10 cm long; heads sessile or nearly sessile in clusters of 2 to 8, 8–9 x 3 mm, with 10 to 12 florets; involucral bracts imbricate, greenish. Corolla tubular, ca. 3.5 mm long, white. Achenes prismatic to cylindrical, ca. 3 mm long; pappus lacking. COLOMBIA. CH: Fuchs et al. 21842 (COL, MO). V: Killip 5335 (NY). C: (E). N: Romero-Castañeda 5121 (COL). ECUADOR. E: Cornejo & Bonifaz 7256 (GUAY, US). Distribution: Eastern Panama to northwestern Ecuador. Ecology: The species occurs mostly in mangroves, though also in adjacent coastal wet forests and rainforests (King and Robinson, 1975). IUCN Red list category: Though common locally, the species is rather poorly represented in herbaria. Due to factors such as a narrow range of distribution of less than 2,000 km2 and a loss of habitat, Tuberostylis rhizophorae is herein categorized as Vulnerable (VU B2abiii). Etymology: The species was first discovered growing on the roots of Rhizophora trees (hence the epithet) by Berthold Carl Seemann (1825–1872) on the voyage of H.M.S. Herald (Blake, 1943). The original publication cited it from the mangroves of southern Darién; however, the type was most likely collected in present northwestern Colombia. BATACEAE (Saltwort Family) Glabrous, succulent, low, herbs or shrubs. Leaves opposite, entire. Flowers bracteate, solitary or in axillary strobili. Staminate flowers with cup-shaped perianth. Stamens 4, alternating with 4 staminodia, the anthers 2-celled, introrse. Pistillate flowers without perianths, each consisting of a single pistil. Ovary 4-loculed, with 1 ovule in each locule; stigma 2-lobed, sessile. Seeds without endosperm.

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50. Batis maritima L., Syst. Nat. (ed. 10) 2: 1289 (1759). Plate 9: D. Vernacular names: romero, vidrillo, vidrio. Terrestrial herbs to subshrubs, usually much branched, to 1 m tall. Stems erect, the older prostrate, cylindric, to 6 mm diam. Branches often opposite. Leaves sessile; blades succulent, frequently trigonous, 10–30 x 4–8 mm, apiculate at apex, glabrous. Inflorescences staminate and pistillate, axillary, on sessile to subsessile spikes, 5–10 x 3–5 mm; floral bracts green, ca. 2 mm long, in 4 rows, imbricate, persistent; stigma persistent. Fruits ± ovoid, irregular, 5–15 x 5 mm. ECUADOR. M: Cornejo 1718 (GUAY). G: Cornejo & Castro 282 (GUAY, QCNE). O: Holm-Nielsen 22822 (AAU). PERU. T: (S). P: (E). Distribution: Southern United States to Peru and Brazil, also on the Galapagos Islands. Ecology: Batis maritima is a widespread halophyte (Tomlinson, 1986). In our area, it occurs in open sunny places, just above the tidal level of disturbed mangroves. Apparently it is restricted to the life zones of dry forest to dry thorn scrub in Ecuador and adjacent northwestern Peru. In mangroves, Batis maritima occurs most frequently as a fringe element, but it is also found surrounding shrimp ponds, and occasionally on beaches. Biological interactions: The plants are eaten by donkeys (Madsen et al., 2001). Uses: Batis maritima contains high concentrations of bromine and iodine. The leaves, when chewed, have a salty taste; they are eaten raw in salads to purify the blood, and as a treatment for rheumatism and “gota”. An infusion of branches and leaves is consumed for several months to manage eczema and psoriasis (Bernal and Correa, 1989). This species is cultivated as an ornamental in the Galapagos islands. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word, referring to the coastal distribution of this species. BIGNONIACEAE (Trumpet-creeper Family) Woody trees, shrubs or vines, rarely herbs. Leaves usually opposite, palmately or pinnately compound, sometimes simple, the terminal leaflet often replaced by a tendril. Glandular fields often present between the petioles. Inflorescence terminal or axillary, a panicle or raceme. Calyx more or less copular. Corolla gamopetalous with a conspicuous tube and 5 usually imbricate lobes. Stamens inserted on corolla tube, didynamous, usually 4, a posterior staminode usually present. Ovary superior, the ovules numerous; style slender; stigma bilamellate. Fruit a silique with dehiscence perpendicular or parallel to septum or a berry or amphisarcum; seeds usually winged, few times with reduced wings and corky.

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Key to the species 1. Shrubs or trees; leaves simple or frequently 3-foliolate; tendrils absent 2. Leaves frequently 3-foliolate, lepidote beneath; fruit oblong-cylindric, dehiscent Tabebuia palustris 2. Leaves simple, glabrous beneath; fruit globose, indehiscent Amphitecna latifolia 1. Lianas; leaves 2-foliolate; tendrils present 3. Tendrils simple; calyx spathaceous; corolla magenta without; seeds thick, wingless Phryganocydia phellosperma 3. Tendrils trifid; calyx cupular; corolla bright yellow; seeds thin, surrounded by a membranaceous wing Anemopaegma chrysanthum

51. Amphitecna latifolia (Mill.) A. H. Gentry, Taxon 25: 108 (1976). Plate 10: A. Bas.: Syn.:

Crescentia latifolia Mill. Crescentia cucurbitina L. Dendrosicus saxatilis Raf. Crescentia obovata Benth. Crescentia palustris Forsyth ex Seem. Enallagma latifolia (Mill.) Small Amphitecna obovata (Benth.) L. O. Williams Dendrosicus latifolius (Mill.) A. H. Gentry

Vernacular names: calabacillo, totumillo. Small scraggly trees to 10 m tall and 20 cm dbh. Branches usually crooked, the twigs terete or 3- or 4-angled with decurrent ridges below nodes. Leaves simple, alternate to opposite; blade coriaceous, obovate, 7–19 x 4–10 cm, cuneate-attenuate at base, rounded to almost acute and usually apiculate at apex, glabrous beneath, the lateral veins 6–15 per side; petioles poorly defined, 0.2–1 cm long; ± winged. Inflorescence a solitary flower or fascicles of 2–3 flowers borne on larger twigs; pedicels 2–3.5 cm long. Calyx bilabiate, 2.5–3.7 x 1–2.5 cm, dark green, essentially glabrous. Corolla tubular, 3.8–6.2 x 1.2–2.4 cm (at mouth), greenish-white. Fruit amphisarcum, subglobose, 6–9 cm wide, greenish-yellow, glossy, glabrous; seeds thick, corky, 1.3–1.6 x 1.6–1.8 cm, without wings. COLOMBIA. CH: Gentry & Juncosa 40932 (MO). V: Killip 33012 (A, COL, MO). C: Cabrera et al. 4358 (CUVC). N: (S). ECUADOR. E: Little 6339 (F, K, US). M: Neill et al. 11409 (MO, NY). Distribution: Mexico to northwestern Ecuador and Venezuela, and in the West Indies. Ecology: A common tree species restricted to seashores, mangrove fringes, and brackish water swamps along both coasts of America (Gentry, 1973, 1980, 1982). It occurs in moist, wet forest, and rainforest life zones.

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Biological interactions: The flowers are pollinated by bats (Fleming et al., 2009). Dispersal: This is a water-dispersed species. Fruits are found washed up on beaches with still fertile seeds, which germinate only when the fruit is opened (Gentry, 1973, 1980, 1982). Uses: The soft pulp of the fruits has been reported to be edible. The fruit has also been reported to be medicinal. The seeds are used as a purgative, and the pulp as a laxative and remedy for chest infections (Gentry, 1973, 1980). IUCN Red list category: Due to factors such as a widespread range of distribution and a common presence of healthy populations, this species is categorized as of Least Concern (LC). Etymology: The epithet is formed by the Latin words lati (broad) and folia (leaves), referring to the relatively wide leaf blades. 52. Anemopaegma chrysanthum Dugand, Caldasia 4: 307 (1947). Plate 10: B. Vernacular names: bejuco gallinero, tortuguero. Woody lianas. Stem glabrous. Tendrils trifid. Leaves 2-foliolate, opposite; leaflets coriaceous to subcoriaceous, ovate to ovate-elliptic, 9–20 x 4–13 cm, cuneate to rounded at base, acute to shortly acuminate at apex, entire, glabrous, the lateral veins 4–7 per side, minute-lepidote beneath; petioles 3–4 cm long, the petiolules 0.8–3.7 cm long. Inflorescence an axillary raceme, 4–8-flowered; pedicels glandular lepidote. Calyx cupular, 5–9 mm long, light green. Corolla tubular-campanulate, zygomorphic, 5–7

long,

uniformly

bright-yellow.

Capsule

oblong-elliptic,

only

slightly

compressed, 14–20 x 7–8.5 cm; seeds thin, surrounded by a membranaceous wing. COLOMBIA. CH: Juncosa 1657 (MO). V: Gentry & Juncosa 40714 (MO). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz s.n. (GUAY). Distribution: Southern Mexico to Peru. Ecology: In dry, moist, wet forests, and rainforests. Dispersal: The seeds are wind dispersed. Use: In rural areas, the elongate fibrous stems are used to tie wooden chicken cages together, hence “bejuco gallinero”, one of the vernacular names. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Greek prefix chryso (gold) and the Greek word anthos (flower), referring to the bright-yellow corollas. Notes: Among lianas in mangroves, the species can be recognized by the 2-foliolate opposite leaves, the lack of foliaceous pseudostipules, the trifid tendrils, and the bright-yellow corollas.

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53. Phryganocydia phellosperma (Hemsl.) Sandw., Bull. Misc. Inform. Kew 1940(7): 302–303 (1941). Plate 10: C. Bas.: Macfadyena phellosperma Hemsl. Woody lianas. Stem glabrous. Tendrils simple. Leaves 2-foliolate, opposite; leaflets coriaceous to subcoriaceous, ovate, 7–13 x 4–9 cm, truncate at base, acuminate at apex, entire, glabrous, the lateral veins 4–6 per side, brochidrodomous; petioles 0.8–4 cm long, the petiolules 0.8– 2.2 cm long. Inflorescence 1–2 terminal flowers; pedicels 2–3.5 cm long. Calyx spathaceous, erect, 2–3 cm long, greenish-white. Corolla tubular-infundibuliform, zygomorphic, 5–10 cm long, magenta without, the throat white, finely reticulate, inconspicuously glandularpunctate. Capsule broadly-ovoid to almost suborbicular, only slightly compressed, 3.5–6.5 x 4–6 cm; seeds thick, corky, wingless. COLOMBIA. CH: Gentry & Juncosa 40935 (COL, MO). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6953 (AAU, GB, GUAY, QCA). Distribution: Endemic to the mangroves from Costa Rica to northwestern Ecuador. Ecology: In wet forests and rainforests. Dispersal: Its cork-like seeds are adapted for water-dispersal (Gentry, 1973). IUCN Red list category: Although the species is sometimes locally common, it is rather poorly represented in herbaria. Due to factors such as a narrow range of distribution of less than 2,000 km2 and a loss of habitat, Phryganocydia phellosperma is herein categorized as Vulnerable (VUB2abiii). Etymology: The epithet is formed by the Greek word phello (cork) and the Latin word sperma (seeds), referring to the corky seeds. Notes: Among lianas in mangroves, Phryganocydia phellosperma is easily recognizable by its 2-foliolate opposite leaves and simple tendrils. Moreover, its spathaceous calyx is highly distinctive. The cited collection from Ecuador represents the first record of this species for the country (Cornejo, 2003). 54. Tabebuia palustris Hemsl., Biol. Centr. -Amer., Bot. 2(12): 495 (1882). Plate 10: D. Vernacular name: barbasco. Small trees or shrubs to 4 m tall and 5 cm dbh. Aerial roots to 10 cm (or more?) tall occasionally present. Bark smooth, pale gray. Leaves frequently 3-foliolate; leaflets subcoriaceous, narrowly-elliptic or oblong, cuneate to subsessile at base, acute at apex, densely lepidote beneath, the terminal leaflet 10–19 x 2.1–5.6 cm, the lateral leaflets (when present) 5–19 x 1.1–5.1 cm; petioles 2.3–10 cm long, lepidote, the petiolules 1–3.5 cm long, the lateral petiolules (when present) 0.2–1.5 cm long. Calyx irregular bilabiate or 3-labiate, 1–2 x 0.4–1 cm, green, lepidote, persistent. Corolla tubular-infundibuliform, 5–7 cm long, white with yellow throat ridges. Stamens didynamous, the filaments 1.4–2.6 cm long. Ovary linearconical, 4.5–5 x 1 mm. Capsule oblong-cylindric, 8–11 x 1.6–2.6 cm, attenuate at both ends, green, densely lepidote; seeds suborbicular, thick, corky, 1.4–1.8 x 1.8–2.2 cm, brown.

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COLOMBIA. CH: Gentry & Juncosa 40720 (MO). V: Barbosa & Bonifaz s.n. (COL, MO). C: Cabrera 3864 (CUCV). N: (S). ECUADOR. E: Cornejo & Bonifaz 7096 (GUAY, QCNE). Distribution: Pacific coast from Costa Rica to northwestern Ecuador. Ecology: Tabebuia palustris is restricted to coastal mangrove swamps, along river mouths near the fresh/ salt water interchange (Gentry, 1973). It occurs in the wet forest and rainforest life zones. Dispersal: The cork-like seeds are water-dispersed. IUCN Red list category: The species has been categorized as Vulnerable (VU) (Polidoro et al., 2010). However, due to factors such as a narrow range of distribution of less than 2,000 km2 and a loss of habitat, Tabebuia palustris is herein categorized as Vulnerable (VU B2abiii). Etymology: The epithet is a Latin word, referring to the swampy habitat where the type was collected in Panama in 1861 by the US Army Surgeon Sutton Hayes (?–1863). Notes: Out of approximately 80 species in this predominantly wind-dispersed seed genus (sensu lato), the characteristic corky water-dispersed seeds with reduced wings of Tabebuia palustris is only shared with the estuarine T. fluviatilis, which occurs at the Atlantic coast from Venezuela to Brazil (Gentry, 1992). T. palustris has been observed in the field to occasionally present small fasciculate aerial roots, but this has never been reported in the literature. These two adaptations, highly specialized to semiaquatic environments, along with a narrow pattern of distribution and habitat, indicate that Tabebuia palustris must be regarded as a minor mangrove; it is restricted to mangroves, where it often forms fringe populations that grow on soft-muddy-water saturate- and poorly aired soils in hyposaline environments. The species is it said to be called “barbasco”, a vernacular name most commonly applied in our area to Muellera chocoensis (Fabaceae) and to several other species of trees, shrubs and lianas of unrelated families that inhabit the terra firme forests, which have swiftly biodegradable phytochemical components (e.g. rotenone) that can stun or paralyze fish, and are used principally to fish in several South American countries. Yet Tabebuia palustris has not been reported to have such usage. BROMELIACEAE (Bromeliad Family) Perennial herbs, sometimes stoloniferous, terrestrial or epiphytic. Leaves spirally arranged, usually in rosetes, simple, entire, spinose or serrate; primary and secondary nerves absent. Scape lacking or more often present and well-developed, usually bracteate. Inflorescence usually terminal, sometimes lateral, simple or compound; axes with flowers distichous or polystichous. Flowers 3-merous, subtended by a bract, bisexual or sometimes functionally unisexual. Sepals 3. Petals 3. Stamens 6, in 2 series. Ovary superior to inferior; styles 3-parted; stigmas 3. Fruit a dehiscent capsule, baccate or berry; seeds winged, caudate, plumose or unappendaged. Bromeliaceae is an almost exclusively Neotropical family, with only one species occurring in West Africa. Twenty five species of this family are herein reported to occur in the mangrove interior. Populations of all of those species mostly inhabit the terra firme habitats, from dry to rainforest life zones, and none is exclusively restricted to mangroves.

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Key to the species 1. Plants or leaves pendulous to subpendulous 2. Plants often with apparience of curtains; stem wiry, thin, very elongated and branched, to several meters long; leaves distichous, filiform, grayish Tillandsia usneoides 2. Plants never with apparience of curtains; without stem; leaves rosulate, ± strap-like, green 3. Plants with dimorphic leaves, the blades not spiral, thin, flat, narrowly-tapering; seeds bialate Pitcairnia heterophylla 3. Plants with monomorphic leaves, the blades loosely spiral, thick, linear-subterete; seed plumose Tillandsia subulifera 1. Plants or leaves erect 4. Leaf blades with margin spinose; fruit berry or baccate, globose to shortly-oblong 5. Leaf blades thick, involute, with white spots at base, attenuate at apex; inflorescence archingpendulous, the flowers ca. 3.5–4.5 cm long, green Billbergia macrolepis 5. Leaf blades relatively thin, not involute, without white spots at base, obtuse to acute at apex; inflorescence erect or suberect to arching (in Aechmea pyramidalis) but not pendulous, the flowers less than 2 cm long, yellow or pink to purple 6. Plants stoloniferous; inflorescence simple, the flowers polystichously arranged Aechmea nudicaulis 6. Plants not stoloniferous; inflorescence 1- to 3-branched, the flowers distichously arranged 7. Spikes 8–15 x 1.2–2 cm, yellowish–cream, subtended by conspicuous, bright-red bracts Aechmea dactylina 7. Spikes 2–5.5 x 0.5–1 cm, green, subtended by reduced green bracts 8. Leaves at base with teeth ± straight, 3–5 mm long; inflorescence cylindric, the flowers yellow Aechmea angustifolia 8. Leaves at base with teeth retrose, 5–10 mm long; inflorescence broadly pyramidal, the flowers pink or purple Aechmea pyramidalis 4. Leaf blades with margin entire; fruit capsule, cylindric-oblong to cylindric-linear-oblong 9. Inflorescence not branched 10. Flowers arranged in several rows (polystichous) 11. Inflorescence cylindric, the floral bracts of two colors, the lower ones pale-green with brown stripes, the upper ones bright-red; flowers white Guzmania monostachia 11. Inflorescence subglobose, all floral bracts reddish to bright-rose; flowers yellowish Guzmania musaica 10. Flowers arranged in two alternate rows (distichous) 12. Plants with several lateral divergent inflorescences per individual Tillandsia complanata 12. Plants with only one central erect inflorescence per individual 13. Plants bulbous at base; blades abruptly narrowing, thick, linear-subterete

Tillandsia bulbosa

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13. Plants forming a cup at base; blades ligulate, or gradually tapering, thin, attenuate or lineartriangular 14. Leaf blades 1–8 cm wide; scape stout, 5–15 mm diam. 15. Leaves 5–8 cm wide; floral bracts green, brown in fruit, the flowers greenish-white to creamish Werauhia gladioliflora 15. Leaves 1–3 cm wide; floral bracts pink, the flowers purple-blue to fuchsia or violet 16. Spikes 3–3.5 cm wide; corolla spreading, the petals with 2 scales at inner base Tilandsia barclayana var. minor 16. Spikes ca. 1 cm wide; corolla tubular, the petals without scales at inner base Tillandsia latifolia var. divaricata 14. Leaf blades 4–8 mm wide; scape slender, 2–4 mm diam. 17. Flowers widely spreading, forming a ± 90º angle with the rachis Tillandsia narthecioides 17. Flowers ascendent, forming an acute angle with the rachis Tillandsia triglochinoides 9. Inflorescence branched 18. Leaf blades ligulate 19. Plants to 20 cm high; inflorescence arching-pendulous; sepals asymmetric, 7–8 mm long; capsule ovoid, ca. 1.2 cm long, the seeds with coma folded over Catopsis sessiliflora 19. Plants 20 to 80 cm high; inflorescence erect; sepals ± symmetric, 1.5–2.5 cm long; capsule cylindric, 3–5 cm long, the seeds with coma straight 20. Leaf blades 15–40 x 2–5 cm; inflorescence slender, to 50 cm high 21. Leaves spreading; inflorescence (sub)erect, the spikes ca. 5–7, horizontal to suberect; rachis and floral bracts greenish, the floral bracts ca. 1 cm long, shorter or same length than sepals Tillandsia venusta 21. Leaves upright-tubular; inflorescence arching, the spikes 1–4, subpendulous; rachis and floral bracts red, the floral bracts ca. 3 cm long, exceeding sepals Tillandsia dyeriana 20. Leaf blades 40–100 x 5–8 cm; inflorescence stout, to 120 cm high 22. Petals without 2 basal appendages within 23. Inflorescence green; sepals densely lepidote Guzmania hitchcockiana 23. Inflorescence light-yellow; sepals glabrous Guzmania scherzeriana 22. Petals with 2 basal nectary appendages within 24. Sepals 2–2.5 cm long Werauhia cf. kupperiana 24. Sepals 2.5–4.5 cm long 25. Floral bracts foliaceous, partially disintegrating in fruits; sepals acute at apex; fruits ± in same direction than rachis Werauhia ringens 25. Floral bracts coriaceous, not disintegrating in fruits; sepals rounded at apex; fruits perpendicular to rachis

Werauhia sanguinolenta

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18. Leaf blades narrowly-triangular to filiform-attenuate 26. Plants bulbous or globose at base, the leaf blades abruptly narrowing into linear or filiformattenuate, 7–25 cm long 27. Leaf sheats (at base) with margins ciliate, the blades flexible; peduncle bracts green, the flowers yellow Tillandsia disticha 27. Leaf sheats (at base) with margins not ciliate, the blades thick and stiff; peduncle bracts purple-red, the flowers purple Tillandsia bulbosa 26. Plants forming a cup at base, the leaf blades gradually tapering, narrowly-triangular or lorate, 25–40 cm long 28. Inflorescences subdigitate or bipinnate, the floral bracts pink or red, the sepals symmetric 29. Inflorescence subdigitate, the spikes ascendent, forming an acute angle with rachis, the floral bracts red Tillandsia flagellata 29. Inflorescence bipinnate, the spikes widely spreading, ± forming a 90º angle with the rachis, the floral bracts pink Tillandsia latifolia var. divaricata 28. Inflorescence tripinnate, the floral bracts green, the sepals asymmetric Racinaea multiflora 30. Primary bracts narrowly-lanceolate, acuminate at apex, the lower ones exceeding axillary branches at maturity Racinaea multiflora var. multiflora 30. Primary bracts lanceolate, acute or short-acuminate at apex, all shorter than axillary branches Racinaea multiflora var. decipiens

55. Aechmea angustifolia Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 43, pl. 159 (1838). Plate 11: A. Syn.:

Hoplophytum angustifolium (Poepp. & Endl.) Beer Hohenbergia angustifolia (Poepp. & Endl.) Baker Aechmea cummingii Baker Aechmea spicata Baker Aechmea boliviana Rusby Aechmea cylindrica Mez Aechmea dryanderae Harms Aechmea eggersii Mez Aechmea inconspicua Harms Aechmea leucocarpa André Aechmea andradei Gilmartin Holoepiphytic herbs. Leaves rosulate; blades ligulate, 50–70 x 3–7 cm, spinose-serrate. Scape erect, white-flocculose, glabrescent; scape bracts elliptic, acute at apex, serrate, red. Inflorescence erect, bipinnate, cylindric, 15–40 cm long; lowest primary bracts like scape bracts, exceeding axillary spikes, the others abruptly reduced to about size of floral bracts;

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spikes spreading or reflexed, distichously 10-flowered, 20–55 mm long, the rachis undulate. Floral bracts spreading, broadly ovate, 5 mm long, barely exceeding ovary. Sepals asymmetric, 4–5 mm long, free. Petals linear, 8–10 mm long, yellow, bearing 2 scales well above base. Ovary ellipsoid. Fruit berry, white at maturity. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 6964 (GUAY). M: Neill et al. 11433 (QCNE). Distribution: Costa Rica to Venezuela and Bolivia. Ecology: In moist and wet forests. Biological interactions and dispersal: The species frequently houses ants in the basal rosette interior. The fleshy fruit is presumably eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. Use: Outside of mangroves the species is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The generic epithet comes from the Greek word aichme (spearhead or point). The nondistinctive specific epithet is formed by the Latin words angusta (narrow) and folia (leaves), meaning the narrow-leaved Aechmea. 56. Aechmea dactylina Baker, J. Bot., 17: 161 (1879). Plate 11: B. Holoepiphytic herbs. Leaves rosulate; blades ligulate, 40–100 x 6–10 cm, spinose-serrate with ± straight black teeth 5–10 mm long. Scape erect, to 1 cm diam., scape bracts imbricate, lanceolate, acute at apex, entire, bright-red. Inflorescence paniculate, 25–50 cm long, axis whitearachnoid; primary (spike) bracts oblong-lanceolate, spreading or reflexed. Spikes oblong, 8–15 x 1.2–2 cm, yellowish-cream, distichous-flowered, strongly complanate, laterally sharp; floral bracts broadly-ovate to suborbicular, 1.4–1.7 mm long. Sepals lanceolate, 7–8 mm long, alatecarinate, basally connate for 1 mm. Petals ligulate, yellow. Ovary sharply 3-angled. Fruit berry. COLOMBIA. CH: (E). N: (E). ECUADOR. E: Cornejo & Cangá 8155 (GUAY). Distribution: Nicaragua to northwestern Ecuador. Ecology: In moist forests, wet forests, and rainforests. Biological interactions and dispersal: The species frequently houses ants in the basal rosette interior. The fleshy fruit is presumably eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. Use: Outside of mangroves the species is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the basally subdigitate (finger-like) inflorescence.

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57. Aechmea nudicaulis (L.) Griseb., Fl. Britt. W. Indies 593 (1864). Plate 11: C. Bas.: Syn.:

Bromelia nudicaulis L. Billbergia nudicaulis (L.) Lindl. Hoplophytum nudicaule (L.) K. Koch Hohenbergia nudicaulis (L.) Baker Pothuava nudicaulis (L.) Regel Billbergia pyramidata Beer Hoplophytum lanuginosum Beer Billbergia lanuginosa hort. ex Beer Holoepiphytic herbs, stoloniferous, growing in dense clumps. Leaves rosulate; blades ligulate, 30–90 x 6–10 cm, spinose-serrate with coarse black teeth to 4 mm long. Scape slender, erect or arching, white-floccose; scape bracts imbricate, usually congested below inflorescence, lanceolate, acute at apex, entire, pink to bright-red. Inflorescence simple, cylindric, fertile throughout, 5–20 cm long, pale-floccoselepidote, glabrescent. Floral bracts minute or lacking. Flowers ca. 20 mm long. Sepals free, strongly asymmetric, 5–10 mm long. Petals ligulate, 12 mm long, yellow, bearing two fimbriate scales within. Ovary subglobose. Fruit berry, orange at maturity.

COLOMBIA. N: (E). ECUADOR. E: Cornejo & Cangá 8217 (GUAY). Distribution: Mexico to Peru and Brazil. Ecology: In dry, moist, and wet forests. Biological interactions and dispersal: The fleshy fruit is presumably eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. Use: Outside of mangroves the species is occasionally cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Latin word nudus (naked) and the Greek word kaulos (stem), referring to the basal stems often exhibited in a stoloniferous growth pattern. 58. Aechmea pyramidalis Benth., Bot. Voy. Sulfur 173 (1844). Plate 11: D. Syn.: Hohenbergia pyramidalis (Benth.) Baker Aechmea edmonstonei Baker Vernacular name: piñuela de monte. Holoepiphytic herbs. Leaves rosulate; blades ligulate, 60–90 x 4–7 cm, loosely serrate with broad flat retrorse teeth. Scape erect, slender; scape bracts erect, closely enfolding scape, imbricate, lanceolate, acute at apex, red. Inflorescence tripinnate, broadly pyramidal, to 1.2

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m long (incl. scape), white-pilose; axes slender, red; primary bracts like scape-bracts, much shorter than branches; spikes 3–5 cm long, ca. 10-flowered, dense, the rachis angled. Floral bracts broadly ovate, 8 mm long, spinose-acuminate. Flowers distichous. Sepals asymmetric, broadly elliptic, 5 mm long. Petals 8–10 mm long, pink or purple to blue. Ovary 4 mm long. Fruit berry. COLOMBIA. N: (E). ECUADOR. E: (S). G: Cerón 13344 (MO, QCNE). Distribution: Western Colombia, western Ecuador, and northwestern Peru. Ecology: In dry, moist, and wet forests. Biological interactions and dispersal: The flowers are observed to be visited by hummingbirds. The fleshy fruit is presumably eaten by birds, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. IUCN Red list category: The species is persistent in secondary forests and disturbed habitats, and is herein categorized as of Least Concern (LC). Etymology: The epithet refers to the pyramidal shape of the inflorescence. Note: The species type was collected in 1838 by Andrew Sinclair (1796–1861) in the environs of Guayaquil (Bentham and Hooker, 1844; Jørgensen and León, 1999). 59. Billbergia macrolepis L. B. Sm., Contr. Gray Herb. 114: 3–4, pl. 1, fig. 6 (1936). Plate 12: A. Syn.: Billbergia richteriana E. Pereira Holoepiphytic herbs. Leaves rosulate; blades erect, thick, narrowly-lanceolate, to 120 x 4 cm, attenuate at apex, sparsely serrate with nearly straight teeth 3 mm long, densely palelepidote, gray-green, marked with large white spots. Scape decurved, 3–4 mm thick, sparsely white-floccose; scape bracts imbricate, lanceolate, to 24 cm long, acuminate at apex, whitefloccose. Inflorescence flaccid-arching-pendulous, cylindric, to 40 cm long, loose, densely white-farinose. Floral bracts spreading or reflexed, narrowly-oblong or the uppermost ovate, to 35 mm long, cream-colored. Flowers sessile, suberect to spreading. Sepals symmetric, coriaceous, 10 mm long, nerved. Petals oblong, to 45 mm long, acute at apex, bronze-green. Ovary subglobose, 15 mm long, coarsely sulcate. Fruit baccate, globose, ca. 1.5 cm diam. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Bonifaz 1879 (GUAY [2], SEL). Distribution: Costa Rica to Colombia, Venezuela, and northwestern Ecuador. Ecology: In wet forests. Biological interactions and dispersal: Occasionally, the flowers are observed to be visited by hummingbirds during the day. However, the flaccid-pendulous inflorescences, bearing nocturnal flowers, suggest bat pollination. The baccate fruit is presumably eaten by birds, while the small seeds are endozoochorously dispersed.

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Use: This beautiful species could be cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The generic name honors Gustave Johannes Billberg (1772–1844), a Swedish botanist and zoologist. The non-distinctive epithet is formed by the Greek words macro (large) and lepis (scale), meaning the Billbergia with large scales. 60. Catopsis sessiliflora (Ruiz & Pav.) Mez in C. DC., Monograph. Phan. 9: 625 (1896). Plate 12: B. Bas.: Syn.:

Tillandsia sessiliflora Ruiz & Pav. Bromelia sessiliflora (Ruiz & Pav.) Lodd. ex Loudon Catopsis aloides (Schltdl. & Cham.) Baker Catopsis apricoides (Schltdl. & Cham.) Baker Catopsis modesta F. Muell. Catopsis nutans var. erecta Wittm. Holoepiphytic herbs. Leaves rosulate; blades ligulate, 13–20 x 1.2–2.5 cm, rounded and apiculate at apex, obscurely lepidote. Scape erect but arching, slender, glabrous; scape bracts erect, shorter than internodes, broadly-elliptic, apiculate at apex. Inflorescence simple or compound from a few loose spikes, to 11 cm long, glabrous. Spikes divergent, 2–9 cm long, loosely flowered. Floral bracts thin, broadly ovate, obtuse at apex, much exceeded by sepals, nerved. Flowers suberect. Sepals asymmetric, subchartaceous, suborbicular, 7–8 mm long, nerved. Petals lanceolate-ovate, barely exserted, white. Capsule ovoid, distinctly shortbeaked, ca. 12 mm long; seeds with coma apical and folded over.

ECUADOR. E: (S). Distribution: Central America to Peru and Brazil. Ecology: In wet forests. Dispersal: The plumose seeds are wind dispersed. IUCN Red list category: The species is persistent in secondary forests and in disturbed habitats. It is here categorized as of Least Concern (LC). Etymology: The epithet is formed by the Latin words sessilis (sessile) and flora (the goddess of flowers and the season of spring in Roman mythology), referring to the sessile flowers of the species. 61. Guzmania hitchcockiana L. B. Sm., Proc. Amer. Acad. Arts 70: 148, tab. 1, fig. 1 (1935). Plate 12: C. Holoepiphytic herbs. Leaves rosulate; blades ligulate, 60–120 x 5–7 cm, acute at apex, entire, densely lepidote without. Scape erect, stout, longer than leaves; scape bracts imbricate, broadly-ovate to lanceolate, greenish. Inflorescence bipinnate, 40–80 x 15–30 cm, the lateral branches spreading, densely lepidote; pedicels very shot and stout. Floral bracts broadlyovate, obtuse at apex, greenish, much exceeded by sepals. Sepals oblong, ca. 2 cm long,

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connate for 10 mm at base, obtuse at apex, densely lepidote. Corolla tubular, creamish-white, the petals ca. 3 cm long. Capsule cylindric, ca. 2.5–3 cm long. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Cangá 8265 (GUAY). Distribution: Southwestern Colombia and western Ecuador. Ecology: In wet and rain forests. Biological interactions: The flowers have been observed to be visited by hummingbirds. Dispersal: The plumose seeds are wind dispersed. Use: This species may be cultivated as an ornamental. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The genus was named after Anastasio Guzmán (?–1807), a pharmacist from Seville, Spain, and a teacher of botany, who died during an expedition to the Llanganates in Ecuador in search of lost Incan gold. Tafalla is assumed to have suggested the generic name to Ruiz and Pavón (Jørgensen and León, 1999). The epithet honors Albert Spear Hitchcock (1865–1935), an American botanist and agrostologist who in 1923 collected the type from wet forests near Bucay, in western Ecuador. 62. Guzmania monostachia (L.) Rusby ex Mez in C. DC., Monogr. Phan. 9: 905 (1896). Plate 12: D. Bas.: Renealmia monostachia L. Syn.: Tillandsia monostachia (L.) L. Vernacular name: pija de perro. Holoepiphytic herbs. Leaves rosulate; blades ligulate, 20–40 x 1.5–2 cm, acute at apex, entire, soon glabrous. Scape erect, stout, longer than leaves; scape bracts imbricate, ovate, pale green. Inflorescence simple, a cylindric-polystichous spike, 8–18 x 2–3 cm, sterile at apex. Floral bracts imbricate, foliaceous to membranaceous, ovate, acuminate to acute at apex, the fertile ones pale-green with brown stripes, the sterile bright-red, arranged at apex. Sepals coriaceous, obovate, 1–2.2 cm long. Corolla tubular, 2–4 x ca. 0.5 cm, white, the petals high-connate. Capsule cylindric, 2–3 cm long. COLOMBIA. N: Cabezas & Cortés 12 (CUVC). ECUADOR. E: Cornejo & Bonifaz 1877 (GUAY, SEL). G: Cornejo 42 (GUAY). Distribution: Southeastern United States (Florida), Central America to Peru and Brazil, and in the West Indies. Ecology: In dry, moist, and wet forests. Biological interactions: The flowers have been observed to be visited by hummingbirds. Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves the species is occasionally sold as an ornamental. IUCN Red list category: Guzmania monostachia is a widespread species persistent in secondary forests and in heavily disturbed habitats. It is here categorized as of Least Concern (LC).

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Etymology: The non-distinctive specific epithet is fomed by the Greek prefix mono (one) and the Greek word stachys (spike), referring to the simple spike that this species bears. Note: The cited collections are of the variety monostachia. The species has been misreported as Tillandsia cornuta in the REMACAM (Ecociencia, 2006: 50, fig. 7; Ministerio del Ambiente, 2009). 63. Guzmania musaica (Linden & André) Mez, Monogr. Phan. 9: 898 (1896). Plate 13: A. Bas.: Tillandsia musaica Linden & André Holoepiphytic herbs; usually spreading by elongate rhizomes. Leaves rosulate; blades ligulate, 25–70 x 4–8 cm, broadly-acute or rounded and apiculate at apex, entire, obscurely punctulatelepidote. Scape erect, stout, somewhat shorter than leaves; scape bracts densely imbricate, broadly-elliptic, bright-rose. Inflorescence simple, subglobose. Flowers subsessile. Floral bracts broadly-obovate, apiculate at apex, bright-rose to red, about half as long as sepals. Sepals oblong, 2–4.5 cm long, highly connate, obtuse at apex. Corolla tubular, yellowish, the petals ca. 3.5 cm long. Capsule cylindric, ca. 3 cm long. COLOMBIA. CH: (E) V: Killip 5321 (NY). Distribution: Costa Rica to Peru, and Venezuela. Ecology: In wet and rain forests. Biological interactions: No information is recorded, but the bright-rose floral bracts and tubular flowers suggest pollination by hummingbirds. Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves this beautiful species is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet, which means the Musaceae-like Guzmania, refers to the red bracts. The bracts were thought by the authors to resemble most species of Heliconia (a genus that at the time of its basionym publication was placed in Musaceae, though it was later segregated to Heliconiaceae). 64. Guzmania scherzeriana Mez in C. DC., Monogr. Phan. 9: 949 (1896). Syn.: Guzmania guatemalensis L. B. Sm.

Guzmania scherzeriana var. guatemalensis (L. B. Sm.) L. B. Sm. Guzmania herthae Harms Holoepiphytic herbs. Leaves rosulate; blades ligulate, 40–70 x 4–6 cm, acute at apex, entire, glabrous. Scape erect, stout, longer than leaves; scape bracts imbricate, broadly-ovate to lanceolate, yellowish. Inflorescence paniculate, 25–40 x 15–30 cm, the lateral branches

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suberect to spreading; pedicels 4–6 mm long. Floral bracts broadly-ovate or elliptic, obtuse at apex, yellowish, much exceeded by sepals. Sepals elliptic-oblong, 2–2.5 cm long, connate to 10 mm at base, obtuse at apex. Corolla tubular, white, the petals ca. 3.3 cm long. Capsule cylindric, ca. 3 cm long. COLOMBIA. CH: (E). V: Coll. unknown s.n. (CUVC). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá (S). Distribution: Belize to Ecuador. Ecology: In wet and rain forests. Biological interactions: The flowers have been observed to be visited by hummingbirds. Dispersal: The plumose seeds are dispersed by wind. Use: This species may be cultivated as an ornamental. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat in near future, it is herein categorized as Near Threatened (NT). Etymology: The epithet honors Karl von Scherzer (1821–1903), an Austrian explorer who in 1853 collected the type in Costa Rica. 65. Pitcairnia heterophylla (Lindl.) Beer, Fam. Bromel. 68 (1856). Bas.: Puya heterophylla Lindl. Holoepiphytic herbs. Leaves rosulate; blades dimorphic, the outer reduced, castaneous, spinose-serrate, the inner subpendulous, lorate, ca. 30–50 x 1–2 cm, acute at apex, paleflocculose beneath, soon glabrous, deciduous before anthesis. Scape very short and concealed by leaves; scape bracts ovate, white-flocculose. Inflorescence capitate. Floral bracts like the upper scape bracts. Flowers erect; pedicels ca. 3 mm long. Sepals narrowly-subtriangular, ca. 3 cm long, subalate-carinate. Petals lorate, to ca. 5.5 cm long. Capsule cylindric, ca. 2 cm long. COLOMBIA. CH: (E). N: (E). ECUADOR. E: (S). Distribution: Mexico to Peru. Ecology: In dry, moist and wet forests. Dispersal: The caudate seeds are wind dispersed. Use: Outside of mangroves the species is occasionally cultivated as an ornamental. IUCN Red list category: This widespread species, persistent in secondary forests, it is here categorized as of Least Concern (LC). Etymology: The epithet refers to the two different leaf blades form.

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66. Racinaea multiflora (Benth.) M. A. Spencer & L. B. Sm., Phytologia 74(2): 155 (1993). Holoepiphytic herbs. Leaves rosulate; blades narrowly-lanceolate, ca. 20–40 x 1.5–2.5 cm, long caudate-reflexed at apex, densely lepidote throughout. Scape erect, glabrous; scape bracts greatly exceeding internodes, long-caudate at apex, stramineous, lepidote. Inflorescence erect, loosely tripinnate, to 40 cm long; primary bracts lanceolate or narrowlylanceolate, acuminate to acute at apex; branches spreading; spikes oblong, 3–5 x ca. 1 cm, rather densely flowered, subsessile or short stipitate, the rachis geniculate, strongly angled, sulcate, glabrous. Floral bracts sharply carinate, ovate, 3–5 mm long, nerved. Sepals free, asymmetric, sharply carinate, oblong, 5 mm long, acute at apex, nerved. Petals 6 mm long. Stamens and pistil included. Capsule cylindric, ca. 4 cm long. 66a. Racinaea multiflora var. multiflora. Plate 13: B. Bas.: Tillandia multiflora Benth. Primary bracts narrowly-lanceolate, acuminate at apex, the lower ones exceeding axillary branches at maturity. ECUADOR. E: Cornejo & Bonifaz 1883 (GUAY, SEL). Distribution: Western Ecuador. Ecology: In moist forests. Dispersal: The plumose seeds of this variety are wind dispersed. Etymology: The non-distinctive epithet means the many-flowered Racinea. IUCN Red list category: Least Concern (LC). Note: The type of this variety was collected in 1838 by Andrew Sinclair (1796–1861), an Englishman naturalist, in the surroundings of Guayaquil, Ecuador (Bentham and Hooker, 1844; Jørgensen and León, 1999). 66b. Racinaea multiflora var. decipiens (André) M. A. Spencer & L. B. Sm., Phytologia 74(2): 155 (1993). Plate 13: C. Bas.: Tillandsia decipiens André Primary bracts lanceolate, acute or short-acuminate at apex, all shorter than axillary branches. ECUADOR. E: Cornejo & Bonifaz 2617 (GUAY). Distribution: Western Ecuador and northwestern Peru. Ecology: In very dry and dry forests. Dispersal: The plumose seeds of this variety are wind dispersed.

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IUCN Red list category: This variety has a narrow range of distribution, persisting in secondary forests and in disturbed habitats. It is here categorized as of Least Concern (LC). Etymology: The varietal epithet means the deceiving Racinea multiflora. 67. Tillandsia barclayana Baker var. minor (Gilmartin) Butcher, Brom. Soc. Queensland 43(6): 5 (2009). Plate 13: D. Bas.: Vriesea barclayana (Baker) L. B. Sm. var. minor Gilmartin Syn.: Tillandsia lateritia André Holoepiphytic herbs. Leaves rosulate; blades coriaceous, narrowly-lanceolate, 15–25 x 1–2 cm, attenuate at apex, cinereous. Scape erect to subpendular, stout; scape bracts densely imbricate, lepidote. Inflorescence simple, erect, complanate, oblongoid, 5–12 x 2.5–3.5 cm, 9–18-flowered, the rachis straight. Floral bracts equaling or exceeding sepals, imbricate, convex, coriaceous, obovate, 1.8–3 cm long, pink. Sepals free, ecarinate, coriaceous, elliptic, 1.7–2 cm long. Petals violet, bearing two large highly adnate scales at base. Capsule 3.5–5 cm long. COLOMBIA. C: Cabrera 3818 (CUCV). N: (S). ECUADOR. E: Cornejo & Bonifaz 1878 (GUAY). Distribution: Southwestern Colombia and western Ecuador. Ecology: In wet forests. Biological interactions: No information recorded, but the pink color floral bracts and the floral structure suggest pollination by hummingbirds. Dispersal: The plumose seeds are wind dispersed. Use: This species may be cultivated as an ornamental. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat in near future, it is herein categorized as Near Threatened (NT). Etymology: The specific epithet honors George W. Barclay (dates unknown), an Englishman who in 1836 collected the type in Valdivia, Province of Santa Elena, Ecuador. The varietal epithet refers to the smaller size that exhibit the plants of this variety against the somewhat larger plant size that present the typical variety. 68. Tillandsia bulbosa Hook., Exot. Fl. 3: pl. 173 (1826). Plate 14: A. Syn.: Tillandsia pumila Griseb. Holoepiphytic herbs. Leaves rosulate, bulbous at base; blades abruptly narrowing, thickly coriaceous, linear, subterete-involute, 7–25 cm long, appressed-lepidote, spreading. Scape shorter than leaves, covered by purple bracts. Inflorescence spicate, the spikes 1–6, compressed, densely flowered, 4–7 cm long. Floral bracts conduplicate, ovate-lanceolate, 1–1.5 cm long, usually red. Sepals ovate, 12 mm long. Petals ca. 2.5 cm long, purple. Capsule cylindric, 3–4 cm long.

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COLOMBIA. CH: Betancur & Gil 7956 (COL). V: Estudiantes del curso de Ecología Vegetal 13 (CUVC). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 1880 (GUAY, SEL). Distribution: Central America to Brazil and Ecuador, and in the West Indies. Ecology: In wet and rain forests. Biological interactions: No information recorded, but the floral structure suggests pollination by hummingbirds. Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves the species is widely cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the bulbous base. 69. Tillandsia complanata Benth., Bot. Voy. Sulphur 173 (1846). Plate 14: B. Syn.: Tillandsia axillaris Griseb.

Tillandsia complanata subsp. latifolia Gilmartin Holoepiphytic herbs. Leaves rosulate, streaked or suffused with dark purple especially at base; blades ligulate, 20–30 x 3–5 cm, abruptly acute or subobtuse, and apiculate at apex, obscurely punctulate-lepidote, soon glabrous above. Scapes many from axils of leaves, ascending, 1–2 mm thick, glabrous; scape bracts erect, imbricate, narrowly-lanceolate. Inflorescences simple spikes, oblong-lanceolate, 3–7 x 1–2 cm, acute at apex, complanate, 4–24-flowered, glabrous. Floral bracts imbricate, ecarinate, coriaceous, elliptic, 1.5–2.5 cm long, exceding sepals. Sepals lanceolate. Petals ca. 2 cm long, rose or purple. Capsule slenderly cylindric, ca. 4 cm long.

ECUADOR. E: Cornejo & Bonifaz 2615 (GUAY). Distribution: Costa Rica to Bolivia. Ecology: This species has a huge range of ecological distribution, from sea level to paramos; in dry, moist, wet, and rain forests. In dry forests, the ocurrence of Tillandsia complanata is always associated to those localities which are influenced by fog and winds. Biological interactions: The flowers have been observed to be visited by hummingbirds. Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves the species is occasionally cultivated as an ornamental. IUCN Red list category: This species is persistent in secondary forests and in disturbed habitats. It is here categorized as of Least Concern (LC). Etymology: The non-distinctive epithet is a Latin word, that refers to the laterally compressed spikes. Note: The type was collected in 1838 by Andrew Sinclair (1796–1861), an Englishman naturalist, in Atacames, Province of Esmeraldas, northwestern Ecuador (Bentham and Hooker, 1844; Jørgensen and León, 1999).

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70. Tillandsia disticha Kunth, Nov. Gen. Sp. Pl. 1: 292 (1816). Plate 14: C. Syn.: Tillandsia disticha var. gracillima Mez Holoepiphytic herbs. Leaves rosulate, bulbous at base; blades involute-subulate, 15–30 cm x 3–4 mm, filiform-attenuate at apex, covered with appressed cinereous scales. Scape erect; scape bracts imbricate, lanceolate, lepidote. Inflorescence digitate from a few spikes; primary bracts lanceolate; spikes subsessile, oblong, 4–6 x ca. 0.8 cm, dense, complanate. Floral bracts carinate, triangular, 6–10 mm long, nerved. Flowers subsessile. Sepals carinate, glabrous. Petals ca. 13 mm long, yellow. Stamens and pistil included. Capsule slenderly cylindric, ca. 5 cm long. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 1881 (GUAY, SEL). G: Cornejo 8 (GUAY). Distribution: Western Colombia to Bolivia. Ecology: In dry, moist and wet forests. Biological interactions: Ants are often housed in the basal pseudobulb interior. Dispersal: The plumose seeds are wind dispersed. IUCN Red list category: This species is persistent in secondary forests and in disturbed habitats. It is here categorized as of Least Concern (LC). Etymology: The non-distinctive epithet is a Latin word, that refers to the densely alternate (distichous) arrangement of flowers in the spikes. 71. Tillandsia dyeriana André, Énum. Bromél. 8 (1888). Plate 14: D. Holoepiphytic herbs. Leaves upright-tubular; blades ligulate, 15–20 x ca. 2 cm, pale green and dark red/purple-spotted. Scape erect, slender; scape bracts imbricate, lanceolate-elliptic. Inflorescence arching, mostly digitate; primary bracts like scape bracts; spikes 1–4, sessile, subpendulous, oblong, strongly complanate, the rachis slender, merely angled. Floral bracts thin, ca. 30 mm long, exceeding sepals, carinate and incurved at apex, bright-red or glowing golden-yellow, finely nerved. Flowers sessile. Sepals ecarinate, coriaceous, sublanceolate, ca. 22 x 7 mm, acute at apex, reddish. Petals white, the lobes spreading, broad, 6 mm long, rounded at apex. Stamens and pistil deeply included. Capsule over 3.2 cm long. ECUADOR. E: Dodson & Embree 13098 (SEL). Distribution: Endemic to lowlands of western Ecuador. Ecology: In wet forests. Biological interactions: No information recorded, but the bright-red floral bracts and the floral structure suggest pollination by hummingbirds.

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Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves this beautiful species is occasionally cultivated as an ornamental. IUCN Red list category: Critically Endangered (CR A4c; León-Yánez et al. eds., 2011). Etymology: The epithet honors M. T. Dyer (dates unknown), an Englishman botanist. 72. Tillandsia flagellata L. B. Sm., Phytologia 6: 258, t. 1, fig. 9, 10 (1958). Plate 15: A. Syn.: Tillandsia lehmannii Rauh Holoepiphytic herbs. Leaves rosulate; blades narrowly-lanceolate, 30–40 cm long, filiformattenuate at apex, covered with fine cinereous scales. Scape erect or suberect; scape bracts densely imbricate, strongly nerved, lepidote, long-flagellate and contorted linear at apex. Inflorescence subdigitate, ca. 7 spikes; primary bracts like scape bracts, erect; spikes divergent to spreading, narrowly-lanceolate, 4–7 x 1 cm, strongly complanate, densely few-flowered, the rachis narrowly-winged. Floral bracts lanceolate, ecarinate, thin, to 25 mm long, exceeding sepals, lepidote, strongly nerved, fuchsia. Sepals ecarinate, oblong-elliptic, ca. 16 mm long, obtuse at apex, even, lustrous, glabrous. Corolla tubular, the petals white with purplish-blue at distal half. Stamens 6, yellow, exserted. Capsule slenderly cylindric, 3–3.5 cm long. ECUADOR. G: Cornejo 7 (GUAY, SEL). Distribution: Southwestern Ecuador (and northwestern Peru?). Ecology: In dry forests. Biological interactions: No information recorded, but the fuchsia floral bracts and the tubular flowers suggest pollination by hummingbirds. Dispersal: The plumose seeds are wind dispersed. Use: This species may be cultivated as an ornamental. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat in near future, it is herein categorized as Near Threatened (NT). Etymology: The epithet is a Latin derived word flagellum (whip) referring to the narrowly-tapering elongate apices of scape bracts. 73. Tillandsia latifolia var. divaricata (Benth.) Mez in DC., Monogr. Phan. 9: 789 (1896). Plate 15: B. Bas.: Tillandsia divaricata Benth. Syn.: Tillandsia minor Mez & Sodiro ex Mez

Tillandsia kunthiana var. divaricata (Benth.) L. B. Smith Holoepiphytic herbs. Leaves rosulate; blades coriaceous, narrowly-lanceolate, 25–45 x ca. 3 cm, filiform-attenuate and recurved at apex, covered with appressed cinereous scales. Scape erect

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or suberect, stout; scape bracts imbricate, cinereous-lepidote. Inflorescence bipinnate or rarely simple, with spreading spikes; primary bracts usually shorter than spikes; spikes subsessile, lanceolate, 6–12-flowered. Floral bracts densely imbricate, coriaceous, carinate, broadly ovate, to 15 mm long, pink, lustrous. Flowers subsessile. Sepals connate for 8 mm, carinate, 12–17 mm long, sparsely lepidote. Corolla tubular, the lobes to 7 mm long, purple-blue to fuchsia. Stamens included, yellow, the filaments geniculate. Capsule cylindric, 3–4 cm long. ECUADOR. E: Cornejo & Bonifaz 2616, 2618 (GUAY, SEL). Distribution: Ecuador and Peru. Ecology: In dry thorn scrub, very dry, and dry forests. Out of mangroves, it also can be found as a terrestrial herb. Biological interactions: The flowers have been observed to be visited by hummingbirds. Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves the species is cultivated as an ornamental. IUCN Red list category: This species is persistent in secondary forests and in disturbed habitats. It is here categorized as of Least Concern (LC). Etymology: The non-distinctive specific epithet refers to the divaricate spikes arrangement. Note: The type of this variety was collected in 1838 by Andrew Sinclair (1796–1861), an Englishman naturalist, in the coastal dry forests of Santa Elena Province, Ecuador (Bentham and Hooker, 1844; Jørgensen and León, 1999). 74. Tillandsia narthecioides C. Presl, Reliq. Haenk. 1(2): 125 (1827). Plate 15: C. Holoepiphytic herbs. Leaves rosulate; blades subchartaceous, linear-lanceolate, 20–30 cm x ca. 5 mm, long-attenuate and recurved at apex. Scape erect or suberect, slender, scape-bracts tubular-involute, erect, the lower imbricate. Inflorescence simple, loose, oblong, 12–15 x ca. 2.5 cm, to 30-flowered, the rachis completely exposed, glabrous. Floral bracts spreading, scarcely carinate, chartaceous, linear-elliptic, ca. 13 mm long, obtuse at apex, strongly nerved, glabrous. Flowers sessile. Sepals equally, subfree, coriaceous, elliptic, 7–15 mm long, strongly nerved, glabrous. Petals white, the lobes spreading, 6 mm long. Stamens deeply included, exceeding the pistil, the filaments highly connate. Capsule slenderly cylindric, 3–4 cm long. ECUADOR. G: (S). Distribution: Southwestern Colombia to western Ecuador (northwestern Peru?). Ecology: In moist and wet forests. Biological interactions: The flowers have been observed to be visited by hummingbirds.

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Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves the species is occasionally cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to some resemblance of the leaves to those of Narthecium (Nartheciaceae). Note: The type was collected by the Czech botanist Thaddäus Haenke in the environs of Guayaquil, between 1790–1791, during the Alejandro Malaspina’s expedition around the world (1761–1816; Presl, 1825–1935; Jørgensen and León, 1999). 75. Tillandsia subulifera Mez, Repert. Spec. Nov. Regni Veg. 16(448–455): 74 (1919). Plate 15: D. Holoepiphytic, herb, growing (upside-down) with positive geotropism. Leaves contortedtubular; blades thickly coriaceous, stiff, linear-lanceolate, 15–20 cm x ca. 5–7 mm, with faint white cross-bands, abruptly acute at apex. Scape pendulous, stout, concealed by leaves; scape-bracts erect, imbricate. Inflorescence simple, loose, oblong, 5–8 x ca. 2.5 cm, 4–6– flowered, the rachis exposed, geniculate, the pedicels short. Floral bracts erect, chartaceous, elliptic, ca. 20 mm long, broadly-acute and incurved at apex, nerved, greenish, appressedlepidote. Sepals free, chartaceous, elliptic, ca. 22 mm long, nerved, appressed-lepidote. Petals tubular-erect, ca. 32 mm long, white. Stamens exserted. Capsule cylindric, 8–10 cm long. ECUADOR. E: (S). Distribution: Nicaragua to northwestern Ecuador, Venezuela, and Trinidad. Ecology: In wet forests. Biological interactions: No information recorded, but the pendulous habit and the tubular corolla suggest pollination by hummingbirds or bats. Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves the species is occasionally cultivated as an ornamental. IUCN Red list category: This widespread species is persistent in secondary forests and in disturbed habitats; therefore, it is here categorized as of Least Concern (LC). Etymology: The epithet refers to the abruptly acute apex of leaves. 76. Tillandsia triglochinoides C. Presl, Reliq. Haenk. 1(2): 125 (1827). Plate 16: A. Syn.: Tillandsia hartwegiana Brong. ex Baker Vernacular name: pikuyo. Holoepiphytic herbs. Leaves rosulate; blades subchartaceous, linear-lanceolate, 20–30 cm

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x ca. 7 mm, long-attenuate at apex. Scape erect or suberect, slender; scape bracts tubularerect, imbricate, sublanceolate. Inflorescence simple, loose, strongly complanate, narrowlylanceolate, 12–18 x 0.8–1 cm. Floral bracts suberect, ecarinate, submembranous, triangularovate, ca. 2 cm long, enfolding and exceeding sepals, nerved, glabrous. Flowers subsessile. Sepals equally, very short connate, coriaceous, lanceolate-ovate, to 12 mm long, lustrous. Petals ca. 2 cm long, white, the lobes broadly-lanceolate, reflexed at anthesis. Stamens deeply included. Capsule ca. 4 cm long. ECUADOR. E: Cornejo & Bonifaz 2605 (GUAY [2]). G: Cerón 18476 (MO). Distribution: Western Ecuador. Ecology: In dry, moist and wet forests. Phenology: This species has been collected with flowers in March and May, and with fruits from August to November. Biological interactions: No information recorded, but the floral structure suggests pollination by hummingbirds or moths. Dispersal: The plumose seeds are wind dispersed. Use: It may be cultivated as an ornamental. IUCN Red list category: Although it is persistent in secondary forests, due to factors as the small range of distribution and a loss of habitat, it is here categorized as Vulnerable (VU B1abiii). Etymology: The epithet refers to the resemblance of the leaves to those of the genus Triglochin (Juncaginaceae). Note: The type was collected by the Czech botanist Thaddäus Haenke in the environs of Guayaquil, between 1790–1791, during the Alejandro Malaspina’s expedition around the world (1761–1816; Presl, 1825–1935; Jørgensen and León, 1999). 77. Tillandsia usneoides (L.) L., Sp. Pl., ed. 2, 1: 411 (1762). Plate 16: B. Syn.:

Renealmia usneoides L. Dendropogon usneoides (L.) Raf. Strepsia usneoides (L.) Nutt. ex Steud. Tillandsia crinita Willd. ex Beer Tillandsia trichoides Kunth

Vernacular names: barba, barba de viejo, gabazo, musgo, salvaje. Holoepiphytic herbs, pendulous to several meters in length. Stem elongate, to 1 mm diam., flexuose, branched, hanging in festoons, grayish. Leaves distichous; blades filiform, divergent, 25–50 x ca. 1 mm, densely lepidote. Scape absent; flowers solitary, rarely 2 together, pseudolateral. Floral bracts ovate, shorter than sepals. Sepals narrowly-ovate, to 7 mm long, pale green. Petals narrow, 9–11 mm long, paler-green. Capsule slenderly cylindric, 2–2.5 cm long.

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COLOMBIA. N: (S). ECUADOR. E: Cornejo & Bonifaz 1882 (GUAY, SEL). G: Cornejo 5 (GUAY). Distribution: Southeastern United States and Mexico to Argentina and Chile. Ecology: Tillandsia usneoides is a common epiphyte in some windy areas in mangroves as well as in terra firme ecosystems. It occurs in very dry, dry, moist, and wet forest life zones. The presence of this species in very dry and dry life zones is always associated to those localities influenced by fog and winds. In our area, T. usneoides grows hanging down vertically from the branches of the Rhizophora trees mainly. Due to the flaccid and densely intertangled stems and recurved leaves it often has a curtain-like appearance. Dispersal: The plumose seeds are wind dispersed. Use: In many Latin American countries (including those of our area) this species it is often traditionally used and locally sold as a Christmas decoration. Despite the massive harvest of T. usneoides for this purpose, the native populations of this species apparently still remains abundant and healthy. IUCN Red list category: This widespread species is persistent in secondary forests and in heavily disturbed habitats; therefore, it is herein categorized as of Least Concern (LC). Etymology: The epithet refers to the resemblance of the plants of this species to those of Usnea (Lichen). Note: This is one of the first epiphytes reported in the literature from mangroves of Ecuador (Guppy, 1906). However, due to its unusual appearance that differs from most common bromeliads that have rosulate leaves with green and ± expanded leaf blades, and a usually conspicuous inflorescence, T. usneoides could not be recognized by non botanists as a Bromeliaceae. It is often mistaken as a lichen or a moss. 78. Tillandsia venusta Mez & Werclé in Mez, Bull. Herb. Boissier, sér. 2, 5: 108 (1905). Plate 16: C. Holoepiphytic herbs. Leaves rosulate; blades coriaceous, ligulate, 20–30 x 3–4.5 cm, purplespotted. Scape erect, slender; scape bracts membranaceous, broadly lanceolate. Inflorescence loosely bipinnate, composed of a terminal and 2 lateral spikes, ca. 13 cm long, glabrous; primary bracts like scape bracts; spikes subspreading, the rachis straight, angled. Floral bracts membranaceous, ecarinate, elliptic, shorter than sepals, prominently nerved. Flowers dense, subspreading, sessile. Sepals free, ecarinate, coriaceous, lanceolate, ca. 15 mm long, acute at apex. Petals 25 mm long, white, the lobes spreading at anthesis, triangular-ovate, ca. 1.2 x 1 cm. Stamens and pistil deeply included. Capsule ca. 4 cm long. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 6958 (GUAY). Distribution: Costa Rica to Ecuador. Ecology: In moist and wet forests. Biological interactions: The flowers have been observed to be visited by hummingbirds. Dispersal: The plumose seeds are dispersed by wind. Use: Outside of mangroves the species is occasionally cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin derived word from the goddess Venus, that refers to the beauty of this species.

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79. Werauhia gladioliflora (H. Wendl.) J. R. Grant, Trop. Subtrop. Pflanzenwelt 91: 31, f. 2 (1995). Plate 16: D. Bas.: Tillandsia gladioliflora H. Wendl. Syn.: Vriesea gladioliflora (H. Wendl.) Antoine Holoepiphytic herbs. Leaves rosulate; blades ligulate, 40–70 x 5–8 cm, entire, obscurely punctulate-lepidote beneath. Scape erect, very stout, covered by scape bracts; scape bracts imbricate, elliptic, green to brownish-green. Inflorescence simple, erect, subcylindric, 30–70 cm long x 4–5 cm wide, densely flowered; flowers distichous, the pedicels very short and stout. Floral bracts erect, very broadly-ovate, equaling to much exceeding sepals, 4–6.5 x 3–4 cm, coriaceous, ecarinate, obtuse or broadly-subacute at apex. Sepals broadly elliptic, 2–4.5 cm long. Petals greenish to greenish-white or creamish, 4–7 cm long, bearing two scales at base. Capsule 3–4 cm long. COLOMBIA. CH: Fuchs & Zanella 22067 (MO). V: Gentry et al. 40425, 40708 (MO). Distribution: Southern Mexico to Brazil. Ecology: In wet forests and rainforests. Biological interactions: The flowers have been reported to be pollinated by bats (Fleming et al., 2009). Dispersal: The plumose seeds are wind dispersed. Use: This species may be cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the flower resemblance of this species to those of Gladiolus (Iridaceae). 80. Werauhia cf. kupperiana (Suess.) J. R. Grant, Trop. Subtrop. Pflanzenwelt 91: 32. (1995). Plate 17: A. Bas.: Vriesea kupperiana Suess. Holoepiphytic herbs. Leaves rosulate; blades ligulate, ca. 125 x 7 cm, entire. Scape erect, stout, covered by scape bracts; scape bracts lanceolate, 14–20 cm (incl. sheath ca. 4.5 cm long), long-caudate at apex, green. Inflorescence panicle, erect, ca. 120 cm (incl. scape 50 cm); basal spike bracts 10–12 cm long; lateral spikes ca. 13 cm long, the rachis flexuose, 4-ribbed. Floral bracts 3–4 x 2 cm, hyaline at margins. Sepals free, somewhat symmetric, ovate-lanceolate, 2–2.5 cm long, striate within. Mature flowers and fruits not seen. ECUADOR. Esmeraldas: Cornejo et al. 7305 (GUAY). Ecology: In wet forests and rainforests. Dispersal: The plumose seeds are wind dispersed. Use: This species may be cultivated as an ornamental.

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IUCN Red list category: Collections with mature flowers are needed for a final confirmation of its identity. At the present, it is categorized as Data Deficient (DD). Etymology: The epithet honors Mr. Kupper (dates unknown), who in 1932 collected it in sterile condition in Costa Rica. Seven years later, Kupper’s collection flowered in the Botanical Garden in Munchen, and thus was described as new to science. 81. Werauhia ringens (Griseb.) J. R Grant, Trop. Subtrop. Pflanzenwelt 91: 35 (1995). Plate 17: B. Bas: Tillandsia ringens Griseb. Holoepiphytic herbs. Leaves rosulate; blades ligulate, 40–80 x 6–8 cm, entire. Scape erect, stout, covered by scape bracts; scape bracts imbricate, elliptic-lanceolate, pale-green, glabrous. Inflorescence a loosely compound panicle, erect, to 100 cm long (incl. scape 30–50 cm), the branches suberect; lateral spikes ca. 15–20 cm long, the rachis flexuose, subquadrate. Floral bracts broadly-ovate, 3–6 x 2–3 cm, exceeding internodes, straight and carinate at apex. Sepals elliptic, 2.5–3.5 cm long. Petals white or yellow, bearing two scales at base. Capsule 4–5 cm long. COLOMBIA. CH: Betancur & Gil 7919 (CUVC). V: (E). C: (E). N: (E). ECUADOR. E: Cornejo & Cangá 8198 (GUAY). Distribution: Nicaragua to western Ecuador, and in the West Indies. Ecology: In wet and rain forests. Phenology: This species has been collected with flowers in July, and with fruits in March. Biological interactions: It is most likely that the flowers are pollinated by bats. This inner funnel base is inhabited by insects (including mosquitoe) larvae. Dispersal: The plumose seeds are wind dispersed. Use: It may be cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet is derived from the Latin word ringens (gaping), that refers to the gaping corolla. 82. Werauhia sanguinolenta (Cogn. & Marchal) J. R. Grant, Trop. Subtrop. Pflanzenwelt 91: 35 (1995). Plate 17: C. Bas: Vriesea sanguinolenta Cogn. & Marchal Holoepiphytic herbs. Leaves rosulate; blades ligulate, 40–70 x 5–8 cm, entire. Scape erect, stout, covered by scape bracts; scape bracts imbricate, broadly-ovate to lanceolate, 10–25 cm

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long (incl. sheath 6–10 cm long), long-caudate at apex, green, glabrous. Inflorescence fewbranched panicle, erect, to 100 cm long (incl. scape 40–60 cm), the branches suberect; lateral spikes ca. 20–25 cm long, the rachis flexuose, 4-ribbed. Floral bracts asymmetric, becoming secund with flowers, 3–5 x 2–3 cm, more than twice as long as internodes, incurved and carinate at apex. Sepals free, somewhat symmetric, broadly-elliptic or ovate, 3–4.5 cm long, striate and punctulate-lepidote within. Petals white, bearing two scales at base. Capsule ca. 5 cm long, perpendicular to rachis. COLOMBIA. CH: Betancur 7931 (CUVC). V: (E). N: (E). ECUADOR. E: Cornejo & Cangá 8116 (GUAY). Distribution: Nicaragua to Ecuador, and in the West Indies. Ecology: In wet and rain forests. Biological interactions: It is most likely that the flowers are pollinated by bats. This plant is inhabited by scorpions, salamanders, crabs and insects (including mosquitoe) larvae. Dispersal: The plumose seeds are wind dispersed. Use: Outside of mangroves the species is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word, that refers to the blood-like purple-red color that sometimes is present in the leaves. CACTACEAE (Cactus Family) Arborescent, shrubby, herbs or lianas, stem-succulents, rarely non-succulent. Leaves absent, scale-like, subulate or rarely foliaceous, subtending axillary structures termed areoles, which are covered with trichomes and clusters of spines in various forms, and which give rise to flowers and new stems. Flowers solitary, sessile, usually perfect and actinomorphic. Bracts in gradual transition to perianth; subtended areole variously invested with whool, hairs, bristles or spines, or naked. Perianth segments 5 to numerous, composed of sepaloids and petaloids. Stamens 10 to numerous. Ovary inferior, style simple; stigmas separate. Fruit baccate, variously bracteate; seeds numerous.

This family in mangroves is represented by epiphytic species only.

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Key to the species 1. Secondary stems flattened; spines absent 2. Stems 2.5–8 cm wide; flowers 20–30 cm long; fruits oblong, 3–6 cm long, red Epiphyllum rubrocoronatum 2. Stems 0.5–1 cm wide; flowers 6–8 mm long; fruits globose, 6–8 mm long, white Rhipsalis micrantha 1. Secondary stems trigonous; spines present

Hylocereus polyrhizus

83. Epiphyllum rubrocoronatum (Kimnach) Dodson & A. H. Gentry, Selbyana 2: 31 (1977). Plate 17: D. Bas.: Epiphyllum phyllanthus var. rubrocoronatum Kimnach Vernacular name: pitahaya. Holoepiphytic herbs, ± suberect with pendulous branches. Stems flattened, the first order stems terete or trigonous, 0.1–3 m long, the secondary order stems pseudo-petiolate, 1–5 cm long; phylloclade-blade stiff, linear oblong, 30–100 x 2.5–8 cm, coarsely serrate or crenate at margins. Areoles 3–6 cm apart. Spines absent. Flowers salver-shaped, to 30 x 8–10 cm, nocturnal, clove odor fragrant. Sepaloids 10–12, linear-lanceolate, 15–50 x 2–4 mm, greenwhite. Petaloids ca. 25, linear-oblanceolate, 40–50 x 3–4 mm, white. Stamens 50–60, the filaments ca. 30 cm long, red, exserted. Berry oblong, 3–8 x 2–3 cm, longitudinally angled, shiny-red; seeds numerous, 2–3 x 1.8–2.2 mm, black. COLOMBIA. N: (S). ECUADOR. E: Cornejo 1887 (GUAY). G: Cornejo 1 (GUAY). Distribution: Panama to western Ecuador. Ecology: Widespread in the coastal lowland wet forests (Madsen, 1989), from inland to mangroves. Phenology: This species has been collected with flowers from March to May, and with fruits in October. Biological interactions and dispersal: The fruit is most likely eaten by birds and bats, while the small seeds, which are embedded in a fleshy mesocarp, are endozoochorously dispersed. Uses: The fruit pulp is edible. It is said that the hyaline viscid mesocarp is occasionally used as hair gel. IUCN Red list category: This widespread species is persistent in secondary forests and disturbed habitats; therefore, it is here categorized as of Least Concern (LC). Etymology: The epithet means the red-crowned Epiphyllum.

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84. Hylocereus polyrhizus (F. A. C. Weber) Britton & Rose, Cact. 2: 185 (1920). Bas.: Cereus polyrhizus F. A. C. Weber Vernacular names: mulata, pitahaya, pitajaya. Hemiepiphytic and holoepiphytic herbs, to several meters long, sprawling to climbing, profusely and intrincately branched, with numerous aerial roots. Stem trigonous, the segments 50–200 x 3–10 cm; ribs 2–4 cm tall; areoles circular, 2–5 mm wide, sparsely lanate, 2–7 cm apart; spines 2–6 per areole, 2–10 mm long, straight, spreading in various directions. Flowers solitary or few at a time, horizontal, funnel-shaped, 25–31 x 25–30 cm, nocturnal. Bracts 30–50, deltoid to lanceolate, dark green with red margin. Sepaloids ca. 50, green, tinged red at margin. Petaloids oblanceolate, 25–30, 11–14 x 3–5 cm, white with yellow midrib, spreading. Stamens 400–500, the filaments 4–10 cm long, yellow. Berry globose-ellipsoid, 7–12 cm long, red, bearing 25–40 whitering bracts; seeds numerous, 2.6–3 x 1.5–1.7 x 1 mm, black. ECUADOR. G: Cornejo 2 (GUAY). Distribution: Costa Rica to northern Peru. Ecology: In very dry and dry forests. Hylocereus polyrhizus grows in deciduous and semideciduous forest (Madsen, 1989). In Ecuador it is restricted to the dry forests area, occurring from inland to mangroves. Phenology: This species has been collected with flowers in October, and with fruits in February and March. Biological interactions and dispersal: The mature fruit pulp has been observed to be eaten by birds and black wasps called “cubos” (Spanish). It is most likely that monkeys and bats also feed on the mature fruits and disperse the seeds endozoochorously. Uses: The sweet mature pulp is edible (Madsen, 1989). It is said that the grinded stems are used to purify water, those while stirring the water precipitate the suspended particles to make it drinkable. IUCN Red list category: This widespread species is persistent in secondary forests and in disturbed habitats; therefore, it is here categorized as of Least Concern (LC). Etymology: The epithet is formed by the Greek prefix poly (many, numerous) and the Greek word rhiza (root) referring to the numerous roots of this plant. 85. Rhipsalis micrantha (Kunth) DC., Prod. 3: 476 (1828). Bas.: Syn.:

Cactus micranthus Kunth Rhipsalis Tonduzii F. A. C. Weber Rhipsalis Wercklei A. Berger Rhipsalis roseana A. Berger Rhipsalis Rauhiorum Barthlott

Vernacular names: disciplina, pitahayita de monte.

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Holoepiphytic herbs, pendulous 0.5–3 m long, slender. Stems acrotonically jointed, the base of the plant 3–4 angled, the ultimate joints in clusters of one to seven, flattened, more or less succulent, linear to linear-oblong in outline, 5–18 x 0.3–1 cm, entire or crenate at margins; areoles 1–5 cm apart; spines lacking. Flowers lateral, one per areole, funnel-shaped, 6–8 x 6–10 mm, diurnal. Petaloids 5–6, ovate, 4–5 x 2.5–4 mm, white. Stamens ca. 30, the filaments 2–3 mm long. Berry globose, 6–8 mm long, white; seeds numerous, 1.1–1.4 x 0.6–0.7 x 0.4–0.5 mm. ECUADOR. G: Cornejo 8 (GUAY). Distribution: Costa Rica and Venezuela to Peru. Ecology: In dry and moist forests. Biological interactions and dispersal: The whole small fruit is most likely eaten by birds and bats, while the minute seeds, which are embedded in a fleshy mesocarp, are endozoochorously dispersed. IUCN Red list category: This widespread species is persistent in secondary forests and in disturbed habitats; therefore, it is here categorized as of Least Concern (LC). Etymology: The non-distinctive epithet is formed by the Greek prefix micro (small) and the Greek word anthos (flower) referring to the small flowers. Notes: It is said that Rhipsalis micrantha is called “disciplina” (Spanish, meaning discipline) because the elongate fibrous stems are used in lieu of a belt as a punishing strap for children (Valverde, pers. comm.).

CHRYSOBALANACEAE (Cocoa-plum Family) Trees or shrubs. Leaves simple, entire, alternate, stipulate; petioles often 2-glandular. Inflorescence racemose, paniculate or cymose; flowers bracteate and usually bibracteolate. Flowers hermaphrodite, actinomorphic to zygomorfic. Calyx lobes 5, imbricate, often unequal; disk always present, forming a lining to receptacle. Petals 5, inserted on margin of disk, imbricate, deciduous. Stamens 3–100, inserted on margin of disk, in a complete circle or unilateral; staminodes often present, the filaments free. Ovary 1, often villous pubescent; style filiform; stigma 3-lobed or truncate. Fruit a fleshy drupe.

A family represented by two genera in our area: Chrysobalanus and Hirtella. Key to the species

1. Flowers bearing 12–26 stamens, the filaments barely exserted, white; fruits with longitudinal ridges Chrysobalanus icaco 1. Flowers bearing 3 stamens, the filaments conspicuously exserted, distally purple; fruits without longitudinal ridges Hirtella carbonaria

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86. Chrysobalanus icaco L., Sp. Pl. 1: 513 (1753). Plate 18: A. Syn.:

Prunus icaco Labat, nom. illeg. Chrysobalanus pellocarpus G. F. W. Mey. Chrysobalanus purpureus Mill. Chrysobalanus orbicularis Schumach. Chrysobalanus icaco var. pellocarpus (G. F. W. Mey.) Hook f. Chrysobalanus ellipticus Sol. ex Sabine Chrysobalanus icaco var. ellipticus (Sol. ex Sabine) Hook f. Chrysobalanus savannarum Britton Chrysobalanus icaco var. genuinus Stehlé, M. Stehlé & Quentin

Vernacular names: icaco, mata de coca arisca. Small trees or shrubs to 5 m tall. Stipules 1–3 mm long, deciduous. Leaves simple, alternate; blades orbicular to ovate-elliptic, 2–8 x 1.2–6 cm, subcuneate at base, retuse or rounded or with a short blunt acumen not exceeding 2 mm long at apex, glabrous on both surfaces; petioles 2–4 mm long. Inflorescences small terminal and axillary cymules with gray-brown tomentum. Receptacle cupuliform, tomentose on interior and exterior. Calyx green, the lobes 5, rounded to acute at apex. Petals exceding calyx lobes, white, glabrous. Stamens 12–26, the filaments joined for part of their length in small groups, densely hairy. Ovary pilose. Fruit a small fleshy drupe, ovate to obovate, 1.8–5 cm long, pink, rose or red-green to purple at maturity, epicarp smooth with longitudinal ridges, glabrous. COLOMBIA. CH: (E). V: Cuatrecasas 16177 (COL, F). C: (E). N: Cabrera 4098 (CUVC). ECUADOR. E: Cornejo et al. 7231 (GUAY). Distribution: From southeast United States (Florida) through Central America, to southern Brazil, and in the West Indies, and on the west coast through Colombia to northwestern Ecuador; also in Africa. Ecology: Chrysobalanus icaco is a mangrove fringe species, tolerant of hyposaline environments. The species inhabits in the wet forest and rainforest life zones, where sometimes is forming pure stands called “icacales”. Uses: Although the fruit pulp is edible, the species apparently is not cultivated in our area, as is it in the Venezuelan Guayana (Prance, 1998). The fruits of Chrysobalanus icaco are sold in Bogota markets for preserves called “dulce de Icaco”. It is said that the leaves of C. icaco are processed as those of “coca” (Erythroxylum coca) for illicit purposes in illegal drug traffic, to be sold as fake coca instead of-, or mixed in with- the latter species, hence the second vernacular name that means “wild coca bush”. On that way, the leaves of C. icaco, which are harvested from the native populations, are used only to increase the profit per sales volume, but apparently with no any drug effect at all as those of the true coca leaves. C. icaco is cultivated as an ornamental in Colombia. IUCN Red list category: This widespread species of healthy populations is herein categorized as of Least Concern (LC).

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Etymology: “icaco”, one of its vernacular names, was proposed as the generic name for the species (Plumier, 1703). However, Linnaeus (1753) used it as the specific epithet, apparently because it was considered to be of a barbarian origin. 87. Hirtella carbonaria Little, Jour. Wash. Acad. Sci. 38: 88 (1948). Plate 18: B. Vernacular names: carboncillo, carbonero, garrapato. Trees or shrubs to 10 m tall and 20 cm dbh (or more?). Stipules linear, 2–4 mm long. Leaves simple, alternate; blades coriaceous, elliptic to ovate-elliptic, 4.5–9.5 x 2.5–4.5 cm, rounded at base, acuminate at apex, the lateral veins 6–9 per side, sparsely hirsutulous beneath. Inflorescences terminal and axillary panicles, 3–9 cm long; bracts and bracteoles ovate, 1–3 mm long, hirsutulous, persistent; solitary glands on a stalk, 1–2 mm long, arising from junction of pedicel and branch or from pedicel. Flowers 4–5 mm long (without stamens length); receptacle campanulate. Calyx lobes rounded to acute. Petals 5, white, glabrous. Stamens 3, unilateral, with 3 short filamentous staminodes opposite them, the filaments far exceeding calyx lobes, distally purple. Style hirsute on lower portion. Ovary inserted near mouth of receptacle, tomentose. Fruit ellipsoid, ca. 2.5 x 1.7 cm, red when immature, turning black at maturity, glabrous. COLOMBIA. CH: Killip & Cuatrecasas 39130 (COL). V: Cuadros 954 (MO). C: (E). N: (S). ECUADOR. E: Bonifaz & Cornejo 4014 (AAU, GB, GUAY, QCNE). Distribution: Pacific coast of Colombia to northwestern Ecuador. Ecology: Hirtella carbonaria is a mangrove fringe species, tolerant of hyposaline environments, that inhabit in wet forest and rainforest life zones. The species mainly occurs landwards in mangroves, and in the adjacent swampy areas dominated by Mora oleifera, which are known as natales. Phenology: This species has been collected in flowers and fruits throughout the year. The flowers seem to be ephemeral; those are fragile to the touch and produce nectar during nights. Biological interactions: Its pollinators are not known, but may include moths. The fleshy drupes suggest dispersal by birds or bats. Uses: The wood is used in making charcoal. The sweet fruit pulp and the seed endosperm are edible. IUCN Red list category: Due to factors as a narrow range of distribution of less than 2,000 km2 and a loss of habitat, this species is categorized as Vulnerable (VU B2abiii). Etymology: The epithet is a Latinized word from “carbonero” (Spanish), one of its vernacular names that is derived from carbon (=charcoal) referring to the main use of this plant. Note: The type was collected in 1943 by E. L. Little Jr., a North American botanist, in San Lorenzo (REMACAM), Province of Esmeraldas, Ecuador.

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CLUSIACEAE (Mangosteen Family) Trees, shrubs (some epiphytic), lianas, perennial or annual herbs, usually with white, yellow or bright colored latex. Leaves opposite, whorled or alternate, simple, entire; exstipulate. Inflorescences terminal or axillary, cymose, or flowers solitary. Flowers actinomorphic. Sepals 2–14. Petals free, usually 3–6, decussate, imbricate or convolute. Stamens usually numerous. Ovary compound, superior, of usually 3–5 united carpels; styles 1 or as many as the carpels; stigmas ± expanded, often smooth and sticky. Fruit usually a capsule, sometimes a berry or a drupe. Key to the species 1. Leaf blades 5–13 cm wide; inflorescences 3–7 cm long, the peduncle very short; fruits subglobose, 2.5–3.5 cm wide Clusia leptanthera 1. Leaf blades 3–7 cm wide; inflorescences 10–20 cm long, the peduncle elongate; fruits oblong or narrowlycylindric, ca. 1 cm wide

Clusia fructiangusta

88. Clusia fructiangusta Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 33 (1950). Plate 18: C. Vernacular name: matapalo. Hemiepiphytic shrubs or small trees. Latex white. Leaves simple, opposite, sessile; blades coriaceous, obovate, 4–14 x 3–7 cm, cuneate at base, broadly obtuse to suborbicular and shortly acuminate at apex, entire, the lateral veins 12–20 per side, ascendent, glabrous. Inflorescence a terminal panicle, 10–20 cm long, the peduncle elongate. Flowers 1.2–2.5 cm diam. Sepals ca. 1 cm long. Petals pink. Fruit oblong or narrowly-cylindric, ca. 2.5 x 1 cm. COLOMBIA. V: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 7258 (GUAY). Distribution: Nicaragua to Ecuador. Ecology: In wet forests and rainforests. Etymology: The epithet is formed by the Latin words fructus (fruit) and angusta (narrow), that refers to the distinctive narrow fruits. IUCN Red list category: Least Concern (LC). Note: The type was collected in 1944 by José Cuatrecasas (1903–1996), a prolific Spanish botanist, at sea level on the Pacific coast in the mouth of Cajambre River, Isla del Guayabal, Department of Valle del Cauca, Colombia.

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89. Clusia leptanthera Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 52 (1950). Plate 18: D. Vernacular name: matapalo. Hemiepiphytic shrubs or small trees. Branches angulate, subpendulous. Leaves simple, opposite; blade coriaceous, obovate to shortly oblanceolate, 10–21 x 5–13 cm, cuneate to narrowlycuneate at base, rounded to truncate at apex, entire, the lateral veins 15–30 per side, oblique, brochydodromous, the intersecondaries present, glabrous; petioles 10–15 mm long, winged. Inflorescence a terminal raceme, 2–3-flowered, 3–7 cm long, the peduncle ca. 2 cm long. Flowers 4–6 cm diam., porcelain-white. Sepals ca. 1–1.2 cm long. Petals 3–4 x 1.5–2 cm, palepink to white. Fruit subglobose, 3–4 x 2.5–3.5 cm, smooth, light-yellow to white at maturity. COLOMBIA. CH: (E). V: Gentry & Juncosa 40707 (MO). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8147 (GUAY). Distribution: Eastern Panama to northwestern Ecuador. Ecology: This species has been mostly collected in the adjacent terra firme coastal forests, frequently along river margins, in the wet forest and rainforest life zones. Etymology: The epithet is fomed by the Greek prefix lepto (narrow) and the Latin word anthera (anther) referring to the narrow anthers that called attention of Cuatrecasas when describing this species (B. Hammel, pers. comm.). IUCN Red list category: Least Concern (LC). Note: The type was collected in 1945 by José Cuatrecasas near sea level in Buenaventura Bay, Department of Valle del Cauca, Colombia. COMBRETACEAE (Indian Almond Family) Trees, shrubs or lianas, spiny or unarmed. Leaves opposite, whorled or spiral, often crowded at tips of branches, simple, entire; petioles often with a pair of glands; exstipulate. Inflorescences axillary and/or terminal; flowers bisexual or both bisexual and staminate in same inflorescence, regular, sessile, in axillary or extra-axillary panicles or globose heads, usually bracteate. Hypanthium more or less two winged, surrounding and adnate to ovary, the upper part deeply or shallowly cupuliform, produced beyond ovary. Calyx lobes 4 or 5 or poorly developed and almost absent. Petals 4 or 5 or none, alternating with sepals, deciduous. Stamens usually twice as many as sepals, the filaments inserted inside upper hypanthium. Intrastaminal disk usually surrounding base of style. Ovary inferior or half inferior, 1-locular; style simple; stigma punctiform or capitate. Fruit indehiscent, often winged or ridged; seed 1.

A family represented in mangroves by two well-known genera: Conocarpus and Laguncularia.

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Key to the species and varieties 1. Leaves opposite; inflorescence paniculate; mature fruits narrowly-obovate, green 2. Inflorescences, floral bracts, calyx, corolla and fruits velutinous to densely pilose Laguncularia racemosa var. racemosa 2. Inflorescences, floral bracts, calyx, corolla and fruits glabrous Laguncularia racemosa var. glabriflora 1. Leaves spiral; inflorescence a raceme of heads; mature fruits in globose heads, orange to brown Conocarpus erectus 90. Conocarpus erectus L., Sp. Pl. 1: 176 (1753). Plate 19: A–E. Syn.: Conocarpus procumbens L. Conocarpus erectus var. procumbens DC. Conocarpus acutifolius Humb. & Bonpl. ex Roem. & Schult. Terminalia erecta Baill. Terminalia erecta var. procumbens (L.) M. Gómez Vernacular names: botoncillo, jabalí, jelí, jelí de semilla, mangle botón, mangle jelí, mangle salado, palo salado. Small trees or shrubs (1.5–)2–8 m tall, to 40 cm dbh. Terminal branchlets angulate. Leaves simple, alternate; blades thinly-coriaceous (chartaceous when dry), elliptic to lanceolate or oblong, 4–10 x 1.5–4 cm, cuneate to attenuate and decurrent onto petiole at base, usually acute and apiculate or acuminate at apex, entire to repand, the domatia in axils of secondary veins and submarginal beneath, the lateral veins 8–14 per side, brochydodromous, the tertiary veins reticulate; petioles short, winged, with a pair of lateral nectary glands. Inflorescence a raceme of heads, axillary and terminal, to 10(–15) cm long, the rachis tomentose to tomentulose, glabrescent; heads at anthesis 3–5 mm diam., bearing ca. 30–50 tiny flowers, usually bisexual. Floral bracts narrowly-triangular, 1–1.5 mm long. Calyx 2/3 gamosepalous, 1–1.5 mm long, the sepals 5, the lobes dentate, acute at apex. Petals absent. Stamens 5(–10), 1–2.5 mm long, alternisepalous, exserted, the filaments white, the anthers 0.2–0.4 mm long, cream-yellowish. Ovary pubescent; style 1.5–2 mm long, white; stigma inconspicuous. Infructescence compound, ± globose, 9–15 x 9–12 mm, suberect to pendulous, orange to brown; individual fruits subwoody, 4–5 x 4–8 mm, lateraly winged, concave within, indehiscent, brown; seed 1. COLOMBIA. CH: Ordóñez et al. 67 (NY). V: Killip 11695 (NY). C: (S). N: Murphy & Parra 710 (MO, NY). ECUADOR. E: Cornejo et al. 7228 (GUAY, QCNE). M: Cornejo 7891 (GUAY). SE: (S). G: Cornejo & Bonifaz 7044 (AAU, GB, GUAY, QCNE). O: Asplund 15868 (NY). GAL: Triviño 61 (GUAY). PERU. T: Smith 10103 (MO). P: (E).

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Distribution: Widely distributed along the tropical shores of West Africa and the Caribbean region, and from southeastern United States (southern Florida) to Brazil, as well as along the west coast of the Americas from Mexico to northwestern Peru, also in the Galapagos islands. Ecology: Conocarpus erectus grows in more consolidate muds, often as a mangrove fringe species, and also in salt marshes and sandy beaches or even on volcanic soils (in the Galapagos islands) to the adjacent terra firme forests, where it is often found to 20 meters above sea level, and exceptionally to 200 m in Colombia (Stace, 2010). This species grows in open sunny places, and occurs in dry thorn scrub, very dry, dry, moist, wet forest, and rainforest life zones. It is noteworthy that individuals which grow on sandy beaches have the tendency to present thicker-coriaceous leaf blades. C. erectus occasionally form woods called “Jelicales” (Rimbach 55, WIS[MAD]), that is a term derived from “jelí”, one of its vernacular names. Phenology: This species has been collected in flowers and fruits throughout the year. Biological interactions: The flowers are fragrant and secrete nectar during the day. Those have been observed to be occasionally visited by several species of small insects as beetles, Sarcophagidae flies and Halictidae bees. The pair of petiolar glands secrete nectar, never salt as it is often mistakenly cited in the literature, but no visitors have been observed. Dispersal: The fruits and seeds are water dispersed. Uses: The bark stores high concentrations of tannins, sometimes over 25 %, and has been reported to be used to tan and medicine (Bernal and Correa, 1990). The leaves contain tannins also (Little, 1969). The wood is incorruptible, compact, heavy and hard, reported to be good for building small ships (AcostaSolís, 1944), and constructions. Nowadays, most individuals usually have trunks of less than 20 cm in diameter, however. The wood burns slowly, and also produces an excellent firewood and charcoal. This species is cultivated as an ornamental in the Galapagos islands. IUCN Red list category: Due to factors as the widespread range of distribution and the common presence of healthy populations in secondary habitats, it is agreed its categorization as of Least Concern (LC) (Polidoro et al., 2010). Etymology: The generic name is formed by the Greek words konos (cone, strobilus) and karpos (fruit) referring to the strobili-like infructescences. The non-distinctive specific epithet is a Latin word, that refers to the upright stance of trees. Notes: Conocarpus erectus is here regarded as a facultative mangrove because has a wider range of ecological distribution. The species was discovered by Sloane in Jamaica during the 17th century. Puerto Jelí, a port located in the Province of El Oro, southwestern Ecuador, owes its name to “Jelí”, one of the vernacular names. Conocarpus erectus forma sericeous (DC.) Stace has been proposed to designate the commonly cultivated ornamental plant of this species. That differs from the typical forma by the leaves that retain a sericeous pubescence until they are old. Extreme examples of C. erectus f. sericeous are very distinctive with densely sericeous leaves of a silvery appearance. C. erectus f. sericeous is more common in the West Indies and in Florida, its natural occurrence in South America is rather rare (Stace, 2007). That forma sericeous is an introduced taxa, that is occasionally cultivated as an ornamental in the city of Guayaquil, and eventually may occur in the mangroves of the region.

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91. Laguncularia racemosa (L.) C. F. Gaertn., Suppl. Carp. 209 (1807). For basyonyms and synonyms see under varieties. Vernacular names: mangle amarillo, mangle blanco, mangle bobo, mangle hembra, palo salado, manoa, manglillo. Trees or shrubs 1.5–10 m tall, to ca. 25 cm dbh. Secondary (sub)superficial roots with many pneumatophores, these thicker-tipped, erect to 10 cm tall, lenticellate. Terminal branchlets with bark which can be pealed out as long narrow strips. Leaves opposite or decussate, arranged along branches; blades coriaceous or slightly suculent, elliptic, oblong or inconspicuously lanceolate, 7–15 x 3.5–7 cm, obtuse or rounded to cordate, and sometimes asymmetric at base, obtuse and inconspicuosly apiculate to retuse at apex, ± entire, with scattered submarginal pits beneath; the lateral veins 9–16(–18) per side; petioles 1–2.5 cm long, reddish, with a distal pair of yellowish to greenish, swollen-like nectary glands adaxially. Inflorescences axillary and terminal racemes, the peduncle 3.8–15 cm long; floral bracts ± orbicular, apiculate to shortly-acuminate (ca. 1.5 mm) at apex, deciduous before or after anthesis. Flowers sessile, greenish in lower half, fragrant. Calyx 2/3 gamosepalous, the sepals 5, the lobes ca. 1–1.2 mm long, rounded to deltoid at apex, green to yellow. Corolla dialipetalous, the petals 5, alternate with sepals, suborbicular, 1–1.5 mm, white. Stamens 10, ca. 1 mm long, arranged in two whorls of 5 each, the inner whorl epipetal, the outer whorl episepal. Ovary inferior; nectary disk gland yellow, arranged on hypantium and surrounding style base; hypanthium ca. 2 mm long, greenish; style ca. 1.3 mm long; stigma capitate, centraly depressed. Fruits indehiscent, narrowly-obovate, 1.4–2 x 0.8–1 cm, longitudinally ribbed, green, crowned by persistent parts of calyx. Distribution: See under varieties. Ecology: See under varieties. Biological interactions: The flowers are visited by at least fifteen species of insects. Among those, the most common pollinators are honey bees, Apis mellifera (Apidae). Secondary visitors are diurnal Syrphid flies as Palpada vinetorum (Plate E: B), P. albifrons, P. aemula, P. sp. 1 and Copestylum pallens vel aff., several unidentified species of flesh fly (Sarcophagidae) and green bottle fly (Calliphoridae), beetles as Olla roatanensis and Paraneda sp. (Coccinellidae), butterflies as Dione juno (Nymphalidae), wasps (Vespidae), small bees (Halictidae) and ants. A video about most of these insect species visiting flowers of L. racemosa is available in this link: https://www.youtube.com/watch?v=XFTcwTR1XlQ The branchlets and leaves have been reported to be eaten by goats (Madsen et al., 2001). Snails of Littorina varia (Littorinidae) as well as caterpillars occasionally feed on the leaves of this species. Dispersal: The fruits and seeds are water dispersed. The root protrudes out of the fruit wall several days after falling on water and/or stranding on mud/ground.

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Uses: Due to the high concentrations (22 %) of tannins (Bernal and Correa, 1990), the wood has been used to tan. The straight trunks, known as “puntal” or “puntalillo” (Spanish), are mainly used as stilts, called “patas” (=legs, Spanish), to build coastal stilt houses on the estuarine water level. The highly resistant wood is less attacked by marine bivalvia drillers (Teredinidae, M. Cruz pers. comm.), known as shipworms or “polilla de agua” (Spanish). The wood is used for canoes, fences, houses, nets, ship frames known as ribs, and charcoal (Madsen et al., 2001). This species can concentrate heavy metals as: Cupper (Cu), lead (Pb), and zinc (Zn) (Villamil, 2010). The shallow secondary horizontal roots when raised over the ground suggest the loss of soils by erosive action of waves or a lixiviating process. Discussion: The paired petiolar glands are extra-floral nectaries and do not excrete salt as has been frequently stated (e.g., Gentry, 1993). Note: Laguncularia racemosa is unique among the Combretaceae trees in the area in having opposite leaves and short pneumatophores, the latter are usually covered by the ground. 91a. Laguncularia racemosa var. racemosa. Plate 20: A, C, E, F. Bas.: Conocarpus racemosus L. Syn.: Laguncularia obovata Miq. Laguncularia racemosa f. longifolia J. F. Macbr. Inflorescences, floral bracts, calyx, corolla and fruits velutinous to densely pilose. COLOMBIA. CH: Gentry & Juncosa 40944 (MO). V: Gentry & Juncosa 40544 (MO). C: (S). N: Espinoza 2958 (NY). ECUADOR. E: Cornejo & Bonifaz 6982 (GB, GUAY, QCA, QCNE). M: Cornejo 1717 (GUAY, QCNE). SE: (E). G: Cornejo & Castro 247 (GUAY, QCNE). O: Bonifaz & Cornejo 3739 (GUAY). GAL: Schimpff 15 (MO). PERU. T: Simpson & Schunke 597 (NY, US). P: (S). Distribution: Widely distributed along the tropical shores of West Africa and the Caribbean region, and from southeastern United States (southern Florida) to Brazil, as well as along the west coast of the Americas from Mexico to northwestern Peru, also in the Galapagos islands. Ecology: Semirestricted to mangroves and occasionally also growing on sandy beaches, in dry thorn scrub, very dry, dry, moist, wet forests, and rainforests. It is often sympatric with Rhizophora and Avicennia, but sometimes forming pure stands (Stace, 2010). Undoubtedly, L. racemosa is a true or major mangrove. L. racemosa is often found growing and replacing on somewhat more consolidates but still muddy soils where Rhizophora have been felled. This is a fast growing tree, flowering and fruiting from before the second year (Little, 1969). Phenology: This variety has been collected in fruits throughout the year. IUCN Red list category: According to the Registro Oficial No. 148 issued on March 16 1993, Laguncularia racemosa has been categorized as an endangered species in Ecuador (INEFAN/GEF, 1998). However, due to factors as the widespread range of distribution, the commonly presence of healthy populations, those usually comprised by abundant individuals, the high rate of seed viability, and the abundant seedlings and juveniles observed in the field, it is agreed its categorization as of Least Concern (LC) (Polidoro et al., 2010).

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Etymology: The generic name is derived from the Latin word laguncula (a flask or bottle), that refers to the shape of fruits of this species. The epithet refers to the racemose inflorescences. Discussions: In the original publication (Macbride, 1930), and subsequently in taxonomical treatments as Flora of Ecuador and Flora Neotropica (Stace, 2007, 2010), Weberbauer 7527 (F) has been cited as the type collection of Laguncularia racemosa f. longifolia (that is a synonym of L. racemosa). In F herbarium such collection has not been found, but Weberbauer 7627 is the only collection available from the same type locality, with the same collector and date (Peru: Tumbes, coastal plain between Tumbes and Zarumilla, Tumbes River near the sea, 30–31 Jan 1927). That Weberbauer 7627 has a label hadwritten as “type” by Macbride himself (Hensold, pers. comm.). Therefore, it is concluded that Weberbauer 7527 is a typewriting error and that Weberbauer 7627 is the correct type of Laguncularia racemosa f. longifolia. The populations observed in San Cristobal (Galapagos islands), differ from the continental populations of this variety by having mature fruits with oblong-obovate shape (vs. narrowly-obovate), these with a rather smooth surface (vs. several times ribbed). Therefore, molecular work including populations of this species from the Galapagos islands is recommended. 91b. Laguncularia racemosa var. glabriflora (C. Presl) Stace, Fl. Ecuador 81: 11 (2007). Plate 20: B, D. Bas.: Laguncularia glabriflora C. Presl, Rel. Haenk. 2: 22 (1831). Syn.: Laguncularia parviflora Shuttlew., nom. nud. Inflorescences, floral bracts, calyx, corolla and fruits glabrous. Full data of the studied collections of Laguncularia racemosa var. glabriflora are presented herein because that variety was previously reported to be known in Ecuador by the type only (Stace, 2007), and was not known to occur in Peru: ECUADOR. Guayas: Guayaquil, 24 Jan 1955 (fl, y fr), Asplund 15227 (NY); W of town, 18 Oct 1955 (fl), Asplund 18189 (NY, US). Outskirts of Guayaquil at road to Salinas, 20 Jan 1985 (fr), Harling & Andersson 21026 (QCA). Prope Guayaquil, in littoribus limosis regionis trop., Feb 1926 (fl), Mille 68 (NY, QCA). 12 km S of Guayaquil near Canal and Puerto Nuevo, 28 Jul 1977 (fr), Iltis et al. E-622 (WIS). REMCH, sector San Alejandro, 79°40’W 2°27’S, 22 Feb 1991 (fr), Cerón 13341 (QCNE). Hda Josefina, Apr 1985 (fl), Elao s.n. (GUAY). Filipo, Oct 1973 (fl), Bonilla s.n. (GUAY). Estero Mongón, 1 Nov 1974 (fl), Bonilla s.n. (GUAY). Isla Puná, vic. of Puná Nueva, 79°55’W 2°44’S, 3 Nov 1987 (fl), Madsen 64098 (GUAY, QCNE); 2 Mar 1989 (fl), Madsen & Eriksen 85870 (MO). El Oro: Arch. Jambelí, Las Huacas, 80°09’W 3°20’S, 6 Feb 1998 (fr), Cornejo & Bonifaz 5981 (GUAY, MO). Arch. Jambelí, Puerto Pitahaya, Nov-Dec 1978 (fl), Albert de Escobar 906-A (QCA); 80°05’W 3°25’S, 6 Feb 1998 (fl, y fr), Cornejo & Bonifaz 5975 (GUAY, MO). PERU. Tumbes: Prov. Tumbes, Estuaries near mouth of Río Tumbes, 80°31’W 3°30’S, 21 Mar 1985 (fl, fr), Smith 10107 (MO); Weberbauer 7621 (MO). Prov. Zarumilla, Estuaries near Puerto Pizarro, 80°28’W 3°30’S, 18 Mar 1985 (fl), Smith 10090 (MO). Puerto Pizarro, 13 Oct 1971 (fl), Ferreyra s.n. (MO); 20 Oct 1971 (fl), Cerrate 5035 (MO). Manglar Cherre, frente a Punta Malpelo, 17 Dec 1977 (fr), Ferreyra 18962 (MO).

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Distribution: On the Pacific coast of America this variety is known only in southwestern Ecuador and adjacent northwestern Peru; and in the Atlantic coast, it has been collected in southwestern Florida, Mexico, and scattered throughout the West Indies. Ecology: In dry thorn scrub, very dry, and dry forests. IUCN Red list category: Due to factors as the reduced range of distribution of this variety in the Pacific coast of America, where it is known as to inhabit in less than 1,000 km2, and a loss of habitat in the region, mainly caused by the shrimp farm construction impact and the incorrect expansion of some cities (e.g., Guayaquil, Machala) over mangroves, destroying significant areas where this variety occurs, it is here propose its categorization as Vulnerable (VU B2abiii). Etymology: The varietal epithet refers to the absence of hairs in the flowers. Note: The type of Laguncularia glabriflora, the variety basyonym, was collected (s.n.) in October 1790 by the Czech explorer and botanist Thaddäus Haenke (1761–1816) on the shore of rio Daule, Guayaquil, Province of Guayas, Ecuador. That seems to be the first botanical collection gathered from the mangroves in the Pacific coast of South America. Remains of the type population of this white mangrove variety are still present at Parque Historico Guayaquil. Discussion: In the protologue, Laguncularia glabriflora was described as having red flowers. However, all of the cited collections have white flowers. Furthermore, I have never seen red flowers for this variety in the numerous field trips and it is considered to be an error as previously suggested (Exell, 1958). Although the remaining morphological differences are not good characters to recognize L. glabriflora as a distinct species, Exell (1958) also suggested that this taxon may represent a variety under L. racemosa, but “without any well-marked geographical distribution”. The new status for Laguncularia racemosa var. glabriflora (Stace, 2007) took almost half of a century to be proposed. CYCLANTHACEAE (Panama hat Family) Perennial herbs, terrestrial or epiphytic; acaulescent or provided with ± short stems. Leaves spirally arranged or distichous, petiolate; blades simple, usually bifid, more seldom flabelliform-parted or entire, uni-, bi- or tricostate. Inflorescence an axillary, peduncled spadix, subtended by 2–11 spathes. Flowers monoecious; staminate and pistillate flowers in spirally arranged groups. Staminate flowers mostly with a cupulate, marginally ± lobed perianth; stamens numerous. Pistillate flowers sometimes entirely coalescent into cycles, or only partly connate with each other or even entirely free. Fruit a fleshy syncarp of united or separate berries; seeds numerous, mostly small, flat or terete.

This Neotropical family is represented in mangroves by one genus comprising two species. Key to the species

1. Stigmas linear seen from above, distinctly uncinate; tepals acute, usually connate in fruit; fruits yellow to orange at maturity Sphaeradenia acutitepala 1. Stigmas lanceolate to elliptic seen from above, not uncinate or only slightly so; tepals subacute or notched, free; mature fruits bright-red at maturity Sphaeradenia killipii

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92. Sphaeradenia acutitepala Harling, Acta Horti Berg. 18(1): 354, fig. 95 D–E (1958). Syn.: Sphaeradenia carrilloana Grayum & Hammel Holoepiphytic climber herbs, growing in clumps. Leaf blades narrowly-bifid, 60–140 cm long, divided 57–20 % of their length, the segments lanceolate to narrowly-triangular, 3.5–12 cm broad; petioles 25–80 cm long. Inflorescences axillary, erect; peduncle 7–27 cm long. Spathes 2 to 4, coriaceous, lanceolate. Spadix cylindrical, 3–8 x 1–1.5 cm, cream; tepals acute; stigmas linear seen from above, distinctly uncinate. Infructescences erect, cylindrical, 4.5–10 x 1.5–2 cm, turning softer, yellow to orange and juicy at maturity. COLOMBIA. CH: Betancur 7953 (COL). V: Gentry & Juncosa 40711 (MO). Distribution: Southern Nicaragua to western Colombia. Ecology: In wet forests and rainforests. Phenology: This species has been collected in fruits from November to May. Biological interactions: It is most likely that the fleshy infructescences are eaten and the seeds endozoochorously dispersed by bats, birds, and monkeys (Croat, 1978; Lobova et al., 2009). Use: This is a potentially ornamental species. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Latin words acutus (acute, pointed) and tepala (tepals) referring to the acute tepals. 93. Sphaeradenia killipii (Standl.) Harl., Acti Horti Berg.17(1): 3 (1954). Plate 21: A. Bas.: Carludovica killipii Standl. Holoepiphytic climber herbs, growing in dense clumps. Stem stout, to more than 1 m long, ca. 3 cm diam. Leaf blades narrowly-bifid, 60–110 cm long, divided 70–32 % of their length, the segments oblong to oblong-lanceolate, 5–11 cm broad; petioles 40–80 cm long. Inflorescences axillary, erect; peduncle 10–17 cm long. Spathes 2 to 4, stiff, coriaceous, lanceolate. Spadix rather narrowly-cylindrical, 4–8 x 1–1.5 cm, yellowish-red; tepals subacute or notched; stigmas lanceolate to elliptic seen from above, not uncinate or only slightly so. Infructescences erect, cylindrical, 7–9 x 2.5–3 cm, turning softer, brigth-red and juicy at maturity. COLOMBIA. V: Killip 34975 (COL, F, the type collection). C: (E). N: (S). ECUADOR. E: Sparre 18352 (S). Distribution: Western Colombia to northwestern Ecuador. Ecology: In wet forests and rainforests.

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Phenology: This species has been collected in flowers in April and August, and in fruits in August and September. Biological interactions: It is most likely that the fleshy infructescences are eaten and the seeds endozoochorously dispersed by bats, birds, and monkeys (Croat, 1978; Lobova et al., 2009). Use: This is a potentially ornamental species. IUCN Red list category: Although it is persistent in secondary forests, due to factors as an increasing loss of habitat in near future, it is herein categorized as Near Threatened (NT). Etymology: The epithet honors Ellsworth P. Killip (1890–1968), a North American botanist who in 1939 collected the type in the mangroves of Buenaventura, Department of Valle del Cauca, Colombia. CYPERACEAE (Sedge Family) Perennial or annual herbs, often appearing as grasses, rarely vine-like. Culms triangular to round in cross section, winged or 4–6-angled, rarely compressed. Leaves basal or basal and cauline, tristichous or less often distichous; blades mostly linear. Inflorescence terminal or terminal and axillary, with few to many spikelets; involucral bracts often present; glumes 1-many per spikelet, usually spirally arranged or distichous. Stamens 1–3. Ovary superior; style 2- or 3-branched or entire. Fruit a nutlet, nutlet-like or a nutlet protected by a utricle.

The species of this family occurs infrequently in the interior of hyposaline disturbed mangroves.

They are more commonly found at fringes and behind mangroves. Key to the species 1. Plants leafless or apparently leafless, forming dense colonies 2. Culms to 3 m tall, blunty trigonous

Schoenoplectus californicus

2. Culms to 1 m tall, terete 3. Culms transversely septate Eleocharis elegans 3. Culms not septate Eleocharis mutata 1. Plants leafy, not forming dense colonies 4. Leaves ± subwhorled, arranged at base or at base and apex of plants, the culms leafless 5. Leaves arranged at base of plants only 6. Leaves not stiff, the leaf sheaths light-green to light-brown at base; spikelets 2.5–6 mm long Fimbristylis dichotoma 6. Leaves stiff, the leaf sheaths blackish-brown at base; spikelets 6–15 mm long Fimbristylis spadicea 5. Leaves arranged at base and apex of plants 7. Spikelets densely arranged, 4–7 mm long

Cyperus ligularis

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7. Spikelets loosely arranged, 10–20 mm long Cyperus odoratus 4. Leaves polystichous, arranged separatedly along culms 8. Stem 5-angled Fuirena umbellata 8. Stem 3-angled 9. Inflorescences narrow, racemose 10. Nutlet 2.6–2.9 mm wide Scleria eggersiana 10. Nutlet 2–2.5 mm wide Scleria melaleuca 9. Inflorescences open, corymbose Rhynchospora corymbosa 94. Cyperus ligularis L., Syst. Nat. ed. 10, 867 (1759). Plate 21: B. Syn.: Mariscus ligularis (L.) Urb. Mariscus rufus Kunth Terrestrial herbs, perennial, often forming dense clumps. Culms to 1 m tall, transversely triangular, without nodes. Leaves arranged only at base; blades 5–12 mm wide, thick, glaucous, usually transverse-lineolate. Bracts long and leaf-like. Spikes in a compound inflorescence, short and very dense; spikelets 4–7 mm long, reddish-brown; scales coriaceous, obtuse, striate, closely appressed. Nutlet narrowly-obovate, 1.5 mm long, sharply trigonous, darkbrown, with a lightly papillose surface. COLOMBIA. CH: Gentry 40933 (COL, MO). V: (E). C: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 1888 (GUAY). Distribution: Southeastern United States (Florida) to Argentina, also in West Africa and Madagascar. Ecology: In dry, moist, and wet forests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the ligule. 95. Cyperus odoratus L., Sp. Pl. 46 (1753). Plate 21: C. Syn.:

Diclidium odoratum (L.) Shrad. ex Nees Pycreus odoratus (L.) Urb. Torulinium odoratum (L.) S.S. Hooper Cyperus macrocephalus Liebm. Cyperus eggersii Boeck. Cyperus engelmannii Steud. Cyperus ferax Rich. Cyperus granadinus Liebm. Cyperus oerstedii Liebm. Terrestrial herbs, annual or short-lived perennial. Culms stout, 0.5–1 m tall, transversely triangular, without nodes. Leaves arranged only at base; blades 5–12 mm wide, shorter than

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culm. Bracts large, leaf-like. Inflorescence compound, the loose heads on rays sometimes 20 cm long; spikelets linear, subterete, 10–20 mm long, yellowish or brownish; scales 2–3 mm long, striate; raquilla strongly winged, the wings becoming corky and enlarged in age, and the spikelet breaking up into single segments. Nutlet oblong, 1.5–2.5 mm long, frequently curved, dark-brown, finely reticulate or papillose. COLOMBIA. CH: Gentry & Juncosa 40933-A (MO). V: (E). C: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 2612 (GUAY, WIS). Distribution: United States to Argentina. Ecology: In dry, moist and wet forests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means having a sweet smell. 96. Eleocharis elegans (Kunth) Roem. & Schult., Syst. Veg. 2: 150 (1817). Bas.: Scirpus elegans Kunth Syn.: Scirpus geniculatus L.

Eleocharis geniculata (L.) Roem. & Schult. Terrestrial herbs. Culms 0.3–1 m tall, transversely septate, terete, leafless. Spikelet solitary, terminal, 1–3 x 0.5–0.9 cm, erect, white to brown, the scales spirally imbricate, acutish; bristles equaling or longer than achene. Nutlet ellipsoid, biconvex to trigonous, granular, yellow-brown; tubercle usually flattened, dark-brown, elongate.

COLOMBIA. CH: (E). V: (E). C: (E). ECUADOR. G: Bonifaz & Cornejo 3907 (GUAY). PERU. T: (E). Distribution: Central Mexico to northern Argentina, and in the West Indies. Ecology: In dry, moist, wet forests, and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means elegant. 97. Eleocharis mutata (L.) Roem. & Schult., Syst. Veg. 2: 155 (1817). Bas.: Scirpus mutatus L. Terrestrial herbs, caespitose, forming dense colonies. Culms 0.4–1 m x 3–6 mm, not septate, terete, leafless. Spikelet solitary, terminal, 1.5–4 x 0.4–0.6 cm, thick, erect, creamish to brown, the scales spirally imbricate, obtuse, greenish; bristles longer than achene; style 3-cleft. Nutlet dark-brown, faintly cancellate; tubercle broader than nutlet apex.

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COLOMBIA. V: Killip & Cuatrecasas 38826 (COL). N: (E). ECUADOR. E: Bonifaz & Cornejo 4006 (GUAY). O: Laegaard 70175 (AAU). PERU. T: (E). Distribution: United States to Bolivia and Paraguay. Ecology: In dry, moist, and wet forests. IUCN Red list category: Least Concern (LC). 98. Fimbristylis dichotoma (L.) Vahl, Enum. Pl. 2: 287 (1805). Bas.: Syn.:

Scirpus dichotomus L. Scirpus diphyllus Retz. Fimbristylis annua var. diphylla (Retz.) Kük. Fimbristylis polymorpha Boeck. Terrestrial herbs, caespitose, annual or perennial. Culms to 0.5(–0.8) m tall, leafy below. Leaf blades narrow, shorter than culms, light-green to light-brown at base. Inflorescence loose and open; spikelets numerous, 2.5–6 mm long, acute, reddish-brown; style 2-fid, prominently fimbriate. Achenes ovoid, biconvex, white or stramineous, with about 10 rows of horizontallyelongated cells on each face and about 10 longitudinally ribs formed by cell margins.

COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 2643 (GUAY). Distribution: Old and New World, subtropics, and tropics. Ecology: In dry, moist, and wet forests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the dichotomous inflorescences. 99. Fimbristylis spadicea (L.) Vahl, Enum. Pl. 2: 294 (1805). Bas.: Syn.:

Scirpus spadiceus L. Scirpus dominguensis Pers. Fimbristylis spadicea f. dominguensis (Pers) Kük. Fimbristylis speciosa Rohde ex Spreng. Terrestrial herbs, caespitose, perennial. Culms to 2 m tall, leafy below. Leaf blades narrow, stiff, indurate and castaneous or blackish at base. Spikelets few or numerous, 6–15 mm long, the scales obtuse, glabrous or puberulous, apiculate, dark-brown with paler veins imbedded in tissue. Nutlet obovate, biconvex, with deep-pitted horizontally elongated cells, and narrowed at base to a minute purplish annulus.

COLOMBIA. CH: Juncosa 2529 (MO, NY). V: Killip & Cuatrecasas 38616 (COL). C: (E). ECUADOR. E: Lægaard 71776 (AAU, QCNE). G: Asplund 15264 (NY).

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Distribution: Southern United States to Brazil. Ecology: In dry, moist, and wet forests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that seems to refer to the spadix-like appearance of the spikelets. 100. Fuirena umbellata Rottb., Descr. Icon. Rar. Pl. 70 (1773). Syn.: Scirpus umbellatus (Rottb.) Kuntze Fuirena paniculata L. f. Fuirena tereticulmis C. Presl Vernacular name: cortadera. Terrestrial herbs, caespitose, perennial. Culms leafy, 0.5–1.5 m tall, 5-angled, stout, sometimes pilose above, often bulbous at base. Leaf blades 7–16 x 0.8–2.5 cm, usually scabrous. Spikelets oblong, 6–10 mm long, clustered, the scales spirally imbricate, pubescent, aristate, with spreading or recurved awns; bristles ovate, membranous, not stipitate. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6984 (GUAY). Distribution: Mexico to Paraguay. Ecology: In dry, moist, wet forests, and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the umbellate inflorescences. 101. Rhynchospora corymbosa (L.) Britton, Trans. New York Acad. Sci. 11: 84 (1892). Syn.:

Scirpus corymbosus L. Rhynchospora aurea Vahl. Rynchospora corymbosa (L.) Britton Dichromena corymbosa (L.) J. F. Macbr. Terrestrial herbs, caespitose, perennial. Culms scabrous above, 3-angled, with several nodes, stout, 0.6–2.5 m tall. Leaves arranged along culm; blades 1–2 cm wide. Several dense corymbs forming an interrupted inflorescence 20–40 cm long; spikelets very numerous, 6–7 mm long, containing a single achene. Nutlet narrowly-obovate, 2–3 mm long, faintly reticulate, darkbrown, the corky thickened beak equaling achene; bristles 6, exceeding achene, upwardly barbed.

COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6970 (GUAY, WIS). Distribution: Mexico to Paraguay, and Argentina. Ecology: In dry, moist, wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the corymbose inflorescences.

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102. Schoenoplectus californicus (C. A. Mey) Soják, Čas. Nár. Mus., Odd. Přír. 140 (3–4): 127 (1972). Bas.: Syn.:

Elytrospermum californicum C. A. Mey Scirpus californicus (C. A. Mey) Steud Scirpus riparius J. Presl & C. Presl Scirpus tatora Kunth

Vernacular name: totora. Terrestrial herbs, caespitose, perennial. Culms closely tufted, to 3 m x 2–4 mm thick near inflorescence, blunty trigonous; apparently leafless. Leaves few, basal, submersed. Inflorescence 4–12 cm long, decompound with a number of usually drooping branches, altogether with 50 to 150 spikelets; spikelets laceolate-ovoid, 6–11 mm long, of 30–50 flowers; scales ovate to obovate, ca. 3 mm long, dark-brown; bristles 2–4, subligulate, reddish-brown; styles mostly bifid. Nutlet obovate, ca. 2 mm long, apiculate, brown. COLOMBIA. N: (S). ECUADOR. E: Bonifaz & Cornejo 4007 (GUAY). Distribution: Southern United States to Bolivia. Ecology: In moist and wet forests. This species mostly occur in Andean lakes. Use: The stems are used to elaborate handicrafts. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the North American state of California, where the type was collected. 103. Scleria eggersiana Boeck, Beitr. Cyper. 2: 41 (1890). Vernacular name: cortadera. Terrestrial herbs. Culms coarse 1–2 m tall, triangular, suberect, nearly smooth. Leaves cauline, distantly disposed; blades coriaceous, rigid, flat or somewhat plicate, ca. 30 (or longer) x 1–2.5 cm, scabrous, especially on veins and margins; sheats 3-winged, scabrous on angles, otherwise glabrous. Inflorescence paniculate, narrow, ca. 30 cm long; hypogynium-margin densely brown-ciliate, cup-shape, 3-lobed. Nutlet globose or subglobose, 2.6–2.9 mm wide, white, shining, smooth, terminated by conic, pale, more or less deciduous style-base. COLOMBIA. N: (E). ECUADOR. E: Dodson et al. 14568 (MO, NY, QCNE). Distribution: Southern Mexico to Ecuador. Ecology: In moist and wet forests. IUCN Red list category: Least Concern (LC). Etymology: The epithet honors Baron Henrik F. A. Von Eggers (1844–1903), a Danish soldier and botanist who collected the type in Cuba.

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Note: Scleria eggersiana closely resembles S. mitis Bergius. Koyama relegates Scleria eggersiana to subspecies rank of S. mitis, but further study must be carried out to prove Koyama’s circumscription correct (Camelbeke and Goetghebeur, 1998). 104. Scleria melaleuca Rchb. ex Schltdl. & Cham., Linnaea 6: 29 (1831). Syn.:

Scleria pterota Presl Scleria pterota var. melaleuca (Rchb. ex Schltdl. & Cham.) Uittien Scleria boliviana Palla ex Buchtien Scleria pratensis Lindl. ex Nees

Vernacular names: cortadera, wam bio tape (Cayapa). Terrestrial herbs. Culms 0.3–1 mm tall, triangular, rather stout, nearly glabrous. Leaves cauline, distantly disposed; blades 15–40 x 0.7–1.5 cm, scabrous on margins. Panicles axillary, pyramidal, 5–15 cm long, sparsely branched, green or brownish, the bracts inconspicuous; hypogynum depressed, 3-lobed, the lobes broad, rounded, ciliate or glabrous. Nutlet depressed-globose, 2–2.5 mm wide, smooth, dark-brown to black. COLOMBIA. CH: (E). V: (E). C: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 2676 (GUAY). Distribution: Southern Mexico to Argentina, also in tropical Africa. Ecology: In moist, wet forests, and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a combination of two Greek words: melas (black) and leukos (white), probably referring to the color variability of the fruits. ERICACEAE (Blueberry Family) Terrestrial or epiphytic shrubs, subshrubs, lianas or perennial herbs. Stipules absent. Leaves alternate, rarely opposite, verticillate, or lacking and then replaced by bract-like scales, simple, petiolate. Inflorescences axillary or rarely terminal, racemose, paniculate, fasciculate or flowers solitary; pedicels usually bibracteolate, the bracteoles persistent. Flowers mostly bisexual, actinomorphic or slightly zygomorphic, mostly (3–)5(–7)-merous. Calyx gamosepalous. Corolla commonly gamopetalous, cylindric, campanulate or urceolate, terete or angled. Stamens (6–)10(–14), in 2 whorls, usually twice as many as petals or rarely just as many, equaling the corolla in overall length or ½–1/3 the corolla length. Pistil single; ovary superior or inferior, 4–5(–10)-carpellate. Fruit a loculicidal or septicidal capsule, berry, or drupe, with a usually persistent calyx. Seeds small, usually numerous. This is a predominantly Andean family. The previously reported occurrence of several species of this family in mangroves swamps (Luteyn, 1983) has been rather unnoticed.

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Key to the species 1. Leaf blades 0.7–2 cm long, the petioles 1–2 mm long; inflorescences 1(–2) flowers, axillary Sphyrospermum buxifolium 1. Leaf blades 4–30 cm long, the petioles 3–15(–22) mm long; inflorescences racemose, terminal and/or axillary 2. Leaf blades with pinnate lateral veins 3. Leaf blades with several glands beneath, truncate to subcordate at base; pedicels 1.4–2 cm long, with apical bracteoles; flowers with corollas 2–2.3 cm long, red at anthesis

Macleania smithiana

3. Leaf blades many glandular-punctate beneath, cuneate to obtuse at base; pedicels 3–4 cm long, with basal bracteoles; flowers with corollas 2.7–3.1 cm long, white at anthesis (turning pink to red at postanthesis)

Thibaudia pachypoda

2. Leaf blades with plinerved lateral veins 4. Leaf blades glandular-punctate beneath; inflorescences 2–4-flowered, the rachis 0.2–0.6 cm long Cavendishia urophylla 4. Leaf blades not glandular-punctate beneath; inflorescences many-flowered (except occasionally Anthopterus wardii), the rachis 0.7–17(–20) cm long 5. Inflorescences axillary, 2–11-flowered; floral bracts inconspicuous, less than 1 cm long; corollas obovoid-urceolate or subglobose 6. Leaf blades obtuse to subcordate at base; petioles 3–5 mm long; inflorescences with rachises 1.2–4 cm long; calyces not articulate with pedicels; calyces and corollas winged Anthopterus wardii 6. Leaf blades cuneate at base; petioles 10–15 mm long; inflorescences with rachises 0.7–1.2 cm long; calyces articulate with pedicels; calyces and corollas not winged

Psammisia occidentalis

5. Inflorescences terminal, 15–35(–52)-flowered; floral bracts conspicuous, (1.5–)2–4(–7) cm long; corollas cylindric to conical-cylindric 7. Leaf blades obtuse to often cordate at base; flowers with corollas (1.5–)1.8–3 cm long Cavendishia callista 7. Leaf blades cuneate to attenuate at base; flowers with corollas 0.7–1.1 cm long Cavendishia palustris 105. Anthopterus wardii Ball, Hooker’s Icon. Pl. 15: 1465 (1884). Plate 21: D. Syn.: Thibaudia wardii (Ball) Hoerold Hemiepiphytic or holoepiphytic shrubby lianas. Leaves loosely spiral; blades stiff, coriaceous, elliptic to ovate-elliptic, 4–10 x 1.5–3.5 cm, obtuse to subcordate at base, acuminate at apex, the lateral veins 3–5(–7)-plinerved from at or near base, puberulous proximally along midrib above, glabrous elsewhere; petioles 3–5 mm long. Inflorescences axillary, racemose, 2–10-flowered,

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the rachis 1.2–4 cm long; floral bracts oblong-lanceolate, ca. 3 mm long; pedicels 1–1.5 cm long, the bracteoles similar to floral brats, but smaller. Flowers not articulate with pedicels. Calyx 5-winged, 0.8–1 cm long, orange to red, glabrous. Corolla 5-lobed, 5-winged, obovoidurceolate, 1–1.3 x 1–1.2 cm, orange to reddish-orange to the throat where turning green and distally white at lobes. Berry light purple to blue at maturity; seeds numerous, minute. COLOMBIA. V: Killip 34955 (NY). Distribution: Panama to western Ecuador. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The fleshy fruit is most likely eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. Use: This species is occasionally cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet honors Mr. Richard Ward (dates unknown), who collected the type near sea level in the forests near Buenaventura, Department of Valle del Cauca, Colombia. 106. Cavendishia callista Donn. Sm., Bot. Gaz. 20(1): 5, t. 2 (1895). Plate 22: A. Syn.:

Cavendishia longiflora Donn. Sm. Cavendishia amalfiensis Mansf. Cavendishia duidae A. C. Sm. Cavendishia bullata A. C. Sm. & Standl. Cavendishia hispida A. C. Sm. Cavendishia gleasoniana A. C. Sm. Cavendishia praestans A. C. Sm. Cavendishia phelpsiae Camp Cavendishia salicifolia Maguire, Steyermark & Luteyn

Vernacular name: matapalo. Hemiepiphytic or holoepiphytic shrubs. Leaves alternate; blades coriaceous to thickly hard coriaceous, ovate, lanceolate, elliptic, oblong or rarely obovate, (4.5–)7–23(–35) x 2.5– 10(–13) cm, obtuse to often cordate at base, acute to long-acuminate at apex, glabrous to moderately short-pubescent above, glabrous, glabrate, or densely hispid beneath especially along nerves, the lateral veins (3–)5-plinerved from at or near base; petioles 6–13(–22) mm long. Inflorescences terminal, racemose, at anthesis viscid, 15–35(–52)-flowered, encircled at base by bracts, the rachis 4–12(–20) cm long; floral bracts oblong, obovate, or oblanceolate, (1.5–)2–4(–6) x (0.7–)1–2.5(–3.5) cm, rounded or acute at apex, pink to deep rose; pedicels 0.6–1.5(–2) cm long, the bracteoles basal. Flowers articulate with pedicels. Calyx 5–9(–11)

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mm long, viscid, pinkish-green at anthesis, glabrous or infrequently sparsely to densely pilosehispid, the lobes triangular, glandular-callose, the callus tissue covering lobes completely or restricted to distal 1/3–1/2. Corolla 5-lobed, cylindric, narrowed to throat, viscid, (1.5–)1.8–3 cm long, white to grayish pearl-white. Mature berry not seen; seeds numerous, minute. COLOMBIA. V: Cuatrecasas 16020 (F, GH, VALLE). Distribution: Guatemala to Amazonia of Brazil. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The cylindric corollas suggest pollination by hummingbirds. The fleshy fruit is most likely eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet callista refers to the beautiful nymph Callisto of Greek mythology. Note: A beautiful, showy, and morphologically very variable species (Luteyn, 1983). 107. Cavendishia palustris A. C. Sm., Amer. J. Bot. 27: 543 (1940). Hemiepiphytic or holoepiphytic shrubby lianas. Leaves alternate; blades coriaceous, lanceelliptic to ovate-lanceolate, 6.5–10(–15) x 2.5–5 cm, cuneate to attenuate at base, ± blunty acuminate at apex, the lateral veins 3–5-plinerved from at or near base, glabrous on both sides; petioles 4–9 mm long. Inflorescences terminal, racemose, 20–35-flowered, the rachis 13–17 cm long; floral bracts somewhat cucullate, oblong, 2.5–3(–7) x ca. 3 cm, rounded at apex, green; pedicels 1–1.8 cm long, the bracteoles basal, fimbriate at margins. Flowers articulate with pedicels. Calyx ca. 4 mm long, pale yellowish-green to green, glabrous; lobes deltate, marginally with one large oblong to crescent-shaped callose gland on each side often obscuring the lobe itself. Corolla 5-lobed, cylindric to conical-cylindric, basally swollen, 0.7–1.1 cm long, red or whitish at base and purplish-violet distally. Berry blue-black to black at maturity; seeds numerous, minute. COLOMBIA. V: Killip 34983 (COL, NY, US, the type collection). Distribution: Western Colombia and northwestern Ecuador. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The fleshy fruit is most likely eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the mangrove swamp habitat where the type species was collected in 1939 by the North American botanist Ellsworth P. Killip (1890–1968), in Buenaventura Bay, Department of Valle del Cauca, Colombia.

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108. Cavendishia urophylla A. C. Sm., J. Arnold Arbor. 27: 110, f. 3 (1946). Plate 22: B. Vernacular name: quereme. Hemiepiphytic or holoepiphytic shrubby lianas. Leaves alternate; blades stiff, brittle, coriaceous, ovate to elliptic, 4–8 x 1.7–3 cm, cuneate to obtuse at base, caudate at apex, 3-plinerved from base but often inconspicuous, glossy above, glandular-punctate and glabrous on both sides; petioles 3–6 mm long. Inflorescences terminal, shortly racemose, 2–4-flowered, the rachis 2–6 mm long; floral bracts elliptic or ovate-oblong, 0.8–2 x 1.3–1.5 cm, rounded at apex, pink or pinkish-white; pedicels 5–7 mm long, the bracteoles basal. Flowers articulate with pedicels. Calyx 8–12 mm long, white, glabrous; lobes triangular, marginally with callose glands on each side. Corolla 5–lobed, cylindric, ca. 3 cm long, white to cream-pink, green at base. Berry yellowish at maturity; seeds numerous, minute. COLOMBIA. V: Gentry et al. 53415 (MO, NY). Distribution: Western Colombia. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The fleshy fruit is most likely eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. IUCN Red list category: Due to the narrow range of distribution and a loss of habitat this species is categorized as Vulnerable (VU B1abiii). Etymology: The epithet is formed by the Greek prefix uro (tail) and the Greek word phyllo (a leaf blade), referring to the caudate apex of leaf blades. 109. Macleania smithiana Luteyn, Fl. Ecuador 54: 131 (1996). Plate 22: C. Vernacular name: femu-piu-tapé (Cayapa). Hemiepiphytic or holoepiphytic lianas. Leaves loosely spiral; blades stiff, coriaceous, ovate to shortly-lanceolate, 7–12 x 4–7 cm, truncate to subcordate at base, with several glands beneath, ± obtuse at apex, the lateral veins pinnate, 2–5 per side, strongly ascendent, glabrous on both sides; petioles ca. 8 mm long. Inflorescences terminal and axillary, short loose racemes, 2–6-flowered; pedicels 1.4–2 cm long, the bracteoles apical, glabrous. Flowers articulate with pedicels. Calyx 5-angled, 1–1.3 cm long, red, glabrous. Corolla 5-lobed, cylindric, 2–2.3 x 0.5 cm, bright-red, densely white-pilose at throat within. Berry yellowish at maturity, glabrous; seeds creamish to light green; seeds numerous, minute. COLOMBIA. CH: Gentry 53862 (NY). V: Killip 34994 (NY). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8201 (GUAY).

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Distribution: Southeastern Panama to northwestern Ecuador. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The bright-red cylindric corollas strongly suggest pollination by hummingbirds. The fleshy fruit is most likely eaten by birds and bats, while the minute seeds, embedded in a sticky and juicy mesocarp, are endozoochorously dispersed. Uses: There are no uses reported for this species from mangroves. In the adjacent terra firme forests of northwestern Ecuador, the Cayapa Indians boil the leaves and the decoction is used to bath swelled tissue, relieving bites of the poisonous Equis snake (Bothrops sp.; Luteyn, 1996). IUCN Red list category: Least Concern (LC). Etymology: The epithet honors A. C. Smith (1906–1999), a North American taxonomist on Ericaceae. Note: The populations from the mangroves of northwestern Ecuador and adjacent southwestern Colombia differ from the species description provided in Flora of Ecuador (Luteyn, 1996) by the bracteoles with glabrous margins. 110. Psammisia occidentalis A. C. Sm., Amer. J. Bot. 27: 542 (1940). Hemiepiphytic or holoepiphytic shrubby lianas. Leaves alternate; blades coriaceous, elliptic, 17–30 x 4.5–11 cm, cuneate at base, acuminate at apex, the lateral veins 5–7-plinerved from at or near base, glabrous on both sides; petioles 10–15 mm long. Inflorescences axillary, shortly-racemose, 8–11-flowered, the rachis 7–12 mm long; pedicels 7–11 mm long, the bracteoles basal. Flowers articulate with pedicels. Calyx 3–6 mm long, red, glabrous. Corolla 5-lobed, subgobose, ca. 4 mm long, yellow or greenish with green lobes, glabrous. Mature fruits not seen; seeds numerous, minute. COLOMBIA. CH: Killip & Cuatrecasas 39110 (A, US). Distribution: Western Colombia. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The fleshy fruit is most likely eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. IUCN Red list category: Data Deficient (DD). Etymology: The epithet refers to the distribution of the species west of the Andes. Note: This is a provisional name. A revision of the genus Psammisia is needed (Pedraza, pers. comm.). 111. Sphyrospermum buxifolium Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 4, pl. 8 (1835). Plate 22: D. Holoepiphytic shrublets. Leaves alternate; blades thick, coriaceous, suborbicular to shortlylanceolate, 0.7–2 x 0.8–2 cm, subcordate at base, subrounded or obtuse at apex, the lateral

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veins 3–5-plinerved from base, glabrous on both sides; petioles 1–2 mm long. Flowers 1(– 2) per axil, usually extending well beyond the leaves; pedicels slender, 1–2 cm long, the bracteoles nearly basal, minute, ciliate. Flowers not articulate with pedicels. Calyx 2–2.8 mm long, green or red, densely pilose to glabrate. Corolla 4-lobed, cylindric-urceolate, ca. 5.5 mm long, terete, white, sparsely pilose to glabrous. Berry whitish and translucent or with a violet or lavender tinge to purple at maturity, pilose or glabrous; seeds numerous, minute. COLOMBIA. V: Killip 34963 (NY). Distribution: Nicaragua to Bolivia. Ecology: In wet forests and rainforests. Biological interactions and dispersal: It has been suggested that bats may feed on the fruits of Sphyrospermum buxifolium and disperse the seeds endozoochorously (Lobova et al., 2009). IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the resemblance of the leaf blades to those of the genus Buxus (Buxaceae). Thibaudia pachypoda A. C. Sm., Jour. Arnold Arb. 27: 97 (1946). Plate 23: A. Holoepiphytic shrubs. Leaves loosely spiral; blades stiff, coriaceous, brittle, ovate-elliptic to elliptic, 8–23 x 3–8.5 cm, cuneate to obtuse at base, acuminate at apex, the lateral veins pinnate, ca. 4–5 per side, strongly ascendent, the blade glandular-punctate beneath, glabrous on both sides; petioles 3–10 mm long. Inflorescences axillary, shortly-racemose, 2–5-flowered, the rachis 3–10 mm long; floral bracts oblong, ca. 3 mm long; pedicels 3–4 cm long, the bracteoles basal. Flowers articulate with pedicels. Calyx 8–10 mm long, pink to rose-red, glabrous. Corolla 5-lobed, cylindric, 2.7–3.1 cm long, white at anthesis, turning pink to red at post-anthesis, glabrous. Berry pink or whitish; seeds numerous, minute. COLOMBIA. V: (E). Distribution: Western Colombia. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The cylindric corollas suggest pollination by hummingbirds. The fleshy fruit is most likely eaten by birds and bats, while the minute seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. IUCN Red list category: Data Deficient (DD). Note: I have not seen collections of this species from mangroves. However, Thibaudia pachypoda is included here on account of its coastal occurrence at mangroves edges, and because it has been reported to occur in mangrove swamps (Luteyn, 1983).

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FABACEAE (Bean Family) Trees, shrubs, herbs or vines. Leaves alternate, usually compound. Flowers actinomorphic (Mimosoideae) or zygomorphic (Caesalpinioideae, Faboideae), bisexual, 5-merous, solitary or in compound inflorescences, axillary or terminal. Sepals free or united, 4 or 5, rarely 1, then spathaceous. Petals free or united, 5. Stamens 5 to many, the filaments free or united, the staminodes sometimes present. Pistil 1, the ovary superior, 1-locular, 1- to many-ovulate. Fruit a legume, dehiscent or indehiscent, 1- to manyseeded, 2-valved, but sometimes modified as a drupe, samara, follicle, or loment; seeds commonly with a coriaceous testa.

In our area this family is represented by five genera and five species. Key to the species

1. Lianas; fruits entirely flattened 2. Leaves unifoliolate, the folioles ovate to shortly lanceolate; inflorescences 2–8 cm long, the flowers zygomorfic, loosely arranged in short racemes

Dalbergia brownei

2. Leaves bipinnate, the folioles oblong; inflorescences 25–50 cm long, the flowers actinomorphic, densely arranged in spikes Entada polystachya 1. Shrubs to tall trees; fruits convex without or partially flattened 3. Leaves paripinnate; inflorescences dense spikes, the flowers actinomorphic, white; fruits large, 10– Mora oleifera

15 cm wide

3. Leaves imparipinnate; inflorescences loose panicles or racemes, the flowers zygomorfic, yellow or purple-magenta; fruits small, 1.5–5 cm wide 4. Red latex present; flowers yellow; fruits ± orbicular and strongly asymmetric, winged, veined Pterocarpus officinalis 4. Red latex absent; flowers purple-magenta; fruits cylindric-moniliform, smooth and richly lenticellate Muellera chocoensis 112. Dalbergia brownei (Jacq.) Shinz, Bull. Herb. Boissier 6(9): 731 (1898). Plate 23: B. Bas.: Amerimnon brownei Jacq. Vernacular name: adorote. Woody lianas or apparently riparian shrubs, reaching over 5 cm diam. Leaves 1-foliolate; blades chartaceous to coriaceous, ovate to shortly lanceolate, 3–8 x 1.4–6 cm, widely obtuse to cordate at base, acuminate-apiculate at apex, glabrous above, inconspicuosly appressedpuberulous beneath, the lateral veins 7–10 per side; petioles 7–25 mm long. Inflorescence a

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raceme or panicle, 2–8 cm long; pedicels 1.5–4 mm long. Sepals 4–5 mm long, green. Petals ca. 1 cm long, white, glabrous. Stamens 5–7 mm long, the filaments fused over its lower half. Ovary oblong, ca. 6 x 1.5 mm, complanate laterally; style ca. 2.5 mm long. Fruit oblong, 2–5 x 0.7–1.3 cm, strongly complanate, attenuate or cuneate at base, obtuse and apiculate at apex, irregularly sinuate and occasionally constricted, green, glabrous. COLOMBIA. CH: Gentry & Juncosa 40934 (MO). V: (E). C: (E). N: Ramírez 5200 (NY). ECUADOR. E: Cornejo & Bonifaz 6933 (AAU, GUAY, K). Distribution: Southern Mexico to Venezuela, northwestern Ecuador, and in the West Indies. Ecology: Dalbergia brownei is a usually common weed in open mangrove areas and in the adjacent estuarine riverbanks, often inhabiting the edge of the upper tidal level in the wet forest and rainforest life zones. The species often presents a shrub-like appearance, which seems to be an adaptation against the water currents where the tidal level rises higher than usual. D. brownei also grows in the natales; it is often associated with Pterocarpus officinalis. Phenology: It has been collected in flower throughout the year. Biological interactions: The flowers are fragrant during the day, suggesting insect pollination. The petals shed on the water are eaten by fish. The plants occasionally are grazed by cattle. Dispersal: The flattened seeds have been reported to float on water, “owing to surface tension, but once sunk they will not resurface”. They are also wind dispersed to a lesser extent (Croat, 1978). IUCN Red list category: This widespread species, commonly represented by numerous healthy individuals, is herein categorized as of Least Concern (LC). Etymology: The epithet honors Patrick Browne (ca. 1720–1790), an Irish physician, historian and botanist who was the first to report its presence from Jamaica as “the shrubby Amerimnon, with simple alternate leaves” (Browne, 1756). Note: The cited collection from Ecuador represents the first record for the country (Cornejo, 2003). 113. Entada polystachya (L.) DC., Prodr. 2: 425 (1825). Plate 23: C. Bas.: Syn.:

Mimosa polystachya L. Adenopodia polystachya (L.) J. Dixon ex Croat Entada polyphylla Benth. Entada plumierii Spreng. Entadopsis polyphylla (Benth.) Britton Entadopsis polystachya (L.) Britton

Vernacular name: bejuco de agua. Woody lianas. Leaves bipinnate, with 2–6 pairs of pinnae, the pinnae usually 6–15 cm long, the leaflets opposite, 6–8 pairs per pinna, oblong, 2–4 x 0.8–1.8 cm, oblique at base, obtuse or emarginate at apex; petioles mostly 4–8 cm long. Inflorescence a large, terminal raceme of slender spikes, 25–50 cm long. Calyx shallowly-cupulate, 0.5–1 mm long, truncate or briefly

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dentate. Petals 5, free, linear, ca. 2 mm long, white, glabrous. Stamens 10, free, somewhat exceding petals. Ovary glabrous. Legume large, oblong, 25–40 x 5–8 cm, strongly complanate, with margins thicker and persistent, transversely articulated, the segments 1-seeded, corkypapirous, shiny-green when inmature, light-brown at maturity. ECUADOR. G: Cornejo & Bonifaz 7043 (GUAY). Distribution: Mexico to Bolivia. Ecology: In dry, moist and wet forests. This species occurs from the forest inland, fresh water wetlands and along rivers to the hyposaline mangroves (Bonifaz, 1988). Biological interactions: The flowers have been observed to be visited by insects. Dispersal: The flattened, light, corky seeds are wind dispersed. Use: It may be cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the many spikes that the inflorescence bears. 114. Mora oleifera (Triana ex Hemsl.) Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 45 (1925). Plate 24: A–D. Bas.: Dimorphandra oleifera Triana ex Hemsl. Syn.: Dimorphandra megistosperma Pittier Mora megistosperma (Pittier) Britton & Rose Vernacular names: alcornoque, mangle nato, nato, nato rojo. Trees to 35 m tall and to 2 m dbh. Buttresses large and acute, which extend horizontally plank-like to several meters long. Terminal branches lenticellate. Leaves alternate, 4-foliolate; leaflets opposite in two pairs, coriaceous, lanceolate, ± asymmetric, 10–20 x 4–8 cm, obtuse to slighly subcordate at base, acuminate at apex, entire, lustrous above, glabrous, the lateral veins 6–12 per side; petioles 2–5 cm long; petiolules 2–3 mm long. Inflorescence a dense axillary spike, erect, ca. 8–12 cm long, growing on a peduncle ca. 3–4 cm long. Flowers sessile. Calyx 5-gamosepalous, campanulate, 3–4 mm long, the lobes rounded, ca. 1 mm long, ciliate at margins. Petals oblong, 4–6 x 2 mm, widely obtuse to truncate at apex, creamish-white, ciliate at margins. Stamens 5, the staminodes 5, 7–8 mm long, exserted. Ovary densely pilose; style 8–10 mm long, exceding stamens. Fruit a pendulous legume, 13–27(–40) x 10–15 cm, brown, lenticellate, glabrous, dehiscent at maturity; seeds 1–2(–3), 10–15 x 10–12 cm, smooth, dark-brown. COLOMBIA. CH: Acevedo-Rodríguez et al. 6859 (NY). V: Killip 5217 (NY). C: Mayor 1 (CUVC). N: Romero-Castañeda 3214 (NY). ECUADOR. E: Bonifaz & Cornejo 4010 (AAU, GB, GUAY, K, QCA). Distribution: Pacific coast, from Costa Rica to northwestern Ecuador. The southernmost limit of

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distribution occurs in northwestern Ecuador; it does not reach the dry forests at southern Ecuador, as previously mapped by West (1977). Ecology: Mora oleifera is a large tree that grows in periodically flooded places, from coastal riverbanks and fresh water swampy areas, seaward to the mangroves. It inhabits moist, wet forest, and rainforest life zones. The species can extend its populations, forming pure stands that reach the outermost seaward border, in very salty water environments (Acosta-Solís, 1966; pers. obs.). The pure stands are known as natales, a term derived from nato, one of its vernacular names. On very soft, compactless soils, the mature and taller individuals of M. oleifera have been frequently observed falling down slowly, because of the absence of morphologically more specialized aerial roots and/or absence of mutual support provided by the neighbouring tree branches or crowns. In mangroves, it is usually associated with Rhizophora spp., Laguncularia racemosa, Avicennia germinans, Pterocarpus officinalis, and Euterpe oleracea. The fruit of M. oleifera is utterly unmistakable, as the seed has been reported to be the largest of any dicot (Hemsley, 1885; Pittier cited in Little, 1969; Gentry, 1993). Biological interactions: The flowers are commonly visited by bees, flies, beetles and butterflies as Aphrissa boisduvalii (Pieridae). Occasionally, the underside of the bark is inhabited by scorpions, which sometimes climb onto the branches and enter the grooved fruits, possibly looking for insects to eat. The seeds are palatable to several crab species, including some edible ones such as the Red crab (Ucides occidentalis) and Blue crab (Cardisoma crassum), which are sold in local markets. It cannot be discarded that some extinct megafauna could feed on the fruits and seeds of this species. Dispersal: At maturity, the fruit longitudinally dehices into two valves, and releases one or two seeds, which are often carried away by the water. After the seeds leave the water, germination is achieved in approximately two days. If the seeds return to the water, they sink and die (Cangá, pers. com.). Uses: The wood is highly compact, hard, but not very smooth, brown-reddish colored, reputed to be more incorruptible, heavier, and stronger than the wood of all mangroves in the area. It is used to build ceilings, houses, fences, and beds, but not often furniture because of its heaviness (Acosta-Solís, 1944). It is also more resistant to sea water than other mangrove woods, and is used to build piers, wharfs, and columns known as “pilares” (Spanish). In addition, the nato wood is highly tolerant to the strong vibration of a wooden boat motor, so it is the preferred wood for building the frame (known as “espejos” in Spanish, in northwestern Ecuador) which supports the boat motor.

During his field observations made between 1939 and 1940 in the Province of Esmeraldas, Acosta-

Solís (1944) noted: “Once nato trees were growing at the Río Esmeraldas mouth. Under river water there are trunks which are still in perfect condition; this indicates the wood high resistence and quality. In the north of the Province of Esmeraldas, I have seen ‘puntales’ [underground support columns] and pieces of nato wood which have been underground for more than 200 years.” The wood, bark, leaves, and fruits contain high concentrations of tannins (Cárdenas and Salinas, 2007). IUCN Red list category: This species has been categorized as Vulnerable (VU) (Polidoro et al., 2010). However, it is necessary to highlight that due to excessive timber use, approximately 60 % of Mora oleifera

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population has been felled in Colombia (Cárdenas and Salinas, 2007); and that a similar, but apparently not formally documented situation occurred to the populations of M. oleifera in Ecuador. Therefore, according to IUCN Red list criteria it is followed its categorization as Endangered (EN A2acd) for Colombia (Cárdenas and Salinas, 2007). Furthermore, it is here proposed the same IUCN categorization (EN A2acd) for the remaining populations of M. oleifera in the whole range of its distribution, or at least in northwestern South America. Selecting seeds for reforestation, field observations and recommendations: For reforestation purposes the seeds should be planted rather than the seedlings. This choice is supported by the fact that the seeds have a high viability rate (ca. 90 %). In contrast, the planted seedlings have a high mortality rate (ca. 90%). The best seeds that must be selected to be planted are those collected from single-seeded fruits found at river mouths, because those are more resistant, and largest in size. Once planted, the cotyledons feed the seedlings until the third month; meanwhile the young plant grows, reaching between 30 to 150 cm tall. The cotyledons then start to dry and subsequently, the plant begins growing at a slower pace. Approximately 1 of each 1,000 seedlings has been observed to be albino (Cangá, field obs.). As M. oleifera needs plenty of water, it is highly recommended for the seeds to be planted near water currents or streams. Otherwise channels should be opened in the ground in order to provide a close enough supply of running water, thus avoiding a much delayed growth rate. Etymology: The epithet is formed by the derived Latin word oleum (oil) and the Latin suffix fer (carrying), that pressumably refers to the presence of oils in the seeds. It has not been found such reports in the literature, however. Note: The type of Dimorphandra oleifera, which is housed in Kew, was collected by Sutton Hayes (No. 69) in the swamps of the Rio Grande, Panama. Discussion: This species has been frequently cited as Mora megistosperma. However, the earliest valid name is Dimorphandra oleifera (1885, against Dimorphandra megistosperma, 1915). Therefore, the subsequent combination Mora oleifera (1925) is the correct name for the species. 115. Muellera chocoensis M. Sousa, Acta Bot. Mexicana 100:23 (2012). Plate 23: D. Vernacular names: bagatá, barbasco, barbasquillo. Trees to 6 m tall and to 20 cm dbh. Stipules inconspicuous. Leaves alternate, imparipinnate, 5–7-foliolate, 10–30 cm long; lateral leaflets opposite, chartaceous, lanceolate, 2–12 x 2–4 cm, obtuse to cuneate at base, acuminate at apex, more or less symmetric, entire, the lateral veins 10–15 per side, the tertiaries inconspicuous, glabrous; petioles 4–7 cm long; petiolules ca. 5 mm long. Inflorescence a short ramiflorous raceme, suberect, ca. 3 cm long, the floral bracts absent, the flowers arranged in pairs. Calyx ca. 5 mm long, purplish-brown. Corolla papilionaceous, the petals 5, ca. 12 mm long, purple-magenta. Stamens 10, ca. 9–11 mm long, the lower half fused forming a tube. Fruit dry, indehiscent, subwoody, cylindric-moniliform, 2–16 x ca. 2 cm, glabrous; seeds 1–5. COLOMBIA. CH: Fuchs et al. 21740 (COL, NY, the type). V: Killip 5236 (NY, US). C: (E). N: RomeroCastañeda 5299 (COL). ECUADOR. E: Cornejo & Cangá 8212 (GUAY).

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Distribution: Pacific coast of Colombia and northwestern Ecuador. Ecology: This is a riparian and mangrove species that often inhabit flooded areas from fresh- to brackishwaters ecosystems. It grows in the wet forest and rainforest life zones. Uses: The seeds have a powerful ictiotoxic activity; they are grinded and mixed in water to fish. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the Choco region, where the species occurs (Sousa and Sotuyo, 2012). Note: Muellera chocoensis has been regarded as a minor mangrove (West, 1956). It has been commonly reported in the literature as the widespread Muellera frutescens or M. moniliformis. According to a recent monograph in the genus (Sousa and Sotuyo, 2012), M. chocoensis has been described as a new species restricted to the Pacific coast of Colombia and NW Ecuador and M. frutescens does not occur in our area. 116. Pterocarpus officinalis Jacq., Select. Stirp. Amer. Hist. 283, pl. 183, f. 92 (1763). Plate 25: A–D. Syn.:

Pterocarpus draco L. Moutouchi suberosa Aubl. Pterocarpus suberosus (Aubl.) Pers. Pterocarpus crispatus DC. Pterocarpus belizensis Standl.

Vernacular names: bambudo, bambulo, sangre de gallo, suela, suela de playa. Trees to 30 m tall and to 50 cm dbh. Trunks often curved, twisted or canaliculated. Buttresses prominent, plank-like, sinuously curved, extending horizontally on ground. Latex red, bitter. Stipules 2, squamiform. Leaves alternate, imparipinnate, 5–9-foliolate, 15–40 cm long; lateral leaflets subopposite to alternate, coriaceous, ovate to lanceolate, oblong or elliptic, 3–15 x 1.5–6 cm, cuneate to truncate at base, acuminate at apex, asymmetric except the terminal, entire, the lateral veins 7–11 per side, the tertiaries reticulate, glabrous; petioles 3–6 cm long; petiolules ca. 5 mm long. Inflorescence an axillary panicle, suberect, 5–30 cm long; floral bracts linear, ca. 3 mm long. Calyx ca. 6 mm long, green. Corolla papilionaceous, fragrant, the petals 5, ca. 12 mm long, yellow. Stamens 10, ca. 7 mm long. Fruit dry, indehiscent, subwoody, ± orbicular and strongly asymmetric, 2.8–5 x 3–5 cm, winged, somewhat flattened on one side and irregularly convex on the other, veined, dark-brown, glabrous; seed 1. COLOMBIA. CH: Gentry 53858 (MO). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo et al. 7237 (GUAY). M: Neill et al. 11408 (NY, QCNE). Distribution: Belize and Costa Rica to northwestern Ecuador. Ecology: Pterocarpus officinalis grows along stream margins down to swamps and mangroves. It is often a dominant species in swamps. Phenology: This species has been collected in flower from February to April, and in October and November; and in fruit from December to May, August, and September.

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Biological interactions: The petals shed on the water are eaten by Sabaleta fish. Dispersal: The seeds are water dispersed. Uses: The wood is soft and weak (Little, 1969) and is used as firewood. Many years ago the pieces of buttresses were used as pans to wash gold in rivers. The bark when wounded yield drops of a red resin, which in contact with the air soon solidifies into crimson tears; by its color it was known as “sangre de dragón” (Spanish, meaning dragon’s blood). This was extracted and exported from Colombia to Spain because of its medicinal uses as a hemostatic and an astringent. The trees have been planted as ornamentals in southern Florida and Cuba (Little, 1969). IUCN Red list category: Due to the widespread range of distribution and the common presence of healthy populations, it is herein categorized as of Least Concern (LC). Etymology: The epithet is a Latin word referring to the fact that the resin was traditionally sold for its medicinal properties. GESNERIACEAE (Gesneriad Family). Epiphytic or terrestrial herbs or shrubs. Leaves opposite, rarely whorled or alternate; blades equal to strongly unequal, simple, entire to toothed; petiolate. Inflorescences axillary or terminal. Flowers usually zygomorphic. Calyx of 4–5 lobes. Corolla variously colored, with a tube of 5 connate petals, often oblique in calyx. Stamens 2–4; staminodes 1–3, usually didynamous. Ovary superior to inferior; style usually elongating as stamens; stigma usually 2-lobed. Fruit a berry, or a dry or fleshy capsule, 2- or 4-valvate; seeds numerous.

A family best represented in the terra firme wet forests and rainforests. Key to the species

1. Leaf blades thick, fleshy, the margins enire; fruit a juicy berry Codonanthe crassifolia 1. Leaf blades thin, coriaceous, the margins dentate; fruit a dehiscent capsule Drymonia serrulata

117. Codonanthe crassifolia (Focke) C. V. Morton, Publ. Field Mus. Nat. Hist., Bot. Ser 18(4): 1159 (1938). Plate 26: A. Bas.: Hypocyrta crassifolia Focke Syn.: Episcia hookerii Hanst. Codonanthe confusa Sandwith Holoepiphytic climber herbs. Stem loose to pendulous, to 50 cm long. Leaves opposite, equal in a pair; blades thick and fleshy, elliptic, to ca. 3 x 2 cm, entire, glabrous; petioles to 1 cm long. Inflorescences axillary. Calyx greenish, the lobes triangular, to 6 mm long. Corolla funnelform, broader toward mouth, to 4 x 1 cm, white, sometimes flushed with pink on outer

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surface of lobes. Stamens 4, included. Ovary superior. Fruit a juicy berry, widely ovoid, red at maturity, with persistent calyx; seeds numerous. COLOMBIA. CH: (E). V: Killip 5221 (NY). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6946 (GUAY, US). Distribution: Mexico to Bolivia. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The funnelform corollas suggest pollination by bees. The fleshy fruit is most likely eaten by birds and bats, while the minute seeds, embedded in a sticky and juicy mesocarp, are endozoochorously dispersed. Use: This species may be planted as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the thick leaf blades. 118. Drymonia serrulata (Jacq.) Mart., Nov. Gen. Sp. Pl. 3: 59 (1929). Bas.: Besleria serrulata Jacq. Syn.: Besleria spectabilis Kunth Drymonia spectabilis (Kunth) Mart. ex G. Don Hemiepiphytic climber herbs. Stem appressed, to 3 m long. Leaves opposite, equal in a pair; blades coriaceous, pale-green or sometimes suffused with dull lavender or maroon, dentate at margins. Inflorescences axillary; bracts green. Calyx green. Corolla funnelform, broader toward mouth, 3â&#x20AC;&#x201C;5 cm long, creamy-yellow or greenish-white. Stamens 4, included. Ovary superior. Fruit a fleshy capsule, globose, dehiscing by 2 valves that recoil to reveal dark winered interior. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo 2669 (GUAY, US). Distribution: Mexico to Bolivia. Ecology: In wet forests and rainforests. Biological interactions: The flowers have been reported to be pollinated by diurnal bees of the genus Epicharis (Steiner, 1985). IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the serrulate margin of leaf blades. HIPPOCRATEACEAE (Hippocratea Family) Trees, shrubs, scandent shrubs, or lianas. Branches mostly opposite. Leaves opposite, simple, petiolate; blades with margins entire, undulate or crenate-serrate. Inflorescences axillary. Flowers actinomorphic,

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usually very small, bracteolate, pedicellate. Sepals 5, imbricate, basally united, often persisting. Petals 5, free, inserted under disk, imbricate. Disk always present, variable. Stamens 3(–5), inserted within disk. Ovary superior, 3-locular. Fruits capsules or drupes, dehiscent or indehiscent; capsular fruits with 3 mericarps, free or laterally ± united. Fruits drupaceous with an indehiscent pericarp, leathery, fleshy or woody.

A family represented by one species of lianas in mangroves.

119. Hippocratea volubilis L., Sp. Pl. 2, 1191 (1753). Syn.: Hyppocratea ovata Lam. Woody lianas. Stem richly lenticellate, glabrous. Branches opposite, (sub)horizontal. Leaves simple, opposite; blades chartaceous to coriaceous, elliptic to obovate or lanceolate, 5–15 x 2–6 cm, cuneate at base, acuminate at apex, inconspicuous to softly crenate, glabrous, the lateral veins 6–9 per side, the tertiaries reticulate, prominent on both sides; petioles 2–10 mm long. Flowers 4–6 mm diam. Infructescences 3-capsular, dehiscent, divergent, narrowlyobovate or elliptic, 5–6 x 2.5–3.5 cm, subwoody, strongly complanate, with prominent longitudinal veins, brown at maturity, glabrous; seeds winged, oblong, lateraly curved, ca. 4.5 x 0.8 cm, complanate, yellow. COLOMBIA. CH: (E). V: (E). C: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 7253 (GUAY, MO). Distribution: Mexico to Argentina. Ecology: Hippocratea volubilis is a fringe species of the hyposaline mangroves and terra firme forests inland, inhabiting moist, wet forest, and rainforest life zones. IUCN Red list category: Least Concern (LC). Etymology: It is most likely that the epithet refers to the vegetative variability. LORANTHACEAE (Showy Mistletoe Family) Parasitic shrubs or subshrubs, erect or scrambling. Leaves alternate, decussate or whorled, entire; with or without epicortical roots from base, sometimes also from stem. Inflorescences terminal and axillary, the flowers mostly in monads, dyads, or triads with fused bracts and/or bracteoles. Flowers basically dichlamydeous but calyx reduced to a very inconspicuous calyculus, usually perfect, some species dioecious. Petals 4–7, often brightly coloured when flowers perfect. Stamens epipetalous. Style in many genera slender and often long. Fruit drupaceous.

The members of this family usually are infrequent elements in the mangrove interior. All cited

species are parasites or hemiparasites of trees.

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Key to the species 1. Inflorescence a spike or raceme of single lateral flowers throughout 2. Stems terete; inflorescences axillary or in terminal, compound arrangements, or both Oryctanthus occidentalis 2. Stems angled at least when young; inflorescences in axils of foliar leaves only Oryctanthus florulentus 1. At least the lower part of the inflorescence constructed of triads or dyads, these arranged in spicate, racemose or umbellate fashion 3. Flowers less than 1 cm long, not brighly coloured, in short-pedunculate or sessile triads; epicortical roots from base of plant and often from stem 4. Anthers dorsifixed, the flowers pale greenish yellow Struthanthus orbicularis 4. Anthers basifixed, the flowers reddish or yellowish 5. Flowers perfect; epicortical roots only at base of plants, not from stems; inflorescence a simple, axillary raceme of triads only Phthirusa pyrifolia 5. Flowers unisexual, plants dioecious; epicortical roots both from base and from stems; larger inflorescences branched, either axillary or terminal Phthirusa stelis 3. Flowers more than 3 cm long, brighly colored, in long-pedunculate triads or diads; epicortical roots lacking Psittacanthus divaricatus 120. Oryctanthus florulentus (Rich.) Tiegh., Bull. Mus. Hist. Nat. Paris 2: 339 (1896). Bas.: Loranthus florulentus Rich. Syn.: Loranthus ruficaulis Poepp. & Endl. Oryctanthus ruficaulis (Poepp. & Endl.) Eichler in Mart. Parasitic shrubs, erect. Stems sharply angular until older, when becoming ± terete. Internodes usually 3 cm long or less, youngest twigs with very conspicuous furfuraceous lines continuing along lower midrib of each leaf blade into apex or nearly so. Leaf blades rather thin, broadlylanceolate, 4–7.5 x 2–3.5 cm, acute at base, rounded at apex, venation pinnate; petioles ca. 3 mm long. Spikes axillary only, 1–3 per axil, 2–4 cm long; peduncle less than 2 mm long, the floriferous portion rather slender. Flowers somewhat angular, ca. 2 mm long, greenish, orange or light-red. Anthers very dimorphic. Stigma reaching above upper anthers. Fruit rather bluntly ellipsoid, nearly perpendicular to axis, orange-red. COLOMBIA. CH: (E). V: (E). C: (E). N: (E). ECUADOR. E: Játiva & Epling 2226 (MO, NY, S, US). Distribution: Colombia and Venezuela to Bolivia. Ecology: In wet forests and rainforests. Biological interactions: The fruit is most likely eaten by birds, while the small seeds are endozoochorously dispersed. IUCN Red list category: Least Concern (LC).

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121. Oryctanthus occidentalis (L.) Eichler in Mart., Fl. Bras. 5(2): 89 (1868). Bas.: Loranthus occidentalis L. Parasitic shrubs, erect. Young stem and peduncles light-cinnamon, furfuraceous, later becoming grayish, always terete. Internodes 2–5(–11) cm long. Leaf blades coriaceous, elliptic or orbicular to broadly-lanceolate, obtuse to acute at base and apex, the midrib usually not reaching the apex; petioles to 1 cm long or nearly absent. Spikes axillary, solitary or in groups, 2–4(–12) cm long. Flowers ca. 2 mm long, about 1/3 of which is ovary, with smooth to minutely denticulate calyculus. Petals and anthers dimorphic. Fruit obovoid or ellipsoid to nearly spherical, (2–)3 x 2 mm, blunt and slightly tuberculate at apex, perpendicular, greenish or green and yellow, or reddish. COLOMBIA. CH: (E). V: (E). C: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 7254 (GUAY). Distribution: Costa Rica to Brazil. Ecology: In moist forests, wet forests, and rainforests. Biological interactions: The fruit is most likely eaten by birds, while the small seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet refers to the species distribution, in the Western hemisphere. 122. Phthirusa pyrifolia (Kunth) Eichler in Mart., Fl. Bras. 5(2): 63 (1868). Bas.: Loranthus pyrifolius Kunth Syn.: Passovia pyrifolia (Kunth) Tiegh Vernacular name: matapalo. Parasitic shrubs, erect. Stem much compressed and keeled when young. Epicortical roots from base of plants only. Leaf blades dark-green, shiny, rather thin, broadly-lanceolate, ca. 10 x 4–5 cm, obtuse to truncate at base, somewhat attenuate or even mucronate at apex, venation pinnate; petioles distinct, ca. 1 cm long, somewhat compressed or keeled. Inflorescence mostly one per foliar axil, sometimes clustered but not in terminal groups, 6–8 cm long. Flowers sessile, the lateral with 2 prophylls each, forming a small, dentate floral cup. Petals ca. 1.5 mm long, dark wine-red to pale red. Stamens sessile. Fruit a blunt, ellipsoid berry, ca. 8 x 5 mm, bright orange-red at maturity with dark purple-green base and yellowish apex. COLOMBIA. V: (E). N: (E). ECUADOR. G: Harling & Andersson 19345 (GB, LEA). Distribution: Southern Mexico to Bolivia. Ecology: In moist forests, wet forests, and rainforests.

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Biological interactions: The fruit is most likely eaten by birds, while the small seeds are endozoochorously dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the pear-like shape of the leaf blades, a variable feature. 123. Phthirusa stelis (L.) Kuijt, Taxon 43: 193 (1994). Bas.: Loranthus stelis L. Syn.: Loranthus retroflexus Ruiz & Pav. Phthirusa retroflexa (Ruiz & Pav.) Kuijt Loranthus aduncum G. Mey Phthirusa adunca (G. Mey.) Maguire Loranthus conduplicatus Kunth Loranthus paniculatus Kunth Phthirusa paniculata (Kunth) J. F. Macbr. Loranthus orinocensis Spreng. in Roem. & Schult. Loranthus magdalenae Cham. & Schltdl. Phthirusa maritima Rizzini Phthirusa anastyla Rizzini in Luces & Steyerm. Phthirusa pyramidalis Rizzini in Luces & Steyerm. Phthirusa cothurnata Rizzini Phthirusa perdivergens Rizzini ? Phthirusa rubromicans Rizzini Vernacular name: matapalo. Parasitic climbers. Stem terete or when young slightly compressed, with occasional roots from internodes. Leaf blades broadly ovate, to 14 x 7.5 cm, usually attenuate at apex; petioles ca. 1 cm long. Dioecius. Inflorescences both terminal and axillary, in the former case always and in the latter case usually compound, consisting of a terminal raceme plus 2â&#x20AC;&#x201C;6 lower, lateral racemes, each with about 12 pairs of evenly spaced triads; compound inflorescences to 15 cm long. Triad peduncles to 2 mm long. Buds to 6 mm long. Petals whitish-yellow to light-red. Anther connective forming a small, terminal horn. Fruit ellipsoid, to 10 x 5 mm, brick-red. COLOMBIA. V: (E). N: (E). ECUADOR. G: Harling & Andersson 19338 (GB, LEA). O: Asplund 15874 (S). Distribution: Costa Rica to Bolivia. Ecology: In moist forests, wet forests, and rainforests. Biological interactions: The fruit is most likely eaten by birds, while the small seeds are endozoochorously dispersed.

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Use: The plant is applied as a cataplasm to remove bruising inflammation. IUCN Red list category: Least Concern (LC). Note: The taxonomy and nomenclature of Phthirusa stelis are extremely complex (Kuijt and Kellogg, 1996; Kuijt, 2001). This species has been treated under the name of P. retroflexa (Kuijt, 1986). 124. Psittacanthus divaricatus (Kunth) G. Don, Gen. Hist. 3: 417 (1834). Plate 26: B. Bas.: Loranthus divaricatus Kunth Syn.: Psittacanthus chanduyensis Eichler Psittacanthus ovbovatus (Benth.) Eichler Psittacanthus tumbecensis (Killip) J. F. Macbr.

Psittacanthus weberbaueri Pacz.

Vernacular names: chichihua, hierba de pajarito, liga, suelda con suelda. Parasitic shrubs, suberect. Stem terete, oldest stem with small, brown, dot-like lenticells. Leaf blades sessile to clasping, somewhat leathery, elliptical or ovate-cordate to nearly orbicular, 1.5–7.5 x 1–5 cm, rounded at apex. Inflorescence terminal, sometimes accompanied by two smaller ones in axils of subjacent leaves, each with 4–6 triads; triad peduncle to 10 mm long, the pedicels to 8 mm long. Buds slightly curved, 2–3 mm thick, blunt-tipped where slightly plicate. Petals 30–40 mm long, bright-scarlet with some yellow at apex; petals and stamens dimorphic, the filaments attached to petals ca. 2 cm long. Fruit ovoid, ca. 7 x 5 mm, blunt, blackish, the cupule not accrescent. ECUADOR. G: Hitchcock 20004 (US). O: Asplund 15859 (S). PERU. T: Smith 10104 (MO). Distribution: Ecuador and Peru. Ecology: In thickets, dry thorn scrub, and xerophytic forests. Biological interactions: Based on a report of a similar species in the same genus (Fleming et al., 2009), it is most likely that bats pollinate the flowers of this species. The fruit is eaten by birds, and the small seeds are endozoochorously dispersed. Use: It is said that the species is used as medicine to heal broken bones, hence “suelda con suelda”, one of the vernacular names. That vernacular name is also applied to Catasetum macroglossum, an orchid from western Ecuador (and northwestern Peru?) also occurring in mangroves that is used for the same medicinal purposes as P. divaricatus. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the widely divergent arrangement of branches.

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125. Struthanthus orbicularis (Kunth) Blume in Schult. f., Syst. Veg. 7: 1731 (1830). Plate 26: C. Bas.: Syn.:

Loranthus orbicularis Kunth Notanthera orbicularis (Kunth) G. Don Phthirusa orbicularis (Kunth) Rusby ex De Wild. Loranthus lucarquensis Kunth Strutanthus chordocladus (Oliver) Eichler Spirostylis chordocladus (Oliver) Tiegh

Vernacular name: matapalillo. Parasitic climbers. Roots formed from the young stems specially where touching other stems, frequently establishing new haustorial contacts. Young branches quadrangular, the terminal branchlets pendulous. Leaves first lanceolate and usually recurved, the petioles then prehensile and capturing other branches, forming a tangled network. When leaves are not successful as prehensile organs, then developing a petiole at least 1 cm long; blades lanceolate to obovate or orbicular, to 7 x 6 cm, variable at base and apex. Staminate spike up to 13 cm long; pistillate spike somewhat smaller than male, both bearing sessile triads. Flowers sessile, ca. 5 mm long, greenish. Fruit ellipsoid, 13 x 8 mm, purplish-red, becoming dark-blue. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 7255 (GUAY, QCNE). Distribution: Southern Mexico to Brazil. Ecology: In dry, moist, and wet forests. Biological interactions: The fruit is most likely eaten by birds, while the small seeds are endozoochorously dispersed. Use: The plant is applied as a cataplasm to remove bruising inflammation. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the orbicular shape of leaves, a variable feature. LYTHRACEAE (Loosestrife Family) Herbs, shrubs or trees. Leaves simple, entire, decussate or rarely partly alternate, or verticillate. Flowers solitary or in racemes, rarely in panicles. Bracts often caducous. Flowers hermaphrodite, actinomorphic or zygomorphic. Calyx persistent, the lobes acute, intersepalic appendages generally present. Petals free, sometimes reduced or absent, inserted on rim of calyx tube, mostly caducous. Stamens 11 to numerous, the filaments of unequal length. Disk glandular. Ovary superior, 2-carpellate, usually 1â&#x20AC;&#x201C;2 locular; style filiform; stigma 2-lobed, capitate, discoid or inconspicuous, papillose. Fruit a capsule, sometimes coriaceous, more or less included in calyx, dehiscent or indehiscent. This family is represented by Crenea in mangroves, a ditypic genus known from the Caribbean and the Pacific coast of northern South America (Lourteig, 1989).

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126. Crenea patentinervis (Koehne) Standl., Field Mus., Bot. Ser. 23: 218 (1947). Plate 26: D. Bas.: Dodecas surinamensis var. patentinervis Koehne Syn.: Crenea surinamensis subsp. patentinervis (Koehne) Koehne Vernacular names: achotillo, matapalillo de barrial. Subshrubs or herbs, 0.5–4 m tall. Stem reddish, tetragonous, 4-winged, with smooth or crenulate wings. Leaves decussate; blades lanceolate-elliptic to elliptic-oblong, 3–11 x 1.5– 2.5 cm, cuneate at base, acute or subacute at apex, with narrow scarious margin, the midrib reddish, the secondary veins 10–20 per side, broquidrodomus; petioles 2–5 mm long. Flowers solitary or in 2–5-flowered cymes. Peduncle 5–15 mm long, the pedicels 2.5–3.5 mm long. Calyx ca. 6 mm long, green with reddish edges, the lobes triangular to oblong, acute to shortly acuminate at apex. Petals 4, shortly-clawed, suborbicular to somewhat obovate, ca. 8 mm long, white. Stamens 8–10, the anthers yellow. Fruit capsule, globose, 5–6 mm diam., indehiscent, reddish to red, more or less enclosed in calyx. Seeds very narrow, ca. 2 mm long. COLOMBIA. CH: Gentry & Juncosa 40939 (COL, MO). V: Juncosa 2094 (NY). C: (E). N: Toro 86 (NY). ECUADOR. E: Cornejo & Bonifaz 2716 (GUAY, QCNE). Distribution: Endemic to the mangroves and adjacent riverbanks of the Pacific coast in Colombia and northwestern Ecuador. Ecology: An often frequent species in the mangroves located in wet forest and rainforest life zones. It sometimes forms understory stands, mainly at fringes in secondary mangrove forests. It also occurs in the intertidal zone at adjacent riverbanks. Phenology: This species has been collected in flower and in fruit from February to September. IUCN Red list category: Although the species is often locally common, due to factors such as a narrow range of distribution of less than 2,000 km2 and a loss of habitat, Crenea patentinervis is herein categorized as Vulnerable (VU B2abiii). Etymology: The epithet refers to the widely divergent to patent arrangement of the secondary veins in leaf blades. Note: This is a true mangrove species. As previously noted, “it could well be described as a mangrove, but is exclusively herbaceous and has no morphological adaptations to mangrove ecosystems” (Tomlinson, 1986). MALPIGHIACEAE (Barbados Cherry Family) Trees, shrubs and vines; hairs unicellular, usually medifixed or submedifixed (“T” shape). Stipules usually present. Leaves usually opposite, often bearing glands on petioles and/or blades; blades simple, entire. Flowers usually bisexual, bilaterally symmetrical. Sepals 5, bearing 1(–2) large abaxial glands; petals 5, distinct, clawed, alternating with sepals, imbricate. Stamens mostly 10. Gynoecium superior, comprising (2–)3 carpels; styles mostly 1 per carpel and distinct. Fruits dry or fleshy, dehiscent or indehiscent, samaroid, nut-like or drupaceous.

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Key to the species 1. Lianas, sometimes shrubs; fruits dry, samaroid 2. Petioles with a distal pair of glands 3. Inflorescences with peduncles less than 10 cm long 4. Fruits with triangular-ovate or asymmetric-hemiorbicular wings 5. Samara wing thick Hiraea brachyptera 5. Samara wing membranose 4. Fruits with asymmetric-oblongoid wings 3. Inflorescences with peduncles 10–30 cm long

Hiraea fagifolia Stigmaphyllon ellipticum

Banisteriopsis martiniana subsp. subenervia

2. Petioles eglandular 6. Fruits 2–6 cm long, the wings developed 7. Inflorescences cymose; fruits 2–3.3 x 0.7–1.3 cm 7. Inflorescences panicle of pseudoracemes; fruits 3–6 x 2–4 cm

Banisteriopsis elegans Heteropterys leona

6. Fruits 0.6–1.1 cm long, the wings vestigial

Tetrapterys subaptera

1. Trees; fruits fleshy, drupaceous

Bunchosia plowmanii

127. Banisteriopsis elegans (Triana & Planch.) Sandwith, J. Arnold Arbor. 24: 222 (1943). Bas.: Banisteria elegans Triana & Planch. Syn. Banisteria elegans subsp. cordata Nied.

Banisteria elegans subsp. cordata var. ciliata Nied.

Banisteriopsis scalariformis Schery Banisteriopsis martiniana var. elegans (Triana & Planch.) Cuatrec. Lianas. Branches appressed-sericeous, glabrescent. Leaves opposite; blades ovate to broadlyelliptic, 7–23 x (3–)5–13 cm, obtuse to cordate at base, acute to acuminate at apex, the lateral veins 5–8 per side, beneath sparsely appressed-sericeous or glabrous, bearing a pair of glands near base on each side of midrib, the glands more numerous towards apex; petioles 0.4–1.4 cm long, eglandular. Inflorescences axillary, cymose, the peduncle 10–35 cm long; flowers 4(–6) per umbel. Sepals 2–3 x 1.5–1.8 mm. Glands 1–2 x 0.6–1 mm. Petals clawed, yellow, lacerate to fimbriate at margins. Stamens unequal. Samara wing foliaceous, asymmetricoblanceolate, 2–3.3 x 0.7–1.3 cm, sparsely sericeous to glabrate.

COLOMBIA. CH: (E). V: Killip & Cuatrecasas 38987 (NY). Distribution: Guatemala to Peru.

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Ecology: In wet forests and rainforests. Biological interactions: Based on a pollination report in some species of this genus (Sigrist and Sazima, 2004), it is most likely that oil-collecting bees are the pollinators. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the elegant appearance of this species. 128. Banisteriopsis martiniana subsp. subenervia Cuatrec., Webbia 13: 501 (1958). Plate 27: A. Syn.: Banisteriopsis martiana var. laevis Cuatrec. Lianas. Branches glabrescent. Leaves opposite; blades elliptic, 9–17 x 5–8 cm, subcordate at base, shortly acuminate or mucronate at apex, the lateral veins 5–8 per side, beneath very sparsely pilose and bearing a pair of glands near base on each side of midrib, and distantly with several inconspicuous scattered glands in an inframarginal row; petioles 0.3–1 cm long, with a distal pair of prominent sessile glands beneath. Inflorescences axillary, dichasia or thyrses, the peduncle 10–30 cm long; flowers 2–5 per umbel. Sepals ca. 2 x 1.5 mm. Glands 1–1.5 x 1–1.3 mm. Petals clawed, yellow, erose or denticulate at margins. Stamens unequal. Samara not seen. ECUADOR. E: Cornejo & Cangá 8135 (GUAY, NY, MICH, QCNE). Distribution: Colombia and Venezuela to Brazil. Ecology: In wet forests. Biological interactions: Based on a pollination report in some species of this genus (Sigrist and Sazima, 2004), it is most likely that oil-collecting bees are the pollinators. IUCN Red list category: Least Concern (LC). 129. Bunchosia plowmanii W. R. Anderson, Contr. Univ. Mich. Herb. 16: 66, fig. 5 (1987). Plate 27: B. Small trees or shrubs 1.5–4 m tall. Branches sericeous or subvelutinous to soon glabrate. Leaves opposite; blades elliptical or slightly obovate, 7–14 x 3.5–6.5 cm, cuneate at base, mostly abruptly shortly-acuminate and with a small gland at apex, beneath persistently sericeous or subvelutinous and bearing a pair of glands near base on each side of midrib, and distantly 1–10 or more glands in an inframarginal row. Inflorescences axillary and rarely terminal pseudoracemes, 3–10 cm long, bearing (4–)8-26 flowers; pedicels 4–8 mm long, usually muricate. Sepals extending 1–2 mm beyond glands. Glands 8, 1.7–2.7 mm long. Petals light-yellow, clawed, with margin erose-denticulate. Stamens 2.5–3.3 mm long. Fruit drupaceous, ovoid to globose, 5–8 x 4.5–8 mm (dried), orange to red at maturity, glabrous or glabrate, granulate.

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ECUADOR. G: Asplund 15305 (S). Distribution: Southwestern Ecuador to northwestern Peru. Ecology: This species occurs in the dry forest life zone, from sea level to 1200 m (Anderson, 1987). Biological interactions and dispersal: There is no information recorded, but it is most likely that oilcollecting bees pollinate this species, and that birds and bats eat the fleshy and juicy mesocarp of the drupaceous fruits, then exozoochorously disperse the seeds. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat, this species is herein categorized as Near Threatened (NT). Etymology: The epithet honors Timothy C. Plowman (1944–1989), a North American botanist and specialist in Erythroxylaceae (coca family), who collected the type in the Department of Tumbes, Peru (Anderson, 1987). 130. Heteropterys leona (Cav.) Exell, Cat. S. Tomé 123 (1944). Bas.: Banisteria leona Cav. Syn.: Malpighia reticulata Poir. Banisteria multiflora DC. Banisteria magnoliifolia Desv. ex Hamilton Lianas or climbing shrubs. Terminal branches appressed-tomentose, soon glabrescent. Leaves opposite; blades oblong, elliptical, lanceolate, ovate, or oblanceolate, 10–37.5 x 3.5– 19.7 cm, rounded to subcordate at base, acuminate at apex, the lateral veins 7–10 per side, beneath sparsely sericeous or sometimes glabrate or rarely glabrous; petioles 0.5–1.8 cm long, eglandular. Inflorescences axillary, a panicle of pseudoracemes, the peduncle 6–18(– 30) cm long. Sepals 2.5–4 mm long, the lateral ones exceeding the glands by ca. 1 mm. Petals clawed, yellow or pale yellow, erose or erose-fimbriate at margins. Stamens unequal. Samara wing asymmetric-oblanceolate, 3–6 x 2–4 cm. COLOMBIA. CH: (S). V: (E). Distribution: Mostly inhabiting the Atlantic coastal regions from Belize to northernmost Brazil, as well as on the Pacific side of Colombia (Choco, Valle); rare in Jamaica; present in the Old World, in coastal West Africa, from Senegal to Angola (Anderson, 2001). Ecology: Present in riverine ecosystems, swamps, flooded forests, beaches, mangroves, and lagoon edges, all in wet forest and rainforest life zones from sea level to 200(–450) m (Anderson, 2001). Biological interactions: Based on a pollination report of some species of this genus (Sigrist and Sazima, 2004), it is most likely the flowers are pollinated by oil-collecting bees. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to Sierra Leona, Africa, from where the type collection was gathered.

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131. Hiraea brachyptera Triana & Planch., Ann. Sci. Nat., Bot., sér. 4, 18: 332 (1862). Lianas or shrubs. Leaves opposite; blades coriaceous, lanceolate-oblong, 8–17 x 4–7 cm, cuneate to obtuse at base, acuminate to cuspidate at apex, drying brown, the lateral veins ca. 7 per side; petioles ca. 1 cm long, with a distal pair of prominent sessile glands. Inflorescences apical on short lateral branches; flowers often 3-umbellate, these borne in cymes. Sepals ca. 3 mm long. Glands 1–2 mm long. Petals clawed, yellow, erose-dentate at margins. Stamens unequal. Samara wing triangular-ovate, thick, sericeous. COLOMBIA. CH: Fuchs et al. 22153 (COL). V: Killip & Cuatrecasas 38626 (COL, MO). C: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 7251 (AAU, GUAY, MICH). Distribution: On the Pacific side of Colombia to northwestern Ecuador. Ecology: In wet forests and rainforests, often collected in mangroves. IUCN Red list category: Due to factors such as a distribution range of less than 2,000 km2 and a loss of habitat, this undercollected woody species is herein categorized as Vulnerable (VU B2abiii). Etymology: The epithet refers to the short wings of the fruits. Note: Hiraea brachyptera was previously known as endemic to the Pacific coast of Colombia. The cited collection is the first record for Ecuador (W. R. Anderson pers. comm.). 132. Hiraea fagifolia (DC.) A. Juss, Ann. Sci. Nat., Bot., sér. 2, 13: 258 (1840). Bas.: Syn.:

Banisteria fagifolia DC. Hiraea riedleyana A. Juss. Hiraea moricandiana A. Juss. Hiraea blanchetiana Moric. Hiraea bahiensis Moric. Lianas or shrubs. Leaves opposite; blades chartaceous, elliptic, ovate or obovate, 8–18 x 3–7 cm, obtuse to subcordate at base, acuminate at apex, the blade initially sericeous, mostly glabrescent on both sides, except along midvein, usually with several submarginal small foliar glands distally arranged, and 2 slightly larger glands near or on the midvein beneath, the lateral veins 9–12 per side; petioles 0.7–1.2 cm long, with a distal pair of shortly linear glands. Inflorescences axillary, ca. 1.5–4 cm long; flowers often 3-umbellate, these borne in cymes. Sepals all eglandular, otherwise the 4 lateral 2-glandular, orange. Petals clawed, the 4 lateral dentate or laciniate and eglandular, the posterior glandular-fimbriate at margins, yellow. Samara wing membranose, asymmetric-hemiorbicular, 1.3–2.5 x 2–3.8 cm, sericeous, somewhat glabrescent.

COLOMBIA. CH: Acevedo 6863 (MO). Distribution: Southern Mexico to northern Argentina.

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Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to a slight resemblance of the leaf blades to those of the genus Fagus (Fagaceae; W. R. Anderson, pers. comm.). 133. Stigmaphyllon ellipticum (Kunth) A. Juss., Ann. Sci. Nat., Bot., sér. 2, 13: 290 (1840). Plate 27: C. Bas.: Banisteria elliptica Kunth Syn.: Banisteria mucronata DC. Stigmaphyllon mucronatum (DC.) A. Juss. Stigmaphyllon mucronatum var. nicaraguense Nied. Stigmaphyllon ellipticum var. nicaraguense (Nied.) Nied. Vines. Branches glabrous. Leaves opposite; blades ovate to elliptic, 4–10 x 3–6 cm, truncate to cordate at base, acute to mucronate at apex, the lateral veins 7–9 per side, glabrous; petioles 0.6–2 cm long, with a distal pair of prominent sessile glands. Inflorescences apical on short lateral branches, solitary or borne in dichasia or small thyrses, the peduncles 2.5–3.4 cm long; flowers 2–8 per umbel. Sepals 2–3.5 x 1.7–2 mm. Glands 1.1–2.5 x 0.8–1.5 mm. Petals clawed, yellow, lacerate or fimbriate at margins. Stamens unequal. Samara wing coriaceous, asymmetric-oblanceolate, to 2.5 x 0.8 cm, reddish at maturity, mainly at outer margin and apex. COLOMBIA. CH: Forero & Gentry 796 (MO, NY). V: Gentry & Juncosa 40716 (COL, MO). C: (E). N: (E). ECUADOR. E: Bonifaz & Cornejo 3997 (AAU, GUAY, MICH). Distribution: Mexico to Brazil, and Peru. Ecology: Inhabiting ecosystems from the terra firme forests to the fresh water swamps, as well as mangroves and sandy beaches, in dry, moist, and wet forests. Biological interactions: Based on a pollination report of some species of this genus (Sigrist and Sazima, 2004), it is most likely that the flowers are pollinated by oil-collecting bees. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet refers to the elliptical leaf blades, a variable feature. 134. Tetrapterys subaptera Cuatrec., Webbia 13(2): 423–425, fig. 7 (1958). Plate 27: D. Woody lianas or scandent shrubs. Branches appressed-ferrugineus, glabrescent. Leaves opposite; blades chartaceous, lanceolate to ovate-lanceolate, 5–15 x 2–6 cm, obtuse at base, attenuate at apex, with 2–4 glands, glabrous above, inconspicuous sparsely appressed beneath, the lateral veins 6–9 per side; petioles 0.5–1 cm long, eglandular. Inflorescences terminal and axillary at distal portion of branches, cymose-paniculate; flowers often 4-umbellate. Sepals 2–2.5 mm long. Glands 8, ca. 3 mm long. Petals clawed, yellow, denticulate at margins. Stamens unequal. Fruit reduced samara, ovate-acuminate, 0.6–1.1 cm long, the wings vestigial, the lateral wings 2–8 mm long.

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COLOMBIA. CH: (E). V: Killip 5219 (NY). C: (E). N: (S). ECUADOR. E: Bonifaz & Cornejo 4002 (AAU, GUAY, MICH). Distribution: Southeastern Panama to northwestern Ecuador. Ecology: It seems to be restricted to Pacific coast mangroves, in wet forests and rainforests. Biological interactions: Based on a pollination report of some species of this genus (Sigrist and Sazima, 2004), it is most likely that the flowers are pollinated by oil-collecting bees. Dispersal: The fruits bearing vestigial wings are presumably water-dispersed rather than wind-dispersed, which is the usual dispersal method for the genus (Cuatrecasas and Croat, 1980). IUCN Red list category: Due to factors such as a distribution of less than 2,000 km2 and a loss of habitat, this species is herein categorized as Vulnerable (VU B2abiii). Etymology: The epithet refers to the characteristic vestigial wings. MALVACEAE (Mallow Family) Herbs, shrubs or trees, often stellate-pubescent. Leaves alternate, stipulate. Flowers solitary or fasciculate in axils of leaves or grouped in racemes or panicles. Calyx gamosepalous, truncate, 5-toothed, or 5-parted. Petals 5, distinct, usually clawed, adnate to staminal column at base. Androecium monadelphous, the anthers reniform, numerous. Gynoecium superior, 3–40-carpelled; styles 1–40; stigmas truncate, capitate or decurrent. Fruit schizocarpic or capsular, sometimes a berry; mericarp sometimes divided internally by a tongue-like intrusion; seeds reniform or turbinate, rarely arillate.

A family represented in mangroves by Talipariti and Pavonia. Thespesia populnea is a species

recently introduced to Ecuador and a new record for the country. It is occasionally cultivated as an ornamental in General Villamil (Playas, Prov. Guayas), and infrequently is observed on sandy beaches (in Prov. Manabi), presumably escaping cultivation. It still has not been recorded in mangrove areas, but as it has been reported as an associated species (Tomlinson, 1986), it is expected to eventually occur there. Key to the species 1. Woody shrubs or trees; leaf blades deeply cordate, 7–15 cm wide; petals 4–6 cm long Talipariti tiliaceum var. pernambucense 1. Subshrubs or weak shrubs; leaf blades lanceolate to lanceolate-oblong, oblanceolate or narrowlyspathulate 0.5–5 cm wide; petals 0.8–2 cm long 2. Leaf blades lanceolate to lanceolate-oblong, 7–14 cm long; inflorescence an elongated terminal raceme; fruits without spines

Pavonia rhizophorae

2. Leaf blades oblanceolate or narrowly-spathulate, 2–4 cm long; flowers solitary, axillary; fruits 2-spined

Pavonia kearneyi

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

135. Pavonia kearneyi D. M. Bates, Kew Bull. 20: 149 (1966). Subshrubs 0.5 m tall. Stems minutely and sparsely lepidote to glabrate. Stipules vestigial. Leaf blades oblanceolate or narrowly-spathulate, 2–4 x 0.5–1.2 cm, attenuate-cuneate at base, obtuse at apex, entire to ± repand at margins; petioles to 2.5 mm long (the leaves subsessile), sparsely lepidote to glabrate. Flowers solitary, axillary; involucellar bracts 5–6, linear-lanceolate, 1–5 mm long; pedicels 4–8 mm long (to 12 mm in fruit), recurved. Calyx 5–6 mm long, green, the lobes lanceolate, lepidote or glabrate. Petals 8–11 mm long, white to reddish-white. Staminal column included. Fruits ca. 7 mm diam. (excluding spines); mericarps 2-spined, the spines 16–19 mm long. COLOMBIA. CH: Gentry 53854 (CUVC, MO, NY). V: Juncosa 2091 (MO, NY). Distribution: Western Colombia. Ecology: Often growing partially submersed in the intertidal zones at rivermouths and in mangroves, in the rainforest life zone. It is sympatric to Pavonia rhizophorae. IUCN Red list category: Due to factors such as a range of distribution of less than 2,000 km2 and a loss of habitat, Pavonia kearneyi is herein categorized as Vulnerable (VU B2abiii). Etymology: The epithet honors Thomas H. Kearney (1874–1956), a North American botanist and taxonomist of Malvaceae. 136. Pavonia rhizophorae Killip ex Kearney, Leafl. W. Bot. 7: 118 (1954). Plate 28: A. Weak shrubs or subshrubs 1.5–3 m tall. Stems stellate-pubescent. Stipules inconspicuous, deciduous. Leaf blades lanceolate-oblong, 7–14 x 3–5 cm, truncate and palmately 3-nerved at base, acuminate at apex, remotely denticulate to subentire at margins; petioles 1–4 cm long, stellate-pubescent. Inflorescence an elongated terminal raceme; involucellar bracts 6–8, linear-lanceolate, 6–10 mm long; pedicels 0.5–1.5 cm long. Calyx 10–13 mm long, green, the lobes ovate-triangular, internally finely tomentulose. Petals 1.5–2 cm long, externally pubescent, yellowish. Staminal column ca. 15 mm long, the filaments 1–2 mm long. Fruits ca. 1 cm diam., subequal to calyx, obscurely 10-costate; mericarps 5, subligneous. COLOMBIA. CH: Gentry 53856 (CUCV, MO). V: Killip & Cuatrecasas 38834 (CAS, US, the type collection); Gentry et al. 40414 (COL, MO). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 7252 (GUAY). Distribution: Southeastern Panama to northwestern Ecuador. Ecology: Often growing partially submersed in the intertidal zones at rivermouths and in mangroves (Fryxell, 1999), in the wet forest and rainforest life zones. Phenology: It has been collected in flower from December to May, as well as in August, and October; and in fruit mostly from January to June.

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Use: The leaves are pounded in a bit of water, to produce a dense, somewhat viscose liquid called “baba” (=saliva, Spanish), which is used to sift and purify a strong alcoholic drink named “guarapo”. IUCN Red list category: Due to factors such as a range of distribution of less than 2,000 km2 and a loss of habitat, it is herein categorized as Vulnerable (VU B2abiii). Etymology: The epithet refers to the Rhizophora (Rhizophoraceae) association of this species in mangroves, where the type species was found. Notes: Due to its nearly exclusive distribution in mangrove ecosystems, Pavonia rhizophorae has been regarded as a minor mangrove associated species (Chapman, 1975), albeit it does not have morphological adaptations. The type collection of P. rhizophorae was gathered by Ellsworth P. Killip and José Cuatrecasas in 1944, at sea level in El Forge near Buenaventura, Department of Valle del Cauca, Colombia. 137. Talipariti tiliaceum var. pernambucense (Arruda) Fryxell, Contr. Univ. Michigan Herb. 23: 262 (2001). Plate 28: B. Bas.: Syn.:

Hibiscus pernambucensis Arruda Paritium pernambucense (Arruda) G. Don Hibiscus tiliaceus var. pernambucensis (Arruda) I. M. Johnst. Hibiscus tiliaceus subsp. pernambucensis (Arruda) A. Cast. Hibiscus bracteosus DC. Hibiscus arboreus Desv. ex Ham. Hibiscus fragantissimus Sessé & Moc.

Vernacular names: majado, majagua. Trees or shrubs 1–8 m tall; minutely stellate-pubescent. Bark brown. Stipules 1.5–2.5 cm long, deciduous. Leaf blades coriaceous, broadly ovate, 7–15 x 7–15 cm, deeply cordate at base, acuminate at apex, entire or minutely denticulate at margins, markedly discolours, with 1–5 linear nectaries at base of principal veins beneath; petioles commonly subequal to blade; involucel 12–17 mm long; pedicels 0.5–3 cm long, solitary in leaf axils. Calyx dentate, ca. 2.5 cm long, with nectary on midrib of each calyx lobe. Petals 4–6 cm long, bright-yellow fading orange (in post-anthesis). Staminal column ca. 3 cm long. Capsules ca. 2 cm long, dehiscent, very light; seeds reniform, 4–5 mm long, minutely papillate. COLOMBIA. CH: Forero 4390 (COL). V: Devia et al. 2966 (MO). C: (E). N: Ramírez 1229 (NY). ECUADOR. E: Cornejo & Bonifaz 7247 (GUAY, QCNE). M: Bonifaz & Cornejo 3521-A (GUAY, TEX). SE: (S). G: Haught 3351 (NY). O: Cornejo & Bonifaz 5986 (GUAY, TEX). PERU. T: (E). Distribution: Neotropics. Ecology: Occurring in coastal habitats, on somewhat compact, muddy and/or sandy soils, sometimes forming pure stands in mangroves and on beaches, reaching the adjacent riverbanks. Present in very dry, dry, moist and wet forest life zones.

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Biological interactions: The ephemeral flowers have been observed to be occasionally visited by hummingbirds of the genus Amazilia, that feed on the nectar at the inner corolla base without touching the anthers. The flowers are also visited by nectar thieve bumble bees of the genus Bombus. In the Galapagos islands, the flowers of T. tiliaceum are pollinated by Carpenter bees of Xylocopa darwini. However, based on a bat pollination report in some species of Hibiscus (Fleming et al., 2009), which have a very similar, large floral structure to those of Talipariti, it cannot be discarded that the continental populations of T. tiliaceum may be pollinated by bats. The floral buds and linear nectaries located at the base of main veins on the abaxial side of blades are visited by domestic bees. The plants are frequently visited by insects of order Hemiptera. Finally, the leaves of T. tiliaceum var. pernambucense are one of the main food resources for the edible blue crab Cardisoma crassum, as has been observed in the Cangrejal de Olon mangrove in Ecuador. Dispersal: This is a water dispersed species. The very light capsular subglobose fruits, after dehiscence, turn to a little-basket-shape, exposing the small seeds. While aging, the dehisced fruits fall on the water and float for approximately five to ten days. The seeds themselves can also float on the water, for at least two to four days. The seeds have been picked up in open seas, ten to twenty miles from the mouth of the Gulf of Guayaquil (Guppy, 1906). In a transverse cut, the mature seeds show two aerial cavities, pressumably to aid in flotation; also, during the first to second day in the water, the numerous papillae on the outer seed coat (Fryxell, 2001; but not smooth as Tomlinson states, 1986) have been observed to collect many micro-bubbles from the air, probably adding to buoyancy from the outside. Uses: The fibrous bark countains a textil fiber used by natives to elaborate ropes and baskets. The fishermen use it to string fish (e.g., “lisas”, Mugil sp.) which are sold out in the streets. It is said that this species is used to heal the sting of the toadfish known as “pejesapo”, a mud-dwelling fish inhabiting the mangrove waters whose stings cause the flesh to rot. Due to its evergreen foliage and the showy flowers, T. tiliaceum var. pernambucense is cultivated as an ornamental tree/shrub in several countries in South America. It is highly recommended the use of this variety for reforestation and ornamentation of sandy beaches and coastal areas. IUCN Red list category: Due to factors such as a widespread range of distribution and a common presence of healthy populations, the categorization of this species and variety is herein proposed to be of Least Concern (LC). Etymology: The specific epithet refers to the slight resemblance of this species with those of the genus Tilia (Tiliaceae). The varietal epithet refers to the State of Pernambuco, Brazil, where that taxon was initially thought to be restricted. Notes: Talipariti tiliaceum var. pernambucense is easy to recognize by its large cordate leaves and beautiful, conspicuous yellow flowers. It sometimes forms pure stands constituting the Malvacetalia alliance in mangroves and beaches. In contrast to other mangrove forests, which are dominated by the major mangrove species, it is noteworthy that T. tiliaceum var. pernambucense is dominant in the Cangrejal de Olón, a small isolated mangrove forest directly facing the Pacific Ocean. The stands of this species are known as majagual (Acosta-Solís, 1944), a derivation of “majagua”, one of its vernacular names.

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The small village of Majagual from northwestern Ecuador owes its name to it. It is necessary to mention that in Ecuador the vernacular name majagua is also applied to Poulsenia armata, an unrelated tree of the Moraceae family that inhabits terra firme wet forests throughout the Neotropics. Discussion: Talipariti tiliaceum has been called a “semi-mangrove” (Tansley and Fritsch, 1905), but this term has been disregarded (Chapman, 1975). In this work, this species is proposed to be a facultative mangrove due to the following factors: the water dispersal adaptations of the fruits and seeds; the high tolerance to hypersaline environments; characteristic pattern of distribution along river water courses leading to mangroves and beaches; and the tendency to occasionally form pure stands in the interior of the true mangrove habitats. MARCGRAVIACEAE Lianas or shrubs, terrestrial or hemiepiphytic. Stems often slender, ± scrambling. Stipules absent. Leaves alternate or spiral; the blade simple, bearing glands abaxially. Inflorescences terminal, sometimes cauliflorous or in short lateral shoots, racemose or pseudoumbels; extrafloral bracteal nectaries present; bracteoles 2, generally sepal-like. Flowers actinomorphic, bisexual. Calyx persistent, the sepals 4 or 5, imbricate, quincuncial. Petals (3–)4–5(–6), distinct, somewhat connate or fused in a calyptra, imbricate. Stamens 3–100. Gynoecium syncarpous; ovary superior, 2–20-carpelled; style 1; stigma lobed or umbonate. Fruit capsular, (sub)globose, the pericarp rather coriaceous, splitting to expose seeds embedded in pulp-like placentae. This predominantly epiphytic Neotropical family is restricted to moist forest, wet forest, and rainforest life zones (Giraldo-Cañas, 2007). The (sub)globose fruits of Marcgraviaceae, subtended by elongate subwoody terminal peduncles, dehisce at maturity. The numerous small exposed seeds have been reported to be eaten and endozoochorously dispersed by bats and birds (Lobova et al., 2009). Key to the species 1. Leaf blades slightly cordate at base; rachis of inflorescences 22–30 cm long; pedicels 4–6(–7) cm long; stamens ca. 50 Schwartzia chocoensis 1. Leaf blades usually cuneate at base; rachis of inflorescences (7–)8–20 cm long; pedicels 1–3(–3.5) cm long; stamens 5 or ca. 14 2. Leaf blades revolute at margins, the lateral veins inconspicuous beneath; nectaries attached to rachis; stamens ca. 14 Pseudosarcopera diaz-piedrahitae 2. Leaf blades not revolute at margins, the lateral veins prominent beneath; nectaries attached to pedicels; stamens 5 3. Leaf blades 6–8 cm wide; petioles 10–13 mm long; pedicels 2–3 cm long Souroubea cf. guianensis 3. Leaf blades 2.5–5 cm wide; petioles 4–8 mm long; pedicels 1–2 cm long Souroubea intermedia

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138. Pseudosarcopera diaz-piedrahitae (Giraldo-Cañas) Giraldo-Cañas, Caldasia 29(2): 206, fig. 1 (2007). Bas.: Schwartzia diaz-piedrahitae Giraldo-Cañas Vernacular names: mangosté, manguté. Holoepiphytic to hemiepiphytic scandent shrubs. Leaves spiral; blades coriaceous, elliptic to oblong, (3–)5–12.5(–14) x (1.3–)1.8–4.5(–6.5) cm, cuneate at base, slightly apiculate at apex, revolute and thick at margins, with 0 to 6 pairs of glands beneath, these arranged 6–10 mm from margins, the lateral veins inconspicuous, glabrous; petioles short. Inflorescences terminal, densely racemose, the rachis (7–)8–11 cm long; pedicels (1.6–)2.2–2.8(–3.5) cm long, spirally arranged. Nectaries attached to rachis, just beneath each pedicel. Flowers 24 to 54. Sepals 5, imbricate, 2-seriate, orbicular, fleshy, 2–2.5 mm long. Petals 5, basally connate, ovate, fleshy, reflexed, 5–7 x 4–5 mm. Stamens 14, the filaments 2.4–2.7 mm long, adnate to petal bases. Ovary superior. Stigma subsessil. Fruits fleshy globose, apiculate, 8–11 mm diam., light green. Seeds 10–15, 2–3 mm long. COLOMBIA. CH: (E). V: Gentry 53423 (MO). Distribution: Western Colombia and northwestern Ecuador. Ecology: In rainforests. Biological interactions and dispersal: No information is recorded, but most likely after the fruits dehisce, birds and bats feed on the seeds and surrounding tissues, digest the fleshy funicles and/or placentas, and disperse the seeds endozoochorously (Lovoba et al., 2009). Use: As a depurative remedy. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat it is herein categorized as Near Threatened (NT). Etymology: The epithet honors Professor Santiago Díaz Piedrahita (COL), a Colombian botanist and taxonomist of the Asteraceae family (Giraldo-Cañas, 2001). 139. Schwartzia chocoensis Giraldo-Cañas, Revista Acad. Colomb. Ci. Exact. 25(97): 478, fig. 1 (2001). Holoepiphytic to hemiepiphytic scandent shrubs. Leaves spiral; blades coriaceous, elliptic to oblong, (12–)15–22 x (3.5–)5.5–7.8(–8.2) cm, slightly cordate at base, shortly apiculate at apex, slightly revolute and thick at margins, with 2 to 7 pairs of glands beneath, these arranged 0.5–1.5(–3) mm from margins, the lateral veins weak on both sides, glabrous; petioles short. Inflorescences terminal, densely racemose, the rachis 22–30 cm long; pedicels 4–6(–7) cm long, spirally arranged. Nectaries sacciform, attached to pedicels, pendulous, reddish. Flowers (5–)20 to 54. Sepals 5, imbricate, 2-seriate, orbicular, fleshy, 2–2.5 mm long. Petals 5, basally connate, ovate, fleshy, reflexed, 6–12(–13) x 3.8–6 mm. Stamens ca. 50, the filaments 1–3 mm long, adnate to petal bases. Ovary superior. Stigma subsessil. Fruits fleshy globose, apiculate, 14–17 mm diam., light green. Seeds ca. 30, 3–4 mm long.

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COLOMBIA. CH: (E). V: Gentry & Juncosa 40423 (COL, MO). Distribution: Western Colombia and northwestern Ecuador. Ecology: In rainforests. Biological interactions and dispersal: No information is recorded, but most likely after the fruits dehisce, birds and bats feed on the seeds and surrounding tissues, digest the fleshy funicles and/or placentas, and disperse the seeds endozoochorously (Lovoba et al., 2009). IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the Choco region, where the species occurs (Giraldo-Cañas, 2001). 140. Souroubea guianensis Aubl., Hist. Pl. Guian. 1: 244, t. 97 (1775). Holoepiphytic to hemiepiphytic lianas. Leaves spiral; blades coriaceous, elliptic to obovateelliptic, 10–15 x 6–8 cm, cuneate at base, usually obtuse at apex, the lateral veins prominent, glabrous beneath; petioles 10–13 mm long. Inflorescences terminal, loosely racemose, the rachis ca. 10–15 cm long; pedicels 2–3 cm long, spirally arranged. Flowers not seen. Fruits fleshy globose, apiculate, ca. 20–25 mm diam., light green. ECUADOR. E: Cornejo & Bonifaz 7238 (GUAY, QCNE). Distribution: Widely distributed in tropical South America. Ecology: In rainforests. Biological interactions and dispersal: No information is recorded, but most likely after the fruits dehisce, birds and bats feed on the seeds and surrounding tissues, digest the fleshy funicles and/or placentas, and disperse the seeds endozoochorously (Lovoba et al., 2009). IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the French Guiana, where it was initially known. 141. Souroubea intermedia de Roon, Acta Bot. Neerl. 18(3): 401, fig. 1 (1969). Holoepiphytic to hemiepiphytic shrubby lianas. Leaves spiral; blades coriaceous, asymmetric, obovate-oblong to oblong, or sometimes lanceolate, 7–14 x 2.5–5 cm, cuneate at base, subacute to obtuse and slightly apiculate at apex, with scattered minute mostly submarginal glands beneath, the lateral veins prominent, glabrous beneath, glabrous; petioles 4–8 mm long. Inflorescences terminal, loosely racemose, the rachis 10–20 cm long; pedicels 1–2 cm long, spirally arranged. Nectaries auriculate, attached to pedicels. Sepals 5, imbricate, transverse-ellictic or orbicular, ca. 1.5 mm long. Petals 5, oblong, 4–6 x 2–3 mm. Stamens 5, the filaments 3–3.5 mm long. Ovary superior. Stigma sessil. Fruits unknown.

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COLOMBIA. CH: Killip & Cuatrecasas 39146 (COL, MO, the type collection). Distribution: Western Colombia and northwestern Ecuador. Ecology: In rainforests. Biological interactions and dispersal: No information is recorded, but most likely after the fruits dehisce, birds and bats feed on the seeds and surrounding tissues, digest the fleshy funicles and/or placentas, and disperse the seeds endozoochorously (Lovoba et al., 2009). IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). MELASTOMATACEAE (Melastome Family) Trees, shrubs, vines, herbs, sometimes epiphytic. Leaves simple, decussate, usually plinerved. Inflorescences terminal or lateral, usually cymose. Flowers regular; hypanthium bearing sepals, petal and stamens on a torus at or near base of calyx lobes. Petals free, usually white to purple, right-contort, usually spreading. Stamens usually twice as many as petals, usually more or less dimorphic at least in size, the anthers opening by 1–2(–4) terminal pores. Ovary superior or inferior; stigma punctiform to capitate; style 1. Fruit a loculicidal capsule enclosed by persistent hypanthium or a berry; seeds usually numerous. Key to the species 1. Terminal branches sharply quadrangular 2. Inflorescences terminal, the flowers many, ca. 1 cm diam., sessile, the filaments strongly curved inwards; fruit baccate

Miconia reducens

2. Inflorescences axillary, the flowers 2–4, ca. 5 cm diam., pedicellate, the filaments erect; fruit capsular Blakea glabrescens 1. Terminal branches subterete or somewhat compressed 3. Petioles without domatia

Conostegia polyandra

3. Petioles with domatia Tococa guianensis 142. Blakea glabrescens Benth., Bot. Sulph. 94 (1844). Plate 28: C. Hemiepiphytic shrubs to 6 m tall. Terminal branches sharply quadrangular. Buds covered by dense brown indumentum. Leaf blades chartaceous, elliptic, 10–15 x 6–10 cm, obtuse at base, rather abruptly and shortly acuminate at apex, covered by sparsely brown indumentum beneath, 5-nerved with secondary veins ca. 1 mm apart; petioles ca. 3 cm long, vaginate. Inflorescences axillary, the flowers 2–4 per node; pedicels 6–20 mm long; bracts free, multinerved. Petals 6, oblongoid, ca. 2.5 cm long, white, tinged purplish at tips, blunt at apex. Stamens 12. Style pink; stigma elliptic-capitate, white. Fruit not seen.

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COLOMBIA. CH: (E). V: (E). C: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 6955 (AAU, GB, GUAY). Distribution: Western Colombia to northwestern Ecuador. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the glabrescent condition of the leaf blades. 143. Conostegia polyandra Benth., Bot. Sulph. 96, pl. 35 (1844). Plate 28: D. Syn.: Miconia rupicola Gleason Vernacular name: mora. Shrubs 1–4 m tall. Terete branchlets, primary leaf veins beneath, inflorescences, and hypanthia (basally) sparsely and caducously furfuraceous with stellulate hairs. Leaf blades thin coriaceous, elliptic to oblong-elliptic, 5–15 x 2.5–5.5 cm, broadly acute to obtuse at base, blunty acute to rounded at apex, distantly serrulate, glabrous, 3-nerved (excluding the tenuous marginals) or shortly pseudo-plinerved; petioles 0.5–2 cm long. Inflorescence a terminal panicle, 4–14 cm long, few-flowered; pedicels 3–6 mm long; bracteoles 3–4 mm long, inserted near middle of pedicels. Hypanthium 4–4.5 mm long. Petals (5– )6–7, obovate, 8–11 x 7–8 mm, white. Stamens 26–33, the filaments 3–3.5 mm long. Stigma not expanded. Fruit baccate, globose, dark-purple at maturity; seeds numerous. COLOMBIA. CH: Fuchs et al. 22133 (COL, MO). V: Otero 154 (CUVC). C: (E). N: Murphy & Parra 719 (MO). ECUADOR. E: Bonifaz & Cornejo 4013 (AAU, GB, GUAY, QCA). Distribution: Nicaragua to Ecuador. Ecology: In wet forests and rainforests, often growing behind sandy beaches. Phenology: This species has been collected in flower and fruit throughout the year. Biological interactions and dispersal: It has been observed that wasps feed on the juicy mesocarp. No further information is recorded, but most likely female buzzing bees pollinate the flowers, while birds and presumably bats eat the small baccate fruits, digest the mesocarp, and disperse the tiny seeds endozoochorously (Kriebel, pers. comm., Lobova et al., 2009). Use: The juicy pulp of the fruit is edible. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet refers to the many stamens borne by the flowers. 144. Miconia reducens Triana, Trans. Linn. Soc. Bot. 28: 106 (1871). Plate 29: A. Shrub or small tree to 5 m tall (also recorded as epiphytic in other places inland). Branches sharply 4-angled, with a prominent interpetiolar ridge at nodes. Leaf blades oblong-elliptic, 18–30 x 6–12 cm, subauriculate at base, acute or short-acuminate at apex, on both sides

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glabrous, shortly 5-plinerved; petioles 2–3 cm long, reddish. Inflorescence a terminal panicle, 10–20 cm long; flowers sessile. Hypanthium ca. 3.4 mm long. Petals 5, oblong, 6–6.5 x 2.5–3 mm, lilac or pink. Stamens ca. 9, the filaments strongly curved inwards. Stigma capitellate, ca. 0.9 mm diam. Fruit baccate, globose; seeds numerous. COLOMBIA. CH: (E). V: Gentry et al. 53405 (MO). C: Cabrera 3262 (CUVC). N: (S). ECUADOR. E: Játiva & Epling 2111 (MO, NY, S, U, US). Distribution: Southern Mexico to Venezuela, and northwestern Ecuador. Ecology: In wet forests and rainforests. Biological interactions and dispersal: No information is recorded, but most likely insects pollinate the flowers, and birds eat the small baccate fruits, digest the mesocarp, and disperse the tiny seeds endozoochorously. IUCN Red list category: Least Concern (LC). 145. Tococa guianensis Aubl., Hist. Pl. Guiane 1: 438, t. 174 (1775). Shrub or small tree to 5(–7) m tall. Stems glabrous, sparsely glandular-pubescent or with a mix of glandular and no-glandular setae. Leaves anisophyllous, seldom isophyllous; blades ovate to oblong-elliptic, the larger blades (5.5–)8–30(–34) x (4–)5–15(–18) cm, the smaller leaves 3.5–10 x 2.5–7 cm, rounded to acute at base, acute to acuminate, caudate or aristate at apex, usually apressed-setose above, 5–(7–)plinerved; petioles 0.5–2 cm long , glandularsetose, the domatia globose, ovoid, or narrowly-ellipsoid (1.5–)2–3(–4.5) x (0.7–)1–2(–3) cm, sometimes present only in larger leaves. Inflorescence a terminal panicle, 5–15(–21) cm long; pedicels 3–9 mm long. Hypanthium 5–7.5 mm long. Petals 5, broadly oblong to ovate, (5.5–)6.5–15 x (3–)4.5–7.5(–9) mm, pink. All stamens of similar length, the filaments 5.5–8.5 mm long. Stigma funnelform, ca. 1.5 mm diam. Fruit baccate, globose, black at maturity; seeds numerous. COLOMBIA. V: (E). Distribution: Southern Mexico to Bolivia. Ecology: In rainforests. Biological interactions and dispersal: No information is recorded, but most likely insects pollinate the flowers, and birds eat the small baccate fruits, digest the mesocarp, and disperse the seeds endozoochorously. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the French Guiana, where it was initially known to occur. Note: This species is included herein based on a report by Cuatrecasas (1958b). However, herbarium collections of Tococa guianensis from mangroves have not been seen (Michelangeli pers. comm., 2010).

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MORACEAE (Mulberry Family) Trees or shrubs, rarely herbaceous, terrestrial or hemiepiphytic stranglers. Milky sap present. Stipules fully amplexicaul to lateral, free or fused. Leaves alternate, in spirals or distichous, petiolate; blade simple, pinnately to subpalmately veined. Inflorescences bisexual or unisexual, racemose, spicate, globosecapitate, discoid-capitate and then often involucrate or urceolate, many- to one-flowered, with or without bracts. Flowers unisexual free or connate or adnate to receptacle. Staminate flowers: tepals 4, stamens 4. Pistillate flowers: tepals 4, pistil 1, stigmas 2. Fruit an achene or more or less drupaceous, or forming a drupaceous structure with enlarged and fleshy perianth, or forming drupaceous or syncarpous structures. This family, widely distributed in the tropics and subtropics, is represented only by the genus Ficus. All species herein recorded are stranglers of trees of red mangrove (Rhizophora spp.), nato (Mora oleifera), and bambudo (Pterocarpus officinalis), in moist to pluvial life zones. The strangler species of Ficus play an important role in the sucession of the terra firme-mangrove vegetation. The inflorescences (figs) of Ficus have been reported to be eaten and the seeds endozoochorously dispersed by birds, howler monkeys (Alouatta spp.), and many species of bats (Lobova et al., 2009). Key to the species 1. Leaf blades cordate to cordate-ovate, the base cordate, the basal lobes overlapping Ficus nymphaeifolia 1. Leaf blades ovate-elliptic to lanceolate, elliptic or obovate, the base cuneate to rounded, the basal lobes if present then never overlapping 2. Terminal stipules densely pubescent to pilose, at least at base; figs 0.5–1 cm diam. 3. Terminal stipules 0.8–1.3 cm long; leaf blades with 6–9 lateral veins per side Ficus brevibracteata 3. Terminal stipules 1.5–3 cm long; leaf blades with ca. 12–15 lateral veins per side Ficus jacobii subsp. mantana 2. Terminal stipules glabrous; figs 1–2.5 cm diam. 4. Leaf blades elliptic to broadly lanceolate, scabrous beneath, the petioles with epidermis flaking off; basal bracts 1–1.5 mm long Ficus maxima 4. Leaf blades broadly obovate, smooth beneath, the petioles with epidermis persistent; basal bracts 4–10 mm long Ficus obtusifolia 146. Ficus brevibracteata W. C. Burger, Phytologia 26(6): 423 (1973). Syn.: Ficus hartwegii var. tumacana Dugand Vernacular name: matapalo.

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Hemiepiphytic stranglers. Terminal branches tomentulose, glabrescent. Terminal stipules 0.8– 1.3 cm long, densely pubescent. Leaf blades coriaceous, ovate-elliptic to ovate-lanceolate, 11–25 x 6–12 cm, truncate to rounded at base, shortly-acuminate at apex, the lateral veins 6–9 per side, pilose beneath; petioles 2–10 cm long, pilose, glabrescent. Figs (sub)globose, 0.5–1 cm diam., light-brown with white spots, tomentose; sessile; basal bracts 2–3 mm long, pubescent. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6950 (BG, GUAY). Distribution: Costa Rica to western Ecuador. Ecology: In moist forests, wet forests, and rainforests. Biological interactions and dispersal: No observations are recorded, but most likely bats, and some birds such as parrots and parakeets (Psittacidae), feed on the figs and disperse the seeds. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the short bracts of figs. 147. Ficus jacobii subsp. mantana C. C. Berg, Blumea 52: 576 (2007). Vernacular name: bu’machi (Cayapa), matapalo. Hemiepiphytic stranglers. Terminal branches glabrous. Terminal stipules 1.5–3 cm long, ± densely pilose at base on abaxial side. Leaf blades coriaceous, narrowly obovate, 15–30 x 8–13 cm, inconspicuosly subcordate at base, distinctively acuminate at apex, densely pilose beneath, mainly on veins, the lateral veins ca. 12–15 per side; petioles 1.5–8 cm long, pilose, glabrescent. Figs globose, 0.6–1 cm diam., densely minutely white-puberulous to subglabrous; pedicels 1–3 mm long; basal bracts 2, 2–2.5 mm long, puberulous. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 6963 (AAU, BG, GUAY). Distribution: Western Ecuador. Ecology: In dry, moist, and wet forests. Biological interactions and dispersal: No observations are recorded, but most likely bats, and some birds such as parrots and parakeets (Psittacidae), feed on the figs of this subspecies and disperse the seeds. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The subspecific epithet refers to Manta, the type locality from Manabi, in western Ecuador. 148. Ficus maxima Mill., Gard. Dict. ed. 8 (1768). Syn.: Ficus radula Humb. & Bonpl. ex Willd. Ficus glaucescens (Liebm.) Miq. Ficus hernandezii (Liebm.) Miq.

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Ficus parkeri Miq. Ficus subscabrida Warb. Ficus bopiana Rusby Ficus sodiroi Rossberg Ficus murilloi Dugand Ficus vicencionis Dugand Ficus chaconiana Standl. & L. O. Williams

Vernacular name: matapalo. Hemiepiphytic stranglers. Terminal branches glabrous. Terminal stipules 1.5–3 cm long, glabrous. Leaf blades coriaceous, elliptic to broadly lanceolate, 9–30 x 4.5–12 cm, cuneate to obtuse at base, acute to acuminate at apex, entire, scabrous beneath, glabrous, the lateral veins 13–17 per side; petioles 1–7 cm long, the epidermis flaking off, glabrous. Figs globose, 1–2 cm diam., green, glabrous; pedicels 4–10 mm long; basal bracts 1–1.5 mm long, glabrous. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6968 (BG, GUAY, QCA). Distribution: Mexico to Paraguay. Ecology: In moist forests, wet forests, and rainforests. Biological interactions and dispersal: For a detailed list of bats that may disperse this species in our area see Lobova et al. (2009). Some birds such as parrots and parakeets (Psittacidae), as well as howler monkeys (e.g., Alouatta palliata) may also feed on the figs, dispersing the seeds. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the large size of trees. 149. Ficus nymphaeifolia Mill., Gard. Dict. (ed. 8) Ficus no. 9 (1768). Plate 29: B. Syn.:

Ficus nymphoides Thunb. Ficus urbaniana Warb. Ficus involuta var. urbaniana (Warb.) Dugand Ficus ierensis Britton Ficus anguina Benoist Ficus pascuorum Pittier Ficus duquei Dugand Ficus duquei var. obtusiloba Dugand

Vernacular name: matapalo. Hemiepiphytic stranglers. Terminal branches lenticellate, glabrous. Terminal stipules 1.5–4 cm long, glabrous. Leaf blades coriaceous, cordate to cordate-ovate, 17–30 x 13–18 cm, cordate at base, the basal lobes overlapping, shortly acuminate at apex, glabrous, 8–12 lateral veins per side; petioles 5–20 cm long, glabrous. Figs (sub)globose, 1.5–2.5 cm diam., velutine; pedicels 2–4 mm long; basal bracts 4–9 mm long, glabrous.

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COLOMBIA. CH: (E). V: (E). C: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 6948 (BG, GUAY, QCA). Distribution: Belize to Bolivia. Ecology: In dry forests, moist forests, wet forests, and rainforests. Biological interactions and dispersal: For a detailed list of bats that may disperse this species in our area see Lobova et al. (2009). Some birds such as parrots and parakeets (Psittacidae), as well as howler monkeys (e.g., Alouatta palliata) may also feed on the figs, dispersing the seeds. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the resemblance of the heart-shaped leaves of this species with those of Nymphaea (Nymphaeaceae, the Water lilies family). 150. Ficus obtusifolia Kunth, Nov. Gen. Sp. (quarto ed.) 2: 49 (1817). Plate 29: C. Syn.: Ficus bonpladiana (Liebm.) Miq.

Ficus gardneriana (Miq.) Miq.

Ficus involuta (Liebm.) Miq.

Ficus proctor-cooperi Standl.

Ficus floresina Pittier

Ficus mattogrossensis Standl.

Ficus chiapensis Lundell

Vernacular name: matapalo. Hemiepiphytic stranglers. Terminal branches glabrous or pubescent. Terminal stipules 1–3 cm long, glabrous. Leaf blades coriaceous, broadly obovate, 7–25 x 6–15 cm, cuneate at base, rounded to blunty acute at apex, entire, smooth and glabrous on both sides, the lateral veins 6–12 per side; petioles 1–4 cm long, glabrous. Figs globose, 1.5–2.5 cm diam., green, glabrous; pedicels 5–8 mm long; basal bracts 2, 4–10 mm long, semicircular, connate, finely pubescent. COLOMBIA. CH: (E). V: (E). C: (E). N: (E). ECUADOR. E: (S). Distribution: Mexico to Bolivia. Ecology: In dry forests, moist forests, wet forests, and rainforests. Biological interactions and dispersal: The fruits are predated and the seeds dispersed by bats. It is most likely that some birds such as parrots and parakeets (Psittacidae), and howler monkeys (e.g., Alouatta palliata) also feed on the figs, dispersing the seeds. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet refers to the widely obtuse apex of the leaf blades.

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MYRSINACEAE Trees or shrubs. Leaves alternate, spiral or subverticillate, often clustered at end of branches, entire, crenulate or serrate, glandular-punctate. Stipules absent. Flowers bisexual or unisexual, usually 4- or 5-parted, regular, small, white, pink or green, in terminal and axillary mostly pedunculate inflorescences, or glomerate on short bract-covered axillary shoots. Calyx mostly ciliate and glandular-punctate, valvate, imbricate, or siniestrorsely contorted, persistent. Corolla regular, usually gamopetalous, rotate to tubular, the petals valvate, imbricate or contorted. Stamens as many as petals and opposite them. Ovary globose, ovoid or clavate, free, sessile, 1-loculed, the ovules numerous or few. Fruit drupaceous, 1-seeded. 151. Ardisia granatensis Mez, Pflanzenr. 4: 236: 86 (1902). Shrubs or small trees to 4 m tall. Leaves loosely spiral; blades chartaceous, oblong to oblongelliptic, 9–16 x 2.5–5 cm, cuneate at base, acuminate at apex, entire and revolute at margins, glandular-punctate and glabrous beneath; petioles 3–6 mm long. Inflorescences terminal, paniculate, subsessile, congested, much shorter than leaves, to 5 cm long, essentially glabrous, with conspicuous cucullate bracts enveloping the mature buds; pedicels 0–4.5 mm long, glabrous. Calyx ca. 3 mm long, greenish, the sepals 5, free, punctate with short oblongish glands. Petals 5, narrowly ovate-elliptic or oblong-elliptic, ca. 7 mm long, connate at base, rounded at apex, sparsely punctate. Stamens ca. 5 mm long. Ovary ovoid, superior. Fruit subglobose, punctate, ca. 7 mm in diam; seeds 1. COLOMBIA. V: Killip 34984 (COL, F, GH). Distribution: Southeastern Panama to western Colombia. Ecology: It occurs in the rainforest life zone, from the terra firme ecosystems to the fresh water and brackish swamps, estuarine ecosystems, and mangroves to the intertidal zones. Biological interactions: No information is recorded, but most likely insects pollinate the flowers, and birds feed on the fruits and disperse the seeds. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the Viceroyalty of New Granada, the Spanish colonial jurisdiction in northwestern South America which subsequently became the separate Republics of Colombia, Venezuela, and Ecuador. NYCTAGINACEAE (Four-o’clock Family) Trees, shrubs, lianas or herbs. Leaves simple, entire, generally opposite, sometimes alternate. Stipules absent. Inflorescences terminal or axillary, the flowers normally bisexual, actinomorphic, subtended by bracts. Perianth usually colored and petaloid but apetalous, uniseriate, the segments 5, at least basally united, often forming an elongate tube, the base of perianth persisting around fruit. Stamens usually 5,

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often unequal. Pistil 1; ovary superior. Fruit a fleshy, coriaceous or woody indehiscent achene or nut, often enclosed within persistent indurated base of perianth to form an anthocarp. 152. Cryptocarpus pyriformis Kunth, Nov. Gen. Sp. 2: 188, pl. 124 (1817). Plate 29: D. Syn.:

Cryptocarpus cordifolius Moric.

Salpianthus pyriformis (Kunth) Standl.

Vernacular names: chope, manglillo salado, monte salado, palo salado, salado. Lianas, sometimes resembling scandent shrubs with sprawling, pendulous branches. Leaves alternate; blades reniform, cordate or deltoid to suborbicular, 1.2–13 x 1.5–12 cm, cordate at base, widely obtuse to emarginate at apex; petioles to 6 cm long. Inflorescences of axillary and subterminal racemes and terminal panicles, arranged toward end of branches, 1–2(–3.5) cm long, white-tomentulose; pedicels to 1 mm long. Calyx ca. 2 mm long, greenish, the sepals 5, divergent at anthesis. Filaments exserted, ca. 3 mm long, white, the anthers medifixed. Fruit subglobose, longitudinally 5-sulcate, ca. 3 mm diam., yellowish. ECUADOR. M: (S). SE: (S). G: Cornejo & Castro 242 (GUAY). O: Asplund 15878 (NY). GAL: Triviño & Cruz 98 (GUAY). PERU. T: Smith 10081 (MO). P: (E). Distribution: Ecuador and Peru, also in the Galapagos Islands. Ecology: Cryptocarpus pyriformis seems to be restricted to the coastal dry thorn scrubs, very dry forests, and dry deciduous forests. It often grows in the mangrove fringe, and in the mangrove interior when disturbed. It also occurs on dunes, sandy beaches, and landwards, to the contiguous terra firme ecosystems. Biological interactions and dispersal: During the day, the flowers have been observed to be visited by flies, bees, butterflies and wasps. The small subglobose fruits are most likely eaten by birds, while the seeds are endozoochorously dispersed. The whole plant is eaten by goats and cattle. Uses: Medicinally employed to treat tooth aches, colds, aches, and pains (Madsen et al., 2001). IUCN Red list category: This species is persistent in secondary forests and in heavily disturbed habitats. It is here categorized as of Least Concern (LC). Etymology: The epithet refers to the pear-like shape of fruits, a variable feature. ORCHIDACEAE (Orchid Family) Terrestrial or epiphytic herbs, rarely vines. Rhizomes delicate to stout, sometimes creeping or condensed. Stems a corm, pseudobulb, ramicaul, or absent, sometimes proliferous. Leaves alternate or distichous, seldom opposite or whorled, sometimes all basal, or reduced to a mere scales, simple, entire, usually parallel-veined. Inflorescences racemose, cymose, spicate or paniculatae, 1-many-flowered, erect or variously pendulous. Flowers bisexual or sometimes unisexual (Catasetum), generally zygomorphic.

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Perianth typically 6 tepals in 2 whorls, all petaloid. Petals 3, the median one strongly differenciated from the others, commonly larger or differing in form and color, forming a lip (labelum). Stamens usually 1, adnate to style, forming a column. Fruit capsular; seeds tiny, numerous, dust-like, wind-dispersed.

Orchidaceae comprises a significant percentage of the associate flora in the mangroves located

in the wet forest and rainforest life zones. With the exception of Elleanthus discolor, the remaining species of this family that occur in mangroves are epiphytic, mainly growing upon the aerial roots and branches of Rhizophoras. The dust-like seeds are wind dispersed. Key to the species 1. Pseudobulb obvious 2. Pseudobulb subcylindric, 7–25 cm long 3. Pseudobulb near apex with short spines from sheaths; leaf blades thin, with obvious thickened, longitudinal veins beneath; inflorescence basal-lateral; capsule 6–9 cm long Catasetum macroglossum 3. Pseudobulb near apex without short spines from sheaths; leaf blades stiffly-coriaceous, the veins inconspicuous; inflorescence apical; capsule ca. 3 cm long 4. Inflorescence a raceme; sepals and petals 7–8 mm wide, the lip irregularly-lanceolate Caularthron bilamellatum 4. Inflorescence a panicle; sepals and petals 3–5 mm wide, the lip orbicular Epidendrum bracteolatum 2. Pseudobulb ovoid to elliptic or shortly-lanceolate, to 5 cm long 5. Leaf blades elliptic to oblong-elliptic, 4–7 cm wide, purple beneath, with obvious thickened, longitudinal veins

Soterosanthus shepheardii

5. Leaf blades lorate or oblong, 0.3–3.5 cm wide, green beneath, the veins inconspicuous 6. Rhizome elongated, ca. 25–50 cm long; inflorescence fasciculate, the flowers bright-orange Ornithidium fulgens 6. Rhizome short; inflorescence a solitary flower, raceme or panicle, the flowers red to purple, greenish, yellowish, cream, or white 7. Inflorescence pendulous, a many-flowered raceme Notylia replicata 7. Inflorescence erect, a solitary flower or a few-flowered raceme, or a many-flowered panicle 8. Inflorescence basal-lateral, a solitary flower 9. Inflorescence several to many times longer than pseudobulb 10. Leaf blades elliptic, ca. 3–5 cm wide, cuneate at base; scape 4–7 cm long Maxillaria hennisiana 10. Leaf blades lorate, ca. 1 cm wide, attenuate at base; scape ca. 15–20 cm long Trigonidium seemannii

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9. Inflorescence of similar length than pseudobulb Camaridium atrovinaceum 8. Inflorescence apical, a raceme or panicle 11. Inflorescence a many-flowered panicle, 10–45 cm long; flowers ca. 3 mm diam. Polystachya concreta 11. Inflorescence a 3–8-flowered raceme, 3–10 cm long; flowers 0.7–4 cm diam. 12. Plants to ca. 30 cm tall; leaf blades 13–18 x 2–4 cm; flower ca. 4 cm diam.; capsule 3-winged Prostechea fragrans 12. Plants to ca. 10 cm tall; leaf blades ca. 5 cm x 3 mm; flower ca. 7 mm diam.; capsule wingless Prostechea pseudopygmaea 1. Pseudobulb absent or if present, then not obvious 13. Stem absent; leaves fan-like arranged Ornithocephalus bryostachys 13. Stem present; leaves usually distichous along stem, or 1–2 from stem apex 14. Leaves 1–2 from stem apex 15. Leaf blades ovate to elliptic or oblong-elliptic; stems wiry, less than 2 mm thick; flowers minute, less than 4 mm diam. 16. Stem 0.4–2 cm long; inflorescences exceeding leaves, the rachis flexuose 17. Leaf blades 4–8 x 3–5 mm, the stems 0.4–0.7 cm long; flowers yellow Platystele cornejoi 17. Leaf blades 15–20 x 6–9 mm, the stems 1.5–2 cm long; flowers purple-red Sarcinula sp. 16. Stem 3–8 cm long; inflorescences shorter than- and protected by- leaves, the rachis straight Trichosalpinx orbicularis 15. Leaf blades lorate; stems 2–5 mm thick; flowers wider than 4 mm diam. 18. Leaf blades 10–17 mm wide; flowers showy; capsules ca. 5–6 cm long Brassavola grandiflora 18. Leaf blades 1–7 mm wide; flowers inconspicuous; capsules 0.5–2 cm long 19. Lip united to apex of columm Epidendrum stangeanum 19. Lip free from apex of columm 20. Plants to 25 cm tall; leaves 3–7 cm long; flowers white; mature capsule 0.5–0.8 cm long 21. Stems bearing 2 leaves at apex, the blades expanded, straight Scaphyglottis prolifera 21. Stems bearing 1 leaf blade at apex, the blades somewhat conduplicate, curved Scaphyglottis sp. nov. 20. Plants 20–60 cm tall; leaves 8–25 cm long; flowers yellow, greenish or brown; mature capsule 1.2–1.5 cm long 22. Stems bearing 2 leaf blades at apex; inflorescences short, to 1.5 cm long, the flowers yellow Scaphyglottis graminifolia 22. Stems bearing 1 leaf blade at apex; inflorescences elongate, 5–15 cm long, the flowers brown to greenish Scaphyglottis cruigera

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14. Leaves several, distichous, arranged along stem 23. Scrambling plants 24. Main stem longer than 50 cm; roots not produced between leaves; leaf blades semiterete, 2–3 mm wide; inflorescences terminal; flowers without a spur produced from lip base Epidendrum microphyllum 24. Main stem shorter than 25 cm; roots produced from between leaves; leaf blades fully expanded, 6–13 mm wide; inflorescences opposite or subopposite to leaves; flowers with a spur produced from lip base Campylocentrum ecuadorense 23. Not scrambling plants 25. Stem 50–200 cm long; leaves with conspicuous longitudinal veins beneath 26. Leaf blades 7–10 cm wide; inflorescence solitary to few flowers, the flowers conspicuous, mostly white 27. Leaf blades 7–8 cm wide Sobralia madisonii 27. Leaf blades 8–10 cm wide Sobralia rhizophorae 26. Leaf blades up to 7 cm wide; inflorescence a densely many-flowered spike, the flowers small, bright orange Elleanthus discolor 25. Stem usually to 40 cm long (to 70 cm long only in Epidendrum macroophorum); leaves with inconspicuous veins beneath 28. Leaf blades articulated with sheaths; inflorescences produced at nodes along stem 29. Plants ca. 25–50 cm long; leaf blades 6–10 cm long; pedicels ca. 2.2 cm long Dichaea trulla 29. Plants to 20 cm long; leaf blades 1–3 cm long; pedicels ca. 1–1.3 cm long Dichaea panamensis 28. Leaf blades not articulated with sheaths; inflorescences produced at apex of stem 30. Plants erect or suberect 31. Leaves densely arranged from a short stem; inflorescence usually a panicle, to 45 cm long; flowers ca. 3 mm diam., the column free from lip Polystachya concreta 31. Leaves loosely arranged along a developed stem; inflorescence always a raceme, to 15 cm long; flowers 0.8–10 cm diam., the column fused to lip 32. Leaf blades lorate; rachis of inflorescence to 1 cm long; flowers pink Dimerandra rimbachii 32. Leaf blades oblong to elliptic; rachis of inflorescence 1.5–15 cm long; flowers yellowish and white, or green 33. Rachis of inflorescence 5–15 cm long; sepals ca. 4 mm long; petals ca. 0.4 cm long; lip undivided, ca. 0.4 cm long; capsule ca. 2 cm long Epidendrum rigidum 33. Rachis of inflorescence to 3 cm long; sepals 1.2–8 cm long; petals 1.1–7 cm long; lip 3-lobed, 0.8–5.7 cm long; capsule 4–15 cm long 34. Sepals 1.2–1.4 cm long; petals 1.1–1.2 cm long; lip 0.8–1.1 cm long Epidendrum littorale

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34. Sepals 2.5–8 cm long; petals 2.5–7 cm long; lip 2–5.7 cm long Epidendrum buenaventurae 35. Leaf blades 1–1.8 cm wide, red-purple beneath 35. Leaf blades 2–4.6 cm wide, light-green beneath 36. Stems to 70 cm long; ovary 22–28 cm; capsule 12–15 cm long Epidendrum macroophorum 36. Stems to 40 cm long; ovary 5.5–8 cm; capsule 5–8 cm long 37. Leaf blades 2–3 cm broad; sepals 4–5.2 cm long; petals 3.5–5 cm long; lip central lobe to 2.7 cm long

Epidendrum pseudonocturnum

37. Leaf blades 2.8–4.6 cm broad; sepals 6–8 cm long; petals 5.5–6.3 cm long; lip central lobe 4–5.7 cm long

Epidendrum cocornocturnum

30. Plants pendulous or ± horizontally arranged 38. Plants pendulous; petals 4–13 mm long; capsule 1–2.5 cm long 39. Many-branched plants; leaf blades 0.4–0.7 cm wide; petals 4–5 mm long, the lip ca. 6 x 1.7 mm; capsule 1–1.2 cm

Epidendrum luckei

39. Few-branched plants; leaf blades 0.7–1.9 cm wide; petals ca. 13 mm long, the lip ca. 10 x 4 mm; capsule ca. 2.5 cm

Epidendrum sculptum

38. Plants ± horizontally arranged; petals 2.5 mm long; capsule ca. 0.5 cm long Epidendrum sp. 153. Brassavola grandiflora Lindl., Edwards’s Bot. Reg. 25: 16 (1839). Syn.:

Brassavola nodosa var. grandiflora (Lindl.) H. G. Jones Holoepiphytic herbs, erect, to ca. 20 cm tall. Rhizomes short. Stems short, terete, to 5 x 0.4 cm, unifoliate at apex, surrounded by tubular scarious sheaths. Leaf blades thick, coriaceous, lorate, to 15 x 1.7 cm. Inflorescence racemose, the scape apical from stem in axil of leaf, suberect and ± arching, to 6 cm long, 2–5-flowered. Lateral sepals and petals lorate, the sepals to 9 x 0.5 cm, the petals to 8 x 0.3 cm, both yellowish-green to greenish. Lip cordiform, to 5 x 4.5 cm, acuminate at apex, at base with an elongate tubular claw that is folded around the column, to 8 cm long. Column terete, to 7 mm long. Capsules to ca. 6 cm long.

COLOMBIA. CH: (E). V: (E). C: (E). ECUADOR. G: Seemann s.n. (see discussion). Distribution: Central America to Ecuador, and introduced in the West Indies. Ecology: In wet forests and rainforests. Biological interactions: Dressler (1981) states that the Brassavola species are usually pollinated by moths. Use: This beautiful species of showy flowers is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the large size of the flowers.

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Discussions: Brassavola grandiflora is commonly considered to be a synonym of the widely cultivated B. nodosa (L.) Lindl. However, in the treatment of Orchidaceae for Flora of Ecuador (Dodson and Luer, 2005) B. grandiflora has been recognized as a valid taxon that differs from B. nodosa by its thick flattened leaves and somewhat larger flowers. The locality of an apparently Ecuadorian collection of B. grandiflora (Seemann s.n.) has been cited in “the mangroves around Guayaquil”, “but no further collections of the species in that habitat have been made even though considerable searching has been done”. It is most likely that an incorrect locality was given to the collection gathered along the coasts of Ecuador during the Voyage of H.M.S. Sulphur, which also traveled along the Pacific coasts of Colombia, where B. grandiflora it is known to occur in mangroves (Dodson and Luer, 2005). 154. Camaridium atrovinaceum (Christenson) M. A. Blanco, Selbyana 31: 52–53 (2013). Plate 30: A. Bas.: Maxillaria atrovinacea Christenson, Richardiana 9(3): 143 (2009). Syn.: Maxillaria vinacea Christenson, Richardiana 9(3): 147 (2009). Holoepiphytic herbs, erect, caespitose, to 35 cm tall. Rhizomes short. Pseudobulbs ovoidellipsoid, compressed, faintly grooved, 3–4 x 1.2–1.5 cm, unifoliate at apex, basally subtended by several old distichous sheaths, ca. 2–3 cm long. Leaf blades coriaceous, lorate, 15–30 x 1–2 cm, obliquely bilobed at apex. Inflorescence a solitary flower, the scape basal-lateral, erect, ca. 3.5 cm long; floral bract narrowly-ovate, ca.12 x 8 mm. Flower red, opaque. Lateral sepals lorate, ca. 14 x 4 mm. Petals lanceolate, ca. 10–12 x 4 mm. Lip 3-lobed, ca. 10 x 4 mm, notched at apex, the lateral lobes suberect. Column arching, ca. 5 mm long. Capsules unknown. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8199 (GUAY, NY). Distribution: Western Colombia to northwestern Ecuador. Phenology: This species has been observed flowering in March and in October. Use: It is occasionally cultivated as an ornamental. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the dark wine color of the flowers. Note: Camaridium atrovinaceum was previously known as endemic to western Colombia. The cited collection represents the first record for Ecuador (Blanco, pers. comm.). 155. Campylocentrum ecuadorense Schltr., Repert. Spec. Nov. Regni Veg. Beih. 8: 171 (1921). Holoepiphytic climber herbs. Roots elongate, produced along full length of stem. Stem cane-like, ascendent. Leaves distichous, arranged along stem; blades coriaceous, oblong to narrowly-elliptic, 2.5–4.5 x 0.6–1.3 cm, asymmetric at apex. Inflorescences raceme, to 3

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cm long, produced opposite or subopposite to leaves; floral bracts tiny. Flowers white, the segments 2.5–3 mm long, the spur produced from lip base, curled forward, 2–3 mm long. Capsule 5–7 mm long. ECUADOR. G: Cornejo 28 (GUAY). Distribution: Western Ecuador. Ecology: In dry forests influenced by fog and moist winds. Campylocentrum ecuadorense in mangroves is rather infrequent. It has been found hanging from Tillandsia usneoides (Bromeliaceae), another epiphyte on Rhizophora. IUCN Red list category: It has been categorized as Least Concern (LC) (León-Yánez et al. eds., 2011). However, due to factors such as a small range of distribution and an increasing loss of habitat, it is herein proposed as Near Threatened (NT). Etymology: The epithet refers to the geographic distribution of the species. Note: The cited collection was gathered close to the locality where the type of C. ecuadorense (Eggers 14358, US) was collected more than a hundred years ago. 156. Catasetum macroglossum Rchb. f., Gard. Chron., n.s. 2: 552 (1877). Plate 30: B. Syn.:

Catasetum sodiroi Schltr.

Vernacular name: suelda con suelda. Holoepiphytic herbs, (sub)erect. Rhizomes short. Pseudobulbs subcylindric, large, to 25 x 4 cm, thick, covered by many imbricating sheaths, with short spines from sheaths near apex. Leaves distichous, arranged at distal portion of pseudobulb; blades thin, to 30 x 8 cm, with obvious thickened, longitudinal veins, deciduous. Inflorescence an erect raceme, to 20 cm long, produced from base of pseudobulb; flowers 3–10, green or yellowish-green. Sepals and petals to 5 x 2.5 cm. Lip hood-shaped in both male and female flowers, the male flowers with an obvious, fleshy, transverse callus inside hood and with elongate antennae from sides of column. Capsule 6–9 cm long. ECUADOR. G: Cornejo 19 (GUAY). Distribution: Western Ecuador (and northwesternmost Peru?). Ecology: In dry forests, moist forests, and wet forests. Biological interactions: The flowers have been reported to be pollinated by Euglossine bees, such as Eulaema bomboides, E. polychroma, and E. cingulata (Apidae; Dodson and Gentry, 1978). Uses: This species is best known in its cultivation as an ornamental. The vernacular name refers to the medicinal use of the pseudobulb in rural areas of coastal Ecuador, where it is applied as a poultice to heal

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broken bones. It is said that the pseudobulbs of Catasetum macroglossum heal faster and better than the traditional methods given to patients in hospitals. The phytochemical compounds from the pseudobulbs involved in that process have not yet been documented. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the large lip. Note: The same vernacular name is also applied to Psittacanthus divaricatus (Viscaceae), an unrelated parasitic shrub that also occurs in mangroves and adjacent dry forests of Ecuador and Peru, which has the same uses and medicinal purposes as C. macroglossum. 157. Caularthron bilamellatum (Rchb.f.) R.E. Schult., Bot. Mus. Leafl. 18: 92, pls. 14, 15 (1958). Plate 30: C. Bas.: Syn.:

Epidendrum bilamellatum Rchb. f. Epidendrum bigibberosum Rchb. f. Holoepiphytic herbs, (sub)erect. Rhizomes short. Pseudobulbs subcylindric, large, 8–20 x 2.5–4 cm, covered by many imbricating sheaths. Leaves 2–5, distichous, arranged at distal portion of pseudobulb; blades stiffly-coriaceous, oblong, (4–)8–15 x 1–2 cm. Inflorescence an erect panicle, 18–50 cm long, produced from apex of pseudobulb; flowers 5–20, white. Sepals and petals 1.3 x 7–8 mm. Lip free, 3-lobed, the callus with two projections, more or less deltoid and diverging each other. Column ca. 8 mm long, winged. Capsule ca. 3 cm long.

COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 2609 (GUAY). Distribution: Mexico to Ecuador and Venezuela, and in the West Indies. Ecology: In moist forests, wet forests and rainforests. Use: This species is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the two divergent projections located in the callus. 158. Dichaea panamensis Lindl., Gen. Sp. Orchid Pl. 209 (1833). Plate 30: D. Syn.:

Epithecia panamensis (Lindl.) Schltr. Dichaeopsis panamensis (Lindl.) Schltr. Dichaea brachypoda Rchb.f. Holoepiphytic herbs, caespitose, subpendulous to suberect. Rhizome short. Stems cane-like, thin, horizontal to suberect, to 20 cm long, surrounded by imbricating, foliaceous sheaths. Leaves distichous, imbricate, deciduous at junction with sheath; blades foliaceous, oblong, 10–30 x 2–3 mm, acute to barely obtuse at apex, glaucous or green. Flowers solitary, axillary,

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ca. 8–10 mm long; pedicels ca. 1–1.3 cm long, thin. Ovary 1.5–2 mm long, smooth. Sepals lanceolate, to 6.5 x 2.5 mm, pale greenish-white. Petals ovate-lanceolate, to 7 x 3 mm, pale greenish-white. Lip tri-lobed, ancoriform, ca. 7 mm long, white. Capsule ca. 10–13 mm long, smooth. COLOMBIA. CH: Fuchs et al. 23303 (MO). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8203 (GUAY, NY). Distribution: Southern Mexico to Brazil. Ecology: In wet forests and rainforests. Use: This species is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the country of Panama, where it was intially known. Note: The color of flowers varies from purple to pale greenish-white (Pupulin, 2007). In our area, only individuals with greenish flowers have been seen. Plants with glaucous as well as with green leaves may be found in a same locality. As previously noted by Pupulin (op. cit.), the glaucous leaf color is a variable feature. In the REMACAM, flowers have been observed during the wet (March) and dry seasons (October). 159. Dichaea trulla Rchb. f., Beitr. Orch. Centrl.-Amer. 104 (1866). Plate 31: A. Syn.: Dichaeopsis trulla (Rchb. f.) Schltr. Dichaea powelli Schltr.

Dichaea brenesii Schltr. Holoepiphytic herbs, subpendulous to suberect. Rhizome short. Stems cane-like, ± horizontally arranged, ca. 25–50 cm long, surrounded by imbricating, foliaceous sheaths. Leaves distichous, imbricate, deciduous at junction with sheath; blades foliaceous, lorate, 6–10 cm x ca. 6–9 mm, green. Flowers solitary, axillary, ca. 15 mm long; pedicels ca. 2.2 cm long, thin. Ovary ca. 2 mm long, smooth. Sepals elliptic-lanceolate, ca. 7–10 mm long, greenish. Petals elliptic-lanceolate, ca. 7–10 mm long, greenish. Lip tri-lobed, broadly trulliform, ca. 8–9 mm long, white. Capsule ca. 20 mm long, smooth.

COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8146 (GUAY). Distribution: Honduras to Bolivia. Ecology: In wet forests. IUCN Red list category: Least Concern (LC). Etymology: Trulla, from the Latin word for scoop or spoon refers to the lip shape.

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160. Dimerandra rimbachii (Schltr.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 17: 44 (1922). Plate 31: B. Bas.:

Epidendrum rimbachii Schltr.

Epidendrum stenopetalum var. tenuicaule Rchb.f.

Dimerandra tenuicaulis (Rchb.f.) Siegerist Holoepiphytic herbs, caespitose, often forming clumps. Rhizome short. Stems cane-like, erect to suberect, to 40 cm long. Leaves distichous, spaced 3–4 cm apart; blades lorate, 5–13 cm x 5–7 mm. Inflorescence apical on stem, very short, to 1 cm long, the new inflorescences formed on old canes each year; flowers 3–5, produced in succession. Sepals and petals to 1.5 x 1 cm, pink. Lip free, 1.5–2 x 0.8–1.5 cm, pink with a white spot at juncture with column. Callus of 3 imbricate lamellae under column with an additional callose ridge on each side. Column to 3 mm long. Capsule ca. 3 cm long.

ECUADOR. E: Cornejo 70 (GUAY [2]). Distribution: Western Ecuador (and southwestern Colombia?). Ecology: In moist and wet forests. Use: This species is cultivated as an ornamental. IUCN Red list category: This persistent species is often locally common in secondary and disturbed habitats in western Ecuador. It is deemed the categorization of the native populations as Near Threatened (NT; León-Yánez et al. eds., 2011). Etymology: The epithet honors August Rimbach (1862–1943), a German collector who gathered the type, which was growing on cocoa trees near Ventanas, Province of Los Rios, Ecuador (Jørgensen and León, 1999). 161. Elleanthus discolor (Rchb.f. & Warsz.) Rchb.f., Ann. Bot. Syst. 6: 480 (1863). Bas.:

Evelyna discolor Rchb.f. & Warsz. Terrestrial herbs, caespitose. Rhizome short. Stems cane-like, terete, over 50 cm long, completely enclosed by striate leaf sheaths. Leaves distichous; blades chartaceous, elliptic to elliptic-lanceolate, to 17 x 7 cm, acuminate at apex, adaxially green, abaxially purplish. Inflorescence terminal, a sessile, polystichous spike; densely many-flowered, the flowers bright orange. Floral bracts ovate-lanceolate, up tp 1.3 cm long. Ovary up to 5 mm long. Dorsal sepal ovate to ovate-elliptic, up to 6.5 x 3 mm; lateral sepals obliquely cymbiform, carinate, to 7.2 x 3 mm. Petals obliquely ligulate to ligulate-subpandurate, up to 6.5 x 1.5 mm. Lip saccate at base, up to 7.5 x 7 mm, bicallose. Column up to 4 mm long. Capsule not seen.

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COLOMBIA. N: (E). ECUADOR. E: Játiva & Epling 2000 (S, UC). Distribution: Nicaragua to Peru. Ecology: This species occurs in the wet forest and rainforest life zones. Use: It is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the two-colored leaf blades. 162. Epidendrum bracteolatum C. Presl, Reliq. Haenk. 1(2): 100 (1827). Syn.:

Epidendrum collare Lindl. Epidendrum leucopyramis F. Lehm & Kraenzl. Holoepiphytic herbs, caespitose. Rhizome short. Pseudobulbs large, 7–20 cm long, thick. Leaves 2–4, distichous, arranged at apex of pseudobulb; blades thickly-coriaceous, oblong, 8–20 x 2–4 cm. Inflorescence a terminal erect panicle, to 70 cm long; flowers 7–50, white. Sepals and petals 2 x 3–5 mm. Lip united to apex of column, orbicular, 1 x 1 cm. Column 9–10 mm long. Capsule ca. 3 cm long.

ECUADOR. G: (S). M: (E). Distribution: Western Ecuador. Ecology: In dry forests. Use: This species is cultivated as an ornamental. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The non-distinctive epithet refers to the bracteate inflorescence. Note: The type was collected between 1790–1791 by the Czech botanist Thaddäus Haenke (1761–1816) in the environs of Guayaquil, during Alejandro Malaspina’s expedition around the world (Presl, 1825– 1835; Jørgensen and León, 1999). 163. Epidendrum buenaventurae F. Lehm. & Kraenzl., Bot. Jahrb. Syst. 26(5): 470 (1899). Plate 31: C. Holoepiphytic herbs, caespitose. Rhizome short. Stems cane-like, laterally compressed, 10–25 cm long. Leaves 3–7, distichous, distributed throughout the apical half of the stem; blades coriaceous, elliptic-lanceolate, 7–15 x 1–1.8 cm, bilobed at apex, dorsally carinate, adaxially green, abaxially red-purple. Inflorescence terminal, shortly-pedunculate, racemose to pluriracemose; producing one flower at a time and during several years, these resupinate, greenish-brown to yellowish green. Ovary ca. 50 mm long. Sepals narrowly elliptic-lanceolate, 25–46 x 1.8–3 mm. Petals linear-lanceolate, 27–45 x 1–1.5 mm. Lip united to column, 3-lobed, 20–40 x 1–1.5 mm, bicallose, the lateral lobes oblique, ovate-triangular; midlobe linear-lanceolate, 23–45 x 0.8–1.7 mm. Column straight or arcuate, 9–17 mm long. Capsule ca. 5 cm long.

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COLOMBIA. V: Gentry et al. 40428 (COL, JAUM, MO, SEL). Distribution: Southern Mexico to Ecuador. Ecology: This species is frequently found at sea level. It occurs in the wet forest and rainforest life zones. Use: It is occasionally cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the port of Buenaventura, in the Department of Valle del Cauca, Colombia, where it was initially known. 164. Epidendrum cocornocturnum Hágsater & Dodson, Icon. Orchid. (Mex.) 3: pl. 326 (1999). Plate 31: D. Holoepiphytic herbs, caespitose, often forming clumps. Rhizome short. Stems cane-like, ca. 30–40 cm long. Leaves 4–6, distichous; blades stiffly-coriaceous, 8–18 x 2.8–4.6 cm. Inflorescence a short raceme, 1–2 cm long, apical on stem; flowers several, produced in succession. Ovary 55–80 mm long. Sepals narrowly-lanceolate, 60–80 x 5–8 mm. Petals linear-lanceolate, 55–63 x 1.7–2.5 mm, yellowish. Lip united to apex of column, 3-lobed, the lateral lobes slightly curved downward, the central lobe linear-lanceolate, slender, 40–57 mm long, white, the callus yellow. Column ca. 19–22 mm long. Capsule ca. 7–8 cm long. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Bonifaz 8261 (GUAY). Distribution: Central and western Colombia to northwestern Ecuador (Hágsater, 1999). Ecology: In wet forests and rainforests. Use: This species is cultivated as an ornamental. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The compound epithet refers to the Municipality of Cocorná, where the type was collected, and “nocturnum” for the group to which the species belong (Hágsater, 1999). 165. Epidendrum littorale Hágsater & Dodson, Icon. Orchid. 2, pl. 153 (1993). Holoepiphytic herbs, caespitose. Rhizome short. Stems cane-like, terete, somewhat flexuous, ca. 15–25 cm long. Leaves 7–9, distichous, arranged along stem; blades coriaceous, elliptic, 3–6.5 x 1–2 cm, bilobed at apex. Inflorescence terminal, shortly-pedunculate, subumbellate; flowers 4–6, resupinate, green. Sepals ovate-elliptic, 12–14 x 5–7 mm. Petals narrowlyoblanceolate, 11–12 x 1.5–2 mm. Lip united to apex of column, 3-lobed, 8–11 x 13–16 mm, bicallose, the lateral lobes semiorbicular. Column arcuate, 6–7 mm long. Capsule not seen. COLOMBIA. V: Killip & Cuatrecasas 38822 (COL, VALLE). Distribution: Western Colombia and northwestern Ecuador. Ecology: In wet forests and rainforests.

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Biological interactions: It is most likely that Epidendrum littorale is pollinated by Noctuid moths, based on a pollination report of E. difforme (Dodson and Gentry, 1978), a species complex from which E. littorale was segregated. Use: This species is occasionally cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the coastal distribution. 166. Epidendrum luckei I. Bock, Die Orchidee 35(3): 100–103, t. 15 (1984). Plate 32: A. Holoepiphytic herbs, pendulous to 1 m long; rooting at base. Stems with many loose, ± horizontal, lateral branches, to 20 cm long. Leaves distichous; blades thickly-oblong, 10–50 x 4–7 mm, emarginate at apex. Inflorescences of short, terminal racemes; flowers 2–5 yellow, surrounded at base by a floral bract, the floral bract ca. 6–7 x 3–4 mm, keeled. Sepals and petals 4–5 x 1–1.5 mm, yellow. Lip united to apex of column, involute-lanceolate, ca. 6 x 1.7 mm. Column 1.8 mm long. Capsule ca. 1–1.2 x 0.8 cm. COLOMBIA. CH: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 6940 (GUAY, QCNE). Distribution: Mexico to Bolivia. Ecology: In wet and pluvial forests, often in swamps. Use: This species is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). 167. Epidendrum macroophorum Hágsater & Dodson, Icon. Orchid. (Mex.) 3: pl. 350 (1999). Plate 32: B. Holoepiphytic herbs, caespitose, often forming clumps. Rhizome short. Stems cane-like, to 70 cm long. Leaves 5–8, distichous; blades stiffly-coriaceous, to 20 x 4 cm. Inflorescence a short raceme, to 3 cm long, apical on stem; flowers few, produced in succession. Ovary 22–28 cm long. Sepals narrowly-lanceolate, 40–50 x 5 mm. Petals linear, 45–70 x 2–3 mm, yellowish. Lip united to apex of column, 3-lobed, the lateral lobes somewhat curved downward, the central lobe linear-lanceolate, slender, to 40 mm long, white, the callus yellow. Column ca. 25 mm long. Capsule 12–15 cm long. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Bonifaz 6934 (GUAY). Distribution: Panama to Ecuador. Ecology: In wet forests and rainforests. Use: This species is cultivated as an ornamental. IUCN Red list category: Although this species is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the extraordinarily long ovary that characterizes it (Hágsater, pers. comm.).

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168. Epidendrum microphyllum Lindl., J. Bot. 3: 85 (1840). Syn.:

Epidendrum sarcophyllum Focke Lanium microphyllum (Lind.) Benth. Lanium colombianum Schltr. Lanium ecudorense Schltr. Lanium peruvianum Schltr. Holoepiphytic scrambling trailing herbs. Stems slender, erect, the basal 1/3 with several small, close sheaths, distichous imbricating, leaf-bearing above. Leaves distichous; blades coriaceous, semiterete, oblong-lanceolate, 1–2.7 x 0.2–0.3 cm, acute at apex. Inflorescence terminal, loosely racemose, to 5 cm long, 5–7 flowered, the peduncle, rachis and pedicellate ovaries hyaline-hirsute. The lateral sepals strongly oblique, ovate-triangular, ca. 5.7 x 2.1 mm, greenish-yellow, abaxially hirsute. Petals linear, ca. 5 x 0.6 mm, semitranslucent greenishyellow. Lip united to column, unlobed, oblong-elliptic, ca. 4.6 x 1.5 mm. Column stout, 3.5 mm long. Capsule not seen.

COLOMBIA. N: Lehman 62209 (W). Distribution: Panama to Bolivia. Ecology: In wet forests and rainforests, often growing in along rivers and in swampy areas. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the small leaf blades. 169. Epidendrum pseudonocturnum Hágsater & Dodson, Icon. Orchid. 2: pl. 178 (1993). Plate 32: C. Holoepiphytic herbs, caespitose. Rhizome short. Stems cane-like, to ca. 40 cm long. Leaves 5–8(–9), distichous, arranged along stem or in upper part of stem; blades stiffly-coriaceous, elliptic to oblong, 8–12 x 2–3 cm, bilobed at apex. Inflorescence a terminal short raceme, to 2.5 cm long; flowers few, produced in succession. Ovary 60–80 mm long. Sepals linearlanceolate, 4–5.2 cm x 2–8 mm, greenish to yellowish. Petals linear to linear-lanceolate, 35– 50 x 2–4 mm, greenish to yellowish. Lip united to apex of column, 3-lobed, 20–40 x ca. 16 mm, bicallose, the lateral lobes slightly curved downward, the central lobe linear-lanceolate, to 27 mm long, straight, white, the callus yellowish. Column 17–25 mm long. Capsule 5–8 cm long. COLOMBIA. V: Killip 34957 (AMES, COL, S, US). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8277 (GUAY). Distribution: Western Colombia and northwestern Ecuador. Ecology: In wet forests and rainforests.

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Use: This species is cultivated as an ornamental. IUCN Red list category: The Ecuadorian populations are threatened by a loss of habitat. The status of the remaining populations in the terra firme forests of Colombia are still unknown. Data Deficient (DD). Etymology: The epithet refers to the resemblance of this plant to that of Epidendrum nocturnum Jacq. 170. Epidendrum rigidum Jacq., Enum. Syst. Pl. 29 (1760). Plate 32: D. Syn.:

Spathiger rigidus (Jacq.) Small Epidendrum cardiophorum Schltr. Epidendrum pium Rchb. f. Holoepiphytic herbs, erect. Rhizome short. Stems cane-like, to 15 cm long. Leaves 3–5, distichous; blades stiffly-coriaceous, to 8 x 2 cm, apex retuse to emarginate, sometimes asymmetric. Inflorescence a terminal raceme, erect, to 15 cm long; flowers 8–10, light-green, each subtended by a floral bract, the floral bract 1.5 x 1 cm. Sepals ca. 4 x 3 mm. Petals ca. 4 x 1 mm. Lip united to apex of column, ca. 4 x 3 mm, retuse to emarginate at apex, sometimes asymmetric. Capsule ca. 2 cm long.

COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 6962 (GUAY, QCNE). G: Cornejo 13 (GUAY). Distribution: Mexico to Bolivia. Ecology: In moist forests, wet forests, and rainforests. Use: Cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: It is most likely that the epithet refers to the rigid aspect of this species. The meaning of the epithet it is not discussed in the original publication, and the word “rigid” or an equivalent term is not mentioned in the meagre original description. 171. Epidendrum sculptum Rchb.f., Bonplandia 2: 89 (1854). Plate 33: A. Syn.:

Epidendrum sculptum var. linearifolium Rchb.f. Epidendrum florijugum Barb. Rodr. Epidendrum biflorum Cogn. Epidendrum boissierianum Schltr. Epidendrum colonense Ames Epidendrum sculptum var. arevaloi Schltr. Holoepiphytic herbs, pendulous to ca. 30 cm long; rooting at base. Stems cane-like, 7–30 cm long, loosely branched, and surrounded by imbricating, foliaceous sheaths, articulate at base of leaf, caducous towards base of stem. Leaves distichous; blades fleshy, stiff, oblong-elliptic, 2–5 x 0.7–1.9 cm, emarginate at apex. Inflorescence a terminal raceme; flowers (1–)2–3,

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surrounded at base by 2 bracts. Sepals and petals linear-oblong, 10–15 x 1.5–3 mm, greenishyellow to yellow, striate. Lip united to apex of column, 10–14 x 4–7 mm, longitudinally veined. Capsule ca. 2.5 cm long. COLOMBIA. V: Killip & Cuatrecasas 38819 (US, VALLE). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 2667 (GUAY). Distribution: Belize to Brazil (Hágsater, 2010). Ecology: In wet forests and rainforests. Use: This species is cultivated as an ornamental. IUCN Red list category: Data Deficient (DD) (Hágsater, 2010). Etymology: The epithet sculptum is a Latin word that means carved, probably referring to the lip margin that is often erose (Hágsater, 2010). 172. Epidendrum stangeanum Rchb.f., Gard. Chron., n.s. 15: 462 (1881). Plate 33: B. Syn.:

Epidendrum glandulosum Ames Holoepiphytic herbs, erect. Secondary stems pseudobulbose, fusiform, 2–4 cm x 2–5 mm, compressed; usually 1-foliolate at apex. Leaf blades fleshy, 4–10 cm x 3–5 mm, subobtuse at apex. Inflorescence a laxiflorous raceme, ca. 7 cm long, to 10-flowered, surrounded at base by one or two floral bracts; floral bract small patent; flowers yellowish-green, the sepals and ovary reddish. Sepals 4.5–5 x 1.2–1.6 mm, densely glandular and verrucose abaxially. Petals ca. 4.5 x 0.5 mm. Lip united to apex of column, ovate-cordate, ca. 3 x 3 mm, 3-nerved and carinate abaxially, acuminate at apex. Column 3 mm long, laterally dentate at apex. Capsule not seen.

COLOMBIA. N: (E). ECUADOR. E: Dodson & Embree 13108 (SEL). Distribution: Guatemala to northwestern Ecuador (Hágsater, 2010). Ecology: In wet forests and rainforests. IUCN Red list category: Data Deficient (DD) (Hágsater, 2010). Etymology: The epithet honors F. Stange (dates unknown), an orchid grower of Consul Schiller in Hamburg, Germany, in the late XIX century (Hágsater, 2010). 173. Epidendrum sp. Plate 33: C. Holoepiphytic herbs, ± horizontally arranged; rooting at base. Stem flexuose, 5–15 cm long. Leaves distichous; blades thick, stiff, oblong-elliptic, ca. 1–2.4 x 0.4–0.6 cm, emarginate at apex. Inflorescence a terminal raceme; floral bracts ovate, ca. 3–6 x 5 mm, cream. Sepals and petals linear-oblong, ca. 2.5 mm long, somewhat persistent in fruit. Capsule ca. 5 x 3.5 mm.

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ECUADOR. E: Cornejo & Cangá 8148 (GUAY). Ecology: In wet forests. IUCN Red list category: Data Deficient (DD). Note: Collections with flowers are needed in order to obtain a complete identification of this taxon. 174. Maxillaria hennisiana Schltr., Orchis 6(7): 117, t. 26, f. 10–18 (1912). Plate 33: D. Holoepiphytic herbs, erect, caespitose, forming dense clumps. Rhizomes short. Pseudobulb subquadrate, laterally somewhat compressed, 2–3 x 1.5–2.5 cm. Leaves petiolate; leaf blades 1, apical on pseudobulb, thick, coriaceous, elliptic, 10–15 x 3–5 cm, the veins inconspicuous; petioles 3–6 cm long. Inflorescences 1-several, basal-lateral, erect, the scape 4–7 cm long; flowers 1, yellow; floral bracts oblong, ca. 1.7 cm; pedicels ca. 2 cm long. Sepals linearlanceolate, ca. 2 cm long. Petals linear-lanceolate, ca. 1.8 cm long. Lip oblong, ca. 0.8 cm, obtuse at apex. Capsule ca. 4 cm long. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8202 (GUAY, NY, QCNE). Distribution: Eastern Panama to western Ecuador. Ecology: In wet forests and rainforests. Use: This species is cultivated as an ornamental. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The epithet honors Mr. H. Hennis (dates unknown), the German orchid collector of the XIX century who gathered the type. Note: It is distinguished by the long-petiolate leaves, subquadrate, dark-green pseudobulbs with dimpled surfaces (Dodson, pers. comm.). 175. Notylia replicata Rchb. f., Otia Bot. Hamburg. 1: 24 (1878). Plate 34: A. Holoepiphytic herbs, suberect. Rhizome short. Pseudobulb 1.5–2 x 1 cm, clothed by sheathing bracts at base; unifoliate at apex. Leaf blades coriaceous, 5–13 x 1.5–3 cm, narrowed to a 1–2 cm long petiole at base. Inflorescence a pendulous raceme, to 18 cm long, from base of pseudobulb; flowers 15–45. Sepals 6 x 2 mm, orange. Petals 5 x 1 mm, yellow-green. Lip triangular, white, orange at base. Capsule 2–2.5 cm long. ECUADOR. E: (S). Distribution: Western Ecuador. Ecology: In moist and wet forests. Use: Cultivated as an ornamental.

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IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Note: It has only been seen sterile in the REMACAM, Province of Esmeraldas, Ecuador. The description of the inflorescences and fruits are taken from the Flora of Rio Palenque (Dodson and Gentry, 1978). 176. Ornithidium fulgens Rchb.f., Beitr. Orchid.-K. C. Amer. 76 (1866). Plate 34: B. Syn.:

Maxillaria fulgens (Rchb.f.) L. O. Williams Holoepiphytic herbs, erect to arching-subpendulous. Rhizome elongate between pseudobulbs. Pseudobulbs ovate to elliptic, ca. 2–3 x 2–2.5 cm, hidden by basal sheaths, unifoliate at apex. Leaf blades coriaceous, lorate, 20–40 x 1–2.5 cm, asymmetric at apex. Inflorescences fasciculate, arising from sheaths of developing leaves; flowers 3–10, bright-orange. Sepals triangular-ovate, 5–8 x 4–5 mm. Petals ovate, 5–6 x ca. 3 mm. Lip pandurate, ca. 7 x 3 mm, yellow. Capsule ca. 3 cm long.

COLOMBIA. V: Gentry et al. 53420 (MO). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8118 (NY). Distribution: Nicaragua to Ecuador. Ecology: In wet forests and rainforests. Use: Cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the brightly colored flowers. 177. Ornithocephalus bryostachys Schltr., Repert. Spec. Nov. Regni Veg. 17: 17 (1921). Holoepiphytic herbs, stiffly pendulous. Rhizome short. Leaves arranged in a rigid fan shape, abscising 1.5–3 cm from base. Blades oblong, to 8 x 1 cm, conduplicate. Inflorescences axillary racemes, subpendulous, 5–7 cm long, thickly pubescent. Sepals and petals 4 x 2 mm, greenish to yellow, the back of sepals thickly pubescent. Lip white, slender, the callus green. Column with an elongate, twisted apex. Capsule 5–7 mm long, densely covered by thick erect hairs. ECUADOR. E: (S). Distribution: Widespread in the Neotropics. Ecology: In wet forests. Use: Cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet derives from the Greek prefix bryo that means moss, and the Greek word stachys that means spike, and refers to the thickly shortly-pubescent rachis of the inflorescences that resemble some mosses.

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178. Platystele cornejoi Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 120: 145, fig. 16 (2010). Plate 34: C. Holoepiphytic herbs, caespitose, forming dense small clumps, to 2 cm tall. Rhizome short. Secondary stems 4–7 mm long, unifoliate at apex. Leaf blades coriaceous, elliptic, 4–12 x 3–5 mm, broadly-obtuse at base, broadly-obtuse to rounded and apiculate at apex; the margins thickened by a marginal vein. Inflorescence an erect loose raceme, arising from base of stem, 2–5 cm long, the peduncle at base covered by stem sheaths, the rachis flexuose; flowers to ca. 15, produced in succession, light-green or yellow, 1.8–2 mm diam.; pedicels ca. 1 mm long. Sepals ovate, ca. 1 x 0.8 mm. Lateral petals lanceolate, ca. 1 x 0.3 mm. Lip upward curved at tip, ca. 0.8 mm long. Capsule 2–2.5 mm long. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8137 (GUAY, QCNE, SEL). Distribution: Northwestern Ecuador and adjacent southwestern Colombia. Ecology: So far, this small species is only known to inhabit the interior of mangrove ecosystems in the REMACAM (Province of Esmeraldas, northwestern Ecuador), and in the adjacent mangroves of Campana (Department of Nariño, Colombia), in the wet forest life zone. It may also occur nearby in the terra firme forests of Chocó. Phenology: Platystele cornejoi has been observed to form scattered populations of locally frequent and commonly flowering caespitose individuals during October, in the dry season. During the rainy season, the populations apparently decline, and individuals of P. cornejoi are rarely found with flowers. Biological interactions: It has been observed to form colonies and grow on well-developed prop roots of Rhizophora. Although there are no observations recorded as to its pollination, tiny flies could be involved in the process. Use: This species may well be cultivated as an ornamental. IUCN Red list category: Due to factors such as a narrow range of distribution and a loss of habitat, Platystele cornejoi is here categorized as Endangered (EN B1abiii). Etymology: The epithet honors Xavier Cornejo, who discovered the species and collected the type. Notes: This is the smallest vascular plant of the eastern Pacific coast mangrove swamps. In our area, the only flowers smaller than Platystele cornejoi are those of Conocarpus erectus. P. cornejoi was first collected in 2001. However, that specimen (Cornejo et al. 7232, GUAY) is a poor unicate that has only a single flower, which had remained for years in the herbarium without a determination. It was not successfully re-collected until eight years later in October, when we had the fortune to find it as a locally frequent and blooming epiphyte. 179. Polystachya concreta (Jacq.) Garay & Sweet, Orquideología 9: 206 (1974). Plate 34: D. Bas.: Syn.:

Epidendrum concretum Jacq. Onychium flavescens Blume Polystachy flavescens (Blume) J. J. Sm. Holoepiphytic herbs, erect. Rhizome short. Stems elongate, swollen into pseudobulbs at base, clothed with leaf bases. Leaves several; blades lorate, the basal to 7 x 2 cm, the apical to 20

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x 3 cm. Inflorescence paniculate, apical, to 45 cm long, branched at regular intervals, each branch to 5 cm long, becoming erect. Flowers greenish or yellowish to white. Sepals ovate, to 4 mm long. Petals oblong, to 3 mm long. Lip to 4 mm long, covered with loose, globose cells. Capsule 0.8–1.5 cm long. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8195 (GUAY, NY). Distribution: United States to Bolivia, also tropical Africa, and Asia. Ecology: In moist and wet forests. Biological interactions: There are no observations recorded, but it is most likely that the flowers are pollinated by small bees. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that refers to the stiff petals. 180. Prosthechea fragrans (Sw.) W. E. Higgins, Phytologia 82(5): 377 (1997). Bas.: Syn.:

Epidendrum fragrans Sw. Anacheilium fragrans (Sw.) Acuña Epidendrum chimboracensis Schltr. Encyclia chimboracensis (Schltr.) Dressler Prosthechea chimboracensis (Schltr.) W. E. Higgins Holoepiphytic herbs, erect, caespitose, to 40 cm tall. Rhizome to 6 cm long. Pseudobulbs narrow, ± lanceolate, 6–15 x 1–1.5 cm, unifoliate at apex and basally surrounded by a bract, ca. 5 cm long, which flakes away leaving fibers. Leaf blades coriaceous, oblong or lorate, 13–18 x 2–4 cm. Inflorescence of a terminal raceme, 7–10 cm long, 3–8-flowered. Flowers cream with incomplete, purple vertical stripes. Sepals narrowly-elliptic, 2–3 cm long. Petals 2–2.5 cm long. Lip elliptic, 1.5–2.5 cm long, acuminate at apex. Column ca. 5 mm long. Capsule 3.5 cm long, 3-winged.

COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 2671 (GUAY). Distribution: Costa Rica to Peru. Ecology: In moist and wet forests. Biological interactions: This species has been reported to be pollinated by sphecid wasps of the genus Campsomeris (Dodson and Gentry, 1978). IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word, referring to the sweet scent of the flowers. 181. Prosthechea pseudopygmaea (Finet) W. E. Higgins, Phytologia 82: 380 (1997). Bas.:

Hormidium pseudopygmaeum Finet Holoepiphytic herbs, erect, to 15 cm tall. Rhizome creeping. Stems occasionally branched. Pseudobulbs to 3 cm long, ca. 5 cm spaced on rhizome, unifoliate at apex and basally

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surrounded by a foliose bract. Leaf blades lorate, ca. 5 cm x 3 mm. Inflorescence of an erect raceme, from apex of pseudobulb, 2–3 cm long; floral bract linear, ca. 4 mm long. Sepals and petals 4–5 mm long, striate. Lip ca. 3 x 2.5 mm, white. Capsule ca. 1.5 cm long. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 2670 (GUAY). Distribution: Mexico to Ecuador. Ecology: In moist and wet forests. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the resemblance of this species to that of Prosthechea pygmaea. Note: It appears that this is the first report for Ecuador. 182. Sarcinula sp. Plate 35: A. Holoepiphytic herbs, caespitose, forming small clumps, to 4 cm tall. Rhizome short. Secondary stems 15–20 mm long, unifoliate at apex. Leaf blades coriaceous, oblong-elliptic, 15–20 x 6–9 mm, cuneate at base, broadly-obtuse to rounded, inconspicuously notched and curved-mucronulate at apex, the margins thickened by a marginal vein; greenish to purplish beneath. Inflorescence an erect loose raceme, arising from base of stem, ca. 4–5 cm long, the peduncle at base covered by stem sheath, the rachis flexuose; flowers ca. 3–6, produced in succession, light-purple, 3–4 mm diam.; pedicels 1.8–2 mm long. Sepals oblong-lanceolate, ca. 4 x 1 mm. Lateral petals shorter than sepals. Lip oblong, 2–2.5 mm long. Capsule ca. 5 mm long. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8136 (MO). Ecology: In wet forests. Growing intermixed with populations of Platystele cornejoi. Use: This species may be cultivated as an ornamental. IUCN Red list category: Data Deficient (DD). Note: This species has been collected only once, with fruits. Specimens with flowers are needed to obtain a proper identification at the species level. 183. Scaphyglottis crurigera (Lindl.) Ames & Correll, Bot. Mus. Leafl. 10(4): 85 (1942). Plate 35: B. Bas.: Syn.:

Hexadesmia crurigera Lindl. Hexopia crurigera Bateman ex Lindl. Holoepiphytic herbs, caespitose, to 40 cm tall. Rhizome short. Stems cane-like, suberect, to 15 cm long, surrounded by bracts at base, unifoliate at apex. Leaf blades narrowly-lorate, 12– 25 cm x 2–5 mm, ± symmetric at apex. Inflorescences solitary or short geniculate racemes, to 1 cm long, these on relatively long peduncles 5–15 cm long arranged from apex of stem. Sepals and petals brown to greenish. Sepals 3 x 2 mm. Petals 3 x 1 mm. Lip 3-lobed, ca. 4 mm long. Column ca. 4 mm long, somewhat curved and laterally expanded. Capsule ca. 1.5 cm long.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

COLOMBIA. N: (S). ECUADOR. E: Cornejo et al. 7235 (GUAY). Distribution: Southern Mexico to Ecuador. Ecology: In wet forests. IUCN Red list category: Least Concern (LC). 184. Scaphyglottis graminifolia (Ruiz & Pav.) Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 59 (1836). Bas.: Syn.:

Fernandezia graminifolia Ruiz & Pav. Cladobium violaceum Lindl. Scaphyglottis violacea (Lindl.) Lindl. Holoepiphytic herbs, caespitose, to 60 cm tall. Rhizome short. Stems cane-like, suberect, to 30 cm long, 2-foliate and with additional stems between leaves at apex. Leaf blades lorate, 8–20 cm x 4–7 mm, symmetric and apiculate at apex. Inflorescences from apex of stem, to 1.5 cm long. Sepals and petals ca. 4 x 1 mm, yellow. Lip ca. 3.5 x 2 mm. Column ca. 3 mm long. Capsule ca. 1.2 cm long.

COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 6935 (GUAY). Distribution: Central America to Peru. Ecology: In wet forests. Etymology: The epithet refers to the grass-like appearance of the leaf blades. IUCN Red list category: Least Concern (LC). Note: This species is the generic type. 185. Scaphyglottis prolifera Cogn. in Mart., Fl. Bras. 3(5): 75 (1898). Plate 35: C. Syn.:

Epidendrum proliferum Sw. Isochilus proliferum (Sw.) R. Br. ex Lindl. Ponera mapiriensis Kraenzl Scaphyglottis cuneata Schltr. Holoepiphytic herbs, to 25 cm tall, often forming dense clumps. Rhizome short. Stems cane-like, much branched, erect, to 10 cm long, with 2 leaves at apex, and additional stems produced from apex between leaves. Leaf blades lorate, 3–6 cm x 3–6 mm, inconspicuously asymmetric at apex. Inflorescences from apex of each stem, to 5 mm long. Sepals 3–4 x 1–2 mm, white. Petals 2–3 x 1 mm, white. Lip 3–4 x 2–3 mm, white. Column ca. 3 mm long, purple. Capsule 5–8 mm long.

COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 2610 (GUAY). Distribution: Southern Mexico to Bolivia, and in the West Indies. Ecology: In moist forests, wet forests, and rainforests.

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IUCN Red list category: Least Concern (LC). Etymology: The epithet is derived from the Latin word proles referring to the abundant off-shoots. 186. Scaphyglottis sp. nov. Plate 35: D. Holoepiphytic herbs, caespitose, to 20 cm tall. Rhizome short. Stems suberect, stipitate narrow pseudobulbs, 1–2 cm long, basally surrounded by membranous bracts which flakes away leaving fibers, unifoliate and with additional stems at apex. Leaf blades linear, 3–7 cm x 1–3 mm, somewhat conduplicate, divergent and curved, symmetric at apex. Inflorescences to 1 cm long, from apex of stems. Sepals and petals 3–5 mm long, white. Column ca. 3 mm long. Capsule ca. 8 mm long. ECUADOR. E: Cornejo & Bonifaz 7234 (GUAY). Distribution: Northwestern Ecuador (and southwestern Colombia?). Ecology: In wet forests. IUCN Red list category: At the present, according to IUCN, this species cannot be categorized until a formal publication is provided. Note: The only collection, gathered in Aug 2001 in the REMACAM (northwestern Ecuador) and housed in the herbarium GUAY, has been named “Scaphyglottis cornejoi Dodson” by C. H. Dodson himself. It is an unpublished name waiting for formal recognition. Subsequent to the previously mentioned collection date, this species has not been found bearing flowers during several trips to the REMACAM. 187. Sobralia madisonii Dodson, Icon. Pl. Trop. 4: pl. 311 (1980). Plate 36: A. Holoepiphytic herbs, suberect, caespitose, often forming clumps. Rhizome short. Stems canelike, to 1.2 m long (or more?). Leaves distichous; blades chartaceous, shortly-lanceolate, 9–20 x 7–8 cm, acute to shortly acuminate at apex, with conspicuous longitudinal veins beneath. Flowers not seen, produced in sucession from apex of stem. Capsule linear-oblong, 10–12 x 2 cm. ECUADOR. E: Cornejo & Bonifaz 6954 (GUAY). Distribution: Southwestern Colombia to northwestern Ecuador. Ecology: In wet forests and rainforests. Phenology: This species has been collected with fruits in February. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The epithet honors Michael T. Madison (dates unknown), a North American botanist and taxonomist of Araceae who collected the type. Note: Collections with flowers are needed for a definitive confirmation of its identity.

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188. Sobralia rhizophorae Cornejo & C. H. Dodson, Harvard Pap. Bot. 16(1): 53 f. 1, 2 (2011). Plate 36: B. Holoepiphytic herbs, suberect to subpendulous, caespitose, often forming clumps. Rhizome short. Stems cane-like, 1–2 m long. Leaves distichous; blades chartaceous, shortly-lanceolate, 15–25 x 8–10 cm, acuminate at apex, with 9 longitudinal major veins. Inflorescence contracted, terminal. Flowers solitary, produced in sucession from apex of stem. Sepals oblong, ca. 4.4 x 1.1 cm long, yellowish. Petals obovate-oblanceolate, oblique, 3.5–4 x 1.4–1.6 cm, white. Lip hemitubular in natural position, when expanded obovate in outline, ca. 4 x 2.5 cm, festoneate, bearing a yellowish-brown spot within, white with 2 lateral red-purple spots at the distal third, and bearing a longitudinally sulcate midvein without. Column ca. 3 cm long, straight. Capsule not seen. COLOMBIA. N: (S). ECUADOR. E: Cornejo & Cangá 8278 (GUAY). Distribution: Northwestern Ecuador and adjacent southwestern Colombia. Ecology: In wet forests. Phenology: It has been collected with flowers in November. Use: Owing to the beautiful flowers it may be cultivated as an ornamental. IUCN Red list category: Due to factors such as an increasing loss of mangrove habitat and a narrow geographical distribution range, it is herein categorized as Near Threatened (NT). Etymology: The epithet refers to the red mangrove Rhizophora L., upon which the species often growths as an epiphyte. Note: This species was discovered during the field work for this Flora. It is only known by the type collection gathered by the author in the REMACAM. 189. Soterosanthus shepheardii (Rolfe) Jenny, Die Orchidee 37(2): 74 (1986). Plate 36: C. Bas.:

Sievekingia shepheardii Rolfe Holoepiphytic herbs, erect, caespitose. Rhizomes short. Pseudobulb ovoid to narrowly-ovoid, 2–4 x 1.5–2.5 cm, basally subtended by several old distichous sheaths, ca. 2–3 cm long. Leaves petiolate; leaf blades 1–2, apical on pseudobulb, foliaceous, elliptic to oblong-elliptic, 12–20 x 4–7 cm, purple beneath, with obvious thickened, longitudinal veins; petioles 1.5–3 cm long. Inflorescences 1–2, basal-lateral, erect racemes, the scape ca. 10 cm long; flowers ca. 8–15, yellow; floral bracts narrowly-lanceolate, 1–1.5 cm long; pedicels 1–1.8 cm long. Sepals lanceolate, ca. 1.5–2 cm long. Petals obovate-oblong, ca. 1 cm long. Lip subpandurate-oblong, ca. 1 cm long, subacute at apex, the callus papillose. Column ca. 0.8 cm long. Fruit capsule.

COLOMBIA. V: Gentry & Juncosa 40725 (MO). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8120 (GUAY). Distribution: Western Colombia and northwestern Ecuador.

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Ecology: In wet forests and rainforests. Phenology: It has been collected with flowers in late February, during the rainy season. Use: This beautiful species is cultivated as an ornamental. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The epithet honors S. Shepheard (dates unknown), an avid collector of orchids who in 1912 gathered sterile specimens of this species in Río Condoto, Department of Choco, Colombia. Subsequently, it flowered in cultivation in Kew, London, from where it was described as new to science. Note: The plant is similar to a Lycaste, but it is recognized by the erect inflorescences and the purple undersides of the leaf blades (Dressler, pers. comm.). 190. Trichosalpinx orbicularis (Lindl.) Luer, Phytologia 54(5): 396 (1983). Plate 36: D. Holoepiphytic herbs, erect, caespitose. Rhizome short. Stems cane-like, 3–8 cm long, surrounded by spaced spathaceous bracts, these dorsally keeled, ciliate at margin; unifoliate at apex. Leaf blades fleshy, ovate to elliptic, 2.5–4.5 x 1.7–3 cm, obtuse at apex, slightly curved (from lateral view), purplish beneath. Inflorescences of 3 apical racemes, these basally surrounded by a common bract, each with 1–3 flowers, red-purple at anthesis. Sepals ca. 7 mm long, the dorsal sepal lanceolate, the two lateral linear-lanceolate. Lip linear-lanceolate, ca. 4 mm long, ciliate at apex. Column ca. 2 mm long. Capsule ca. 6 mm long. COLOMBIA. N: (S). ECUADOR. E. Cornejo & Cangá 8218 (GUAY, NY, SEL). Distribution: Widespread in Tropical America. Ecology: In wet forests and rainforests. Phenology: In the REMACAM it has been observed with flowers during the dry and rainy seasons. Use: It is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the shape of the leaf blades, originally described as “subrotundo-ovato” (=suborbicular-ovate), a variable feature. 191. Trigonidium seemannii Rchb. f. in Seem., Bot. Voy. Herald 214 (1854). Syn.:

Trigonidium riopalenquense Dodson Holoepiphytic herbs, erect, caespitose, to 45 cm tall, often forming large clumps. Rhizome short. Pseudobulb ovoid to conic-ovoid, 2.5–3.5 x 2–2.5 cm, laterally compressed. Leaf blades 1–2, apical on pseudobulb, coriaceous, lorate, 25–40 x ca. 1 cm. Inflorescences solitary, several, basal-lateral, erect, the scape to 20 cm long. Floral bract ca. 3.5 cm long. Flowers yellowish to pinkish. Sepals spathulate, to 3.1 x 1.5 cm, held erect in a tube, flared near apex. Petals elliptic-lanceolate, ca. 1.5 x 0.5–0.8 cm, grey with a blue-grey spot at apex. Lip 3-lobed, to 8 x 6 mm, the midlobe acute at apex. Column ca. 5 mm long. Fruit capsule.

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COLOMBIA. N: (E). ECUADOR. E: (S). Distribution: Nicaragua to western Ecuador. Ecology: In wet forests and rainforests. Use: It is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet honors Berthold Carl Seemann (1825–1872), a British naturalist of the H.M.S. Herald expedition, who collected the type. PIPERACEAE (Pepper Family) Epiphytic or terrestrial herbs, vines, shrubs or sometimes trees. Leaves usually alternate, opposite or verticillate; petioles often with sheating foliar bases; blades simple, entire, palmately or pinnately veined or plinerved, sometimes peltate. Inflorescence axillary, terminal, opposite, solitary spike or racemose or compound, umbellate or paniculate when compound, usually white or cream. Flowers numerous, usually close together in spiral or ring-like arrangement, without a perianth, bisexual. Stamens usually 1–6, free and/or arising near base of ovary. Ovary superior. Fruit a drupe, dry or fleshy; seeds small, solitary. Key to the species 1. Plants usually prostrate, with elongate creeping stems; leaf blades 0.2–15 cm long; inflorescences spicate 2. Leaf blades 0.2–1.5 cm long 3. Leaf blades deltoid to orbicular-subdeltoid, 1–2.5 cm wide; petioles to 3 cm long Peperomia serpens 3. Leaf blades orbicular to elliptic or elliptic-obovate, narrowly ovate or shortly-lanceolate, 0.2–1 cm wide (to 1.5 cm in P. gorgonillana only); petioles to 1 cm long 4. Plants of a delicate aspect; leaf blades 0.2–0.5 cm long

Peperomia emarginella

4. Plants of a no delicate aspect; leaf blades 0.6–2 cm long 5. Leaf blades usually narrowly-ovate or shortly lanceolate, sometimes to elliptic Peperomia jamesoniana 5. Leaf blades orbicular, suborbicular or elliptic-obovate to elliptic 6. Leaf blades pubescent above, ciliate at margins 7. Leaf blades elliptic to elliptic-obovate

Peperomia gorgonillana

7. Leaf blades orbicular to elliptic-orbicular

Peperomia rotundifolia

6. Leaf blades all glabrous

Peperomia obovalis

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2. Leaf blades 1.5–16 cm long Peperomia jamesoniana

8. Leaf blades 0.3–0.8 cm wide 8. Leaf blades 1–7 cm wide 9. Spikes to 5 cm long

10. Leaf blades narrowly-obovate or narrowly-oblanceolate to elliptic; spikes 2–3 mm diam Peperomia obtusifolia 10. Leaf blades deltoid or ovate to suborbicular; spikes 1–2 mm diam. 11. Leaf blades deltoid to orbicular-subdeltoid Peperomia serpens 11. Leaf blades ovate or ovate-elliptic to suborbicular Peperomia blephariphylla

12. Beak of fruit ending in a knoblike tip 12. Beak of fruit pointed or blunt, but not knoblike

Peperomia ciliaris

9. Spikes 5–20 cm long 13. Leaf blades ovate to lanceolate Peperomia elongata 13. Leaf blades narrowly-obovate or narrowly-oblanceolate to elliptic 14. Leaf blades somewhat acute to rounded or inconspicuously emarginate at apex; spikes 2–3 mm diam. 14. Leaf blades acuminate at apex; spikes 1–2 mm diam.

Peperomia obtusifolia Peperomia portobellensis

1. Plants erect, with short stems; leaf blades 15–30 cm long; inflorescence narrowly-paniculate Peperomia pernambucensis 192. Peperomia blephariphylla Trel. & Yunck., Piperac. N. South Amer. 2: 665, f. 580 (1950). Holoepiphytic appressed climber herbs. Stems prostrate, to 80 cm long or more, sparsely branched, crisp-puberulent to glabrate. Leaves alternate, several to many along stem; blades thinly chartaceous to foliaceous, ovate to suborbicular, 1.3–3(–5) x 1.2–2.5(–3.7) cm, subacute to truncate at base, rounded to obtuse at apex, obscurely subpalmately 5–7-nerved, ± hairy on both surfaces near the margins and densely ciliate; petioles 0.8–4 cm long, puberulent to glabrate. Inflorescence terminal or opposite the leaves; peduncle 1–3 cm long, puberulent or glabrate; spike to 3 cm x 1–2 mm, greenish to light-yellow. Beak of fruit ending in a knoblike tip. COLOMBIA. CH: Fuchs & Zanella 21867 (COL, MO, NY). V: (E). Distribution: Western Colombia. Ecology: In rainforests. IUCN Red list category: Although it is persistent in secondary forests, due to an increasing loss of habitat, it is herein categorized as Near Threatened (NT). Etymology: The epithet is formed by the Greek word blepharis, eyelash, and the Greek suffix phyla, leaf, and refers to the leaf blades of a densely ciliate eyelash-like pubescence.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

193. Peperomia ciliaris C. DC., J. Bot. 4: 144 (1866). Holoepiphytic climber herbs. Stems prostrate, to 80 cm long or more, sparsely branched, densely pilose. Leaves alternate, several to many along stem; blades thickly coriaceous, ovate to ovate-elliptic or suborbicular, 1.5–4 x 1–3 cm, obtuse to truncate-subpeltate at base, widely obtuse to subacute at apex, 5–7-subplinerved, sparsely crisp-pubescent above, glabrescent beneath, densely ciliate near margins; petioles 1–4 cm long, puberulent to glabrate. Inflorescence terminal and axillary; peduncle 1–5 cm long, pilose or glabrate; spike 1–2 cm x 1–2 mm, greenish to white to cream. COLOMBIA. CH: (E). V: Gentry et al. 53373 (MO). Distribution: Costa Rica to northwestern Ecuador. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word referring to the ciliate pubescence. 194. Peperomia elongata Kunth, Nov. Gen. Sp. (quarto ed.) 1: 62 (1815 [1816]). Syn.: Piper elongatum (Kunth) Poir.

Peperomia cuspidigera Sodiro

Piper extensum Roem. & Schult.

Peperomia myriocarpa Miq.

Peperomia japurensis C. DC.

Peperomia surinamensis C. DC.

Peperomia maribiana C. DC.

Peperomia controversa C. DC. Holoepiphytic appressed climber herbs. Stems prostrate, to 70 cm long or more, distally subpendulous. Leaves alternate, ca. 6–12; blades somewhat fleshy, ovate to lanceolate, 1.7–7 x 1–2.5 cm, obtuse to truncate at base, acuminate at apex, obscurely 7-plinerved, glabrous on both sides but ± marginally ciliate at apex; petioles 2–20 mm long, glabrous to pilose. Inflorescence terminal; peduncle 5–10 mm long; spike 5–10 cm x 1–1.5 mm, creamish-green.

COLOMBIA. CH: (E). V: (E). N: (E). ECUADOR. E: Cornejo & Bonifaz 6956 (GUAY). Distribution: Colombia and Venezuela to northern Argentina. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet means the elongated Peperomia.

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195. Peperomia emarginella (Sw. ex Wikstr.) C. DC. in A. DC., Prodr. 16: 437 (1869). Bas.:

Piper emarginellum Sw. ex Wikstr.

Syn.:

Acrocarpidium exile Miq.

Peperomia exilis Griseb. Peperomia arundinacea C. DC. Peperomia emarginella var. glabrior C. DC. Peperomia delicatissima Trel. Holoepiphytic appressed climber herbs of a very delicate net-like appearance. Stems prostrate, to 80 cm long or more, many branched, glabrous. Leaves alternate, many along stem; blades thinly chartaceous or foliaceous, orbicular, orbicular-obovate, obcordate or broader than long, 2–5 x 2–5 mm, rounded to subacute and commonly slightly peltate at base, widely obtuse to subtruncate, commonly emarginate at apex, obscurely palmately 3-nerved, glabrous or rarely with few long hairs beneath, the margins usually ± distantly setose-ciliate; petioles 1–3 mm long, glabrous. Inflorescence terminal; peduncle 4–10 mm long; spike to 2 cm x ca. 0.5–1 mm, loosely flowered, greenish to yellowish-green.

COLOMBIA. CH: (E). V: (E). ECUADOR. E: (S). Distribution: Belize to Bolivia, and in the West Indies. Ecology: In wet forests and rainforests. Use: This species may be cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the emarginate apex of leaf blades. 196. Peperomia gorgonillana Trel. & Yunck., Piperac. N. South Amer. 2: 632, f. 554 (1950). Holoepiphytic appressed climber herbs. Stems prostrate, to 80 cm long or more, crisp-puberulent. Leaves alternate, many along stem; blades somewhat fleshy, elliptic or elliptic-obovate, 0.6–1.5 x 0.5–1.5 cm, cuneate to rounded at base, usually obtuse to rounded at apex, obscurely palmately 3-nerved, ± pubescent on both sides, ciliate; petioles 3–10 mm long, crisp-pubescent. Inflorescence terminal; peduncle 4–15 mm long, ± crisp-pubescent; spike 0.5–4 cm x 1–1.5 mm, greenish. COLOMBIA. CH: (E). C: (E). N: (S). ECUADOR. E: (E). Distribution: Western Colombia and western Ecuador. Ecology: In wet forests and rainforests. IUCN Red list category: Data Deficient (DD). Etymology: The epithet refers to Gorgonilla Island, in the Department of Nariño, southwestern Colombia, where the type was collected by Ellsworth P. Killip in 1939.

FLORA OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS

197. Peperomia jamesoniana C. DC. in A. DC., J. Bot. 4: 137 (1866). Syn.:

Peperomia demissa C. DC. Holoepiphytic appressed climber herbs. Stems prostrate, to 80 cm long or more, many branched, glabrous or glabrescent. Leaves alternate, many along stem; blades foliaceous, narrowly-ovate or shortly-lanceolate to elliptic, 7–20 x 3–8 mm, cuneate at base, obtuse and often notched at apex, obscurely palmately 3-nerved, glabrous on both sides or occasionally villose above, dark-dotted beneath, with a few bristle-like hairs at apex; petioles 1–6(–10) mm long, glabrous. Inflorescence terminal; peduncle 3–10 mm long; spike to 5 cm x ca. 1 mm, greenish.

COLOMBIA. CH: (E). V: (E). ECUADOR. E: (S). Distribution: Costa Rica to Ecuador and Venezuela. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet honors William Jameson (1796–1873), an English collector. 198. Peperomia obovalis Trel. & Yunck., Piperac. N. South Amer. 2: 630, f. 551 (1950). Plate 37: A. Holoepiphytic appressed climber herbs. Stems prostrate, to 60 cm long or more, glabrous. Leaves alternate, many along stem; blades somewhat fleshy, elliptic-orbicular to orbicular, 0.6–1 x 0.6–0.8 cm, widely obtuse to rounded at base and apex, the lateral veins inconspicuous, glabrous; petioles 3–10 mm long, glabrous. Inflorescence terminal; peduncle 3–10 mm long; spike 2–5 cm x 1–1.5 mm, greenish. COLOMBIA. CH: (E). V: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8215 (GUAY). Distribution: Western Colombia and Ecuador. Ecology: In wet forests and rainforests. IUCN Red list category: Data Deficient (DD). Etymology: The non-distinctive epithet refers to the obovate shape of the leaf blades. 199. Peperomia obtusifolia (L.) A. Dietr., Sp. Pl. (sexta ed.) 1: 154 (1831). Plate 37: B. Bas.: Syn.:

Piper obtusifolium L. Piper humile Mill. Piper milleri Roem. & Schult. Peperomia hemionitidifolia Ham. Peperomia commutata Trel. Peperomia bayatana Trel. Peperomia daiquirana Trel. Peperomia lunata Trel.

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Peperomia earlei Trel. Peperomia dodecatheontophylla Trel. Peperomia antoni var. reducta Trel. Holoepiphytic herbs, solitary or forming groups of several individuals, erect, to 20 cm tall. Stem short. Leaves alternate, ca. 3 to 6; blades fleshy, narrowly-obovate or narrowlyoblanceolate to elliptic, 3.5–16 x 2–7 cm, attenuate at base, somewhat acute to rounded or inconspicuously emarginate at apex, the lateral veins inconspicuous; petioles 0.8–2 cm long. Inflorescence terminal and/or axillary, simple or forked; peduncle 2–7 cm long; spike 2.5–20 cm x 2–3 mm, creamish-green.

COLOMBIA. CH: (E). V: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 2724 (GUAY). Distribution: Mexico to Brazil, and in the West Indies. Ecology: In wet forests and rainforests. Use: It is cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet refers to the obtuse apex of the leaf blades. Note: This species has been misreported for the mangroves of REMACAM in northwestern Ecuador as Anthurium citrifolium (Ecociencia, 2006: 49, fig. 3). 200. Peperomia pernambucensis Miq., London J. Bot. 4: 420 (1845). Plate 37: C. Syn.:

Peperomia longifolia C. DC. Peperomia multiplex Leprieur ex C.DC. Peperomia lopezensis Trel. Peperomia paniculata Regel

Vernacular name: amargosa. Holoepiphytic herbs, solitary or few individuals, erect. Stem short. Leaves alternate, ca. 3 to 6; blades coriaceous, elliptic, elliptic-oblong or oblanceolate, 15–30 x 4–10 cm, narrowly cuneate or attenuate at base, acuminate or occasionally obtuse at apex, the lateral veins ca. 7–8 per side, inconspicuous; petioles 2–8 cm long. Inflorescence terminal, narrowly-paniculate; primary peduncle 4–8 cm long, red or reddish, densely puberulent; lateral peduncles ca. 3 mm long, the spikes many, 1–2 cm x 1–2 mm, creamish. COLOMBIA. CH: Fuchs et al. 22161 (COL). V: (E). Distribution: Nicaragua to Brazil. Ecology: In wet forests and rainforests. Use: The plant is reported to be used in malaria treatment. It may be cultivated as an ornamental.

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IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the State of Pernambuco, Brazil, where the species was initially known to occur. 201. Peperomia portobellensis Kongl. Vetensk. Acad. Handl. 147 (1854 [1856]). Plate 37: D. Holoepiphytic climber herbs. Stems prostrate, to 80 cm long or more, crisp-pubescent, glabrescent, glabrescent. Leaves alternate, many along stem; blades coriaceous, narrowlyelliptic, 5–9 x 1.1–2.5 cm, cuneate at base, acuminate at apex, ciliate at margins, the lateral veins inconspicuous; petioles 1–4 mm long, crisp-pubescent. Inflorescence terminal; peduncle ca. 15 mm long, glabrous; spike 5–15 cm x 1–2 mm, greenish. COLOMBIA. CH: (S). Distribution: Southern Mexico to Colombia. Ecology: In wet forests and rainforests. Use: This species may be cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the port city of Portovelo, Province of Colon, Panama, where the type was collected by Johan E. Billberg (1799–1845) in 1826. 202. Peperomia rotundifolia (L.) Kunth, Nov. Gen Sp. (quarto ed.) 1: 65 (1815). Bas.: Syn.:

Piper rotundifolium L. Piper nummularifolium Sw. Peperomia nummularifolia (Sw.) Kunth Acrocarpidium nummularifolium (Sw.) Miq. Peperomia rotundifolia var. subglabrilimba C. DC. Peperomia rotundifolia var. subelliptica Trel. Holoepiphytic appressed climber herbs of a net-like appearance. Stems prostrate, to 60 cm long or more, many branched, crisp-puberulent or somewhat sparsely villous, glabrescent. Leaves alternate or subopposite, many along stem; blades thinly chartaceous, orbicular to elliptic-orbicular, 5–12 x 4–10 mm, subacute or rounded and subpeltate at base, widely obtuse to subtruncate at apex, obscurely palmately 3-nerved, crisp-puberulent, loosely villous, or commonly glabrescent beneath, the margins ± ciliate; petioles 1–10 mm long. Inflorescence terminal; peduncle 3–15 mm long, crisp-puberulent or glabrate; spike to 4 cm x ca. 1 mm, greenish to greenish-yellow.

COLOMBIA. CH: Gómez et al. 431 (HUA, MO). V: (E). N: (E). ECUADOR. E: (E). Distribution: Belize to Bolivia, and in the West Indies. Ecology: In wet forests and rainforests. It has often been collected along riversides.

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Use: This species may be cultivated as an ornamental. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet refers to the orbicular shape of the leaf blades. 203. Peperomia serpens (Sw.) Loud. Hort. Brit. 13 (1830). Bas.: Syn.:

Piper serpens Sw. Verhuellia serpens (Sw.) Miq. Peperomia scandens Ruiz & Pav. Piper scandens (Ruiz & Pav.) Vahl Acrocarpidium scandens (Ruiz & Pav.) Miq. Piper bracteatum Thomps. Peperomia repens Kunth Acrocarpidium repens (Kunth) Miq. Peperomia reniformis Hook. Piper guildinguianum Spreng. Acrocarpidium guildinguianum (Spreng.) Miq. Peperomia myosurus Willd. Peperomia pulicaris Opiz Acrocarpidium pulicare (Opiz) Miq. Peperomia ionophylla Griseb. Holoepiphytic appressed climber herbs. Stems prostrate, to few meters long, branched, crisppubescent. Leaves alternate, many along stem; blades thinly chartaceous to foliaceous, deltoid to orbicular-subdeltoid, 1–3 x 1–2.5 cm, truncate to subcordate at base, acute to obtuse at apex, palmately 3–5-nerved, ± crisp-pubescent on both sides, the margins ciliolate toward apex; petioles 5–30 mm long. Inflorescence axillary and sympodially terminal; peduncle to 4 cm long, crisp-pubescent; spike to 4 cm x ca. 1 mm, greenish-yellow to cream.

COLOMBIA. CH: (E). V: (E). ECUADOR. E: (S). Distribution: Southern Mexico to Bolivia, and in the West Indies. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Latin word serpens (snake), and means the snake-like Peperomia, referring to the reptant-creeping stems. It is a common habit for many other species in this genus. POACEAE (Grass Family) Herbs or subwoody plants with terete, hollow or solid stems (culms), these closed at nodes. Leaves often distichous, usually parallel-veined, consisting of a sheath enveloping culm, a usually flat blade, and between the two, on inside, a membranous hyaline or hairy appendage (ligule). Perianth reduced to 2(–3) small hyaline appendages (lodicules). Flowers (florets) perfect, borne in spikelets consisting of a shortened axis (rachilla) and bracts, the lowest 2 (glumes) empty, the others (lemmas) subtending and

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often partially enclosing a second 2-nerved bract (palea) that bears in its axil. Stamens usually 3(–6) on slender filaments. Pistil 1, bearing a single ovule, stigmas 2(–3). Fruit normally a caryopsis. The grasses listed below are salt tolerant and light demanding species. These are found in openings in the mangroves (Lægaard pers. comm.). Key to the species Paspalum vaginatum 1. Inflorescences of 2 horizontally divergent rows 1. Inflorescences panicles or racemes 2. Inflorescences well-developed panicles, 15–40 cm long 3. Leaf blades ± scabrous above; panicles dense, the spikelets 1–2 cm long, strongly flattened Uniola pittieri 3. Leaf blades smooth above; panicles open, the spikelets ca. 2 mm long, subterete Urochloa mutica Jouvea straminea 2. Inflorescences delicate short racemes, to 5 cm long 204. Jouvea straminea E. Fourn., Bull. Soc. Bot. Belg. 15: 475 (1877). Terrestrial herbs, perennial, tough, wiry, extensively stoloniferous. Culms slender, erect from a dense knotty crown, branching, 10–35(–50) cm tall. Lowermost sheaths very short, crowded, with reduced blades; blades scabrous, 2–6 cm x 1–3 mm. Staminate inflorescences 2–3 cm long, bearing 2–3 spikelets; spikelets 4–8-flowered; glumes acuminate, unequal; lemnas acute or acuminate, ca. 4 mm long. Pistillate inflorescences 2–3 together, not crowded; spikelets 15–25 mm long, 2–4-flowered, slender, pointed, arcuate. COLOMBIA. CH: (E). ECUADOR. E: (S). Distribution: Mexico to Ecuador. Ecology: It usually occurs in salt marshes along the coast, in wet forest and rainforest life zones. It is a salt tolerant species. IUCN Red list category: Least Concern (LC). Etymology: The non-distinctive epithet refers to the appearance of a straw. 205. Paspalum vaginatum Sw., Prodr. Veg. Ind. Occ. 21 (1788). Syn.:

Paspalum littorale R. Br. Digitaria foliosa Lag. Paspalum tristachyum LeConte Digitaria tristachya Schult. Terrestrial herbs, perennial, with low creeping rhizomes and slender wiry stolons. Flowering culms 10–60 cm tall, glabrous. Leaves alternate; blades linear, 2.5–15 x 0.3–0.8 cm. Racemes

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2(3–5), rachis 3-angled, often flexuose especially at apex; spikelets solitary, in 2 rows on 1 side of a narrow winged rachis, imbricate, elliptic, 3–4 mm long, glabrous. First glume wanting or rarely slightly developed. Second glume and sterile lemma equal, thin, weakly 5-nerved, but midnerve of second glume often suppressed, sterile lemma often transverse undulate. Fertile lemma 2.5–3 mm long, slightly concave-convex, clasping palea for only about two-thirds of its length. ECUADOR. E: (S). Distribution: United States to Chile. Ecology: It usually grows along sea coasts, in salt marshes and on sandy beaches; in dry, moist, and wet forest life zones. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the sheath of leaf blades, a common feature in grasses. 206. Uniola pittieri Hack., Oesterr. Bot. Zeitschr. 52: 309 (1902). Plate 38: A. Vernacular names: pajonal, tumbagaspar. Terrestrial herbs, perennial, stout, extensively stoloniferous, glabrous. Culms in large coarse clumps 0.5–1 m tall, some internodes elongate with 2–4 very short ones between them. Leaves appearing in groups; sheats rounded, glabrous, villose in throat and on margins; blades very firm, elongate, attenuate to a fine trip, 8–10 mm wide toward base, usually curled at apex, glabrous beneath, more or less scabrous above and on margins. Panicles usually 20–30 x 4–6 cm, dense; spikelets very shortly pedicellate, very flat, 1–2 x ca. 1 cm. Lemmas ovate, acute, ciliate on keel, the margins densely villose at base, otherwise glabrous. Palea about two-thirds as long as lemma, narrow, the keel not bowed out, ciliate with fine hairs, which protrude beyond margin of lemma. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Cornejo et al. 7223 (GUAY). Distribution: Mexico to western Ecuador. Ecology: Uniola pittieri occasionally occurs in hyposaline disturbed mangroves located in wet forests. It mostly inhabits sandy beaches; in very dry, dry, moist, wet forest, and rainforest life zones. Biological interations: The flowers have been observed to be visited by Trigona bees (Apidae; Duke 11560, MO). IUCN Red list category: Least Concern (LC). Etymology: The epithet honors Henri François Pittier (1857–1950), a Swiss botanist and avid collector of plants who gathered the lectotype, as well as a multitude of new species of plants from Costa Rica, Panama, and Venezuela.

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207. Urochloa mutica (Forssk.) T. Q. Nguyen, Novosti Sist. Vyss. Rast. 1966: 13 (1966). Bas.: Syn.:

Panicum muticum Forssk. Panicum purpurascens Raddi Panicum barbinode Trin. Panicum glabrinode Hack. Brachiaria purpurascens (Raddi) Henrard Brachiaria glabrinodis (Hack.) Henrard Brachiaria mutica (Forssk.) Stapf.

Vernacular name: zeneida. Terrestrial herbs, perennial, 1–2 m tall; growing in dense masses or with single stems, often scrambling in shrubs or other plants. Stems commonly branching, hollow, 3–6 mm diam. Sheaths usually as long as or longer than internodes, glabrous or long hirsute, especially at and near base; ligule a dense rim of erect hairs, ca. 1 mm long. Blades flat, linearlanceolate, 20–40 x 1–2 cm, midrib prominent on lower side, glabrous to sparsely hirsute on one or both sides. Panicle 15–25 cm long, branches raceme-like, erect, spreading, the lower ones 8–12 cm long, all single or few together, the lower with prolonged branchlets, the branchlets and pedicels hirsute. Spikelets ovate, ca. 2 x 1 mm, acuminate at apex. COLOMBIA. N: (E). ECUADOR. E: Cornejo & Bonifaz 6989 (AAU, GUAY). Distribution: Mexico to Argentina. Ecology: Urochloa mutica is an introduced weed (Zuloaga and Morrone in Jørgensen and León, 1999). It is common in fresh water inland swamps (Lægaard pers. comm.; pers. obs.), and is sometimes frequent along riverbanks, reaching to the hyposaline mangroves, in dry, moist, wet forest, and rainforest life zones. Uses: It is grown for forage and is occasionally used as a medicine against paludism (Madrigal 832, MO). IUCN Red list category: Least Concern (LC). Etymology: The epithet derives from the Latin word muticus, meaning blunt. PONTEDERIACEAE (Water hyacinth Family) Annual or perennial aquatic herbs, floating or rooted in mud, rhizomatous or stoloniferous. Stems usually short, often somewhat spongy or succulent. Leave distichous, petiolate, the petioles vaginate, sheating at base; stipules often present. Inflorescences terminal or pseudolateral, spicate, racemose, or paniculate, each subtended by a spathe-like leaf sheath. Flowers bisexual, actinomorphic or zygomorphic with a perianth of 6 imbricate segments, these free or basally connate. Stamens 3 or 6, rarely 1, inserted on the perianth, often unequal, with free filaments. Ovary superior, 3-locular; ovules numerous or solitary; styles slender, the stigmas 3- to 5-lobed. Fruit a 3-valved capsule or indehiscent nutlet; seeds numerous usually ribbed.

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Key to the species 1. Flowers with upper lobe fimbriate at distal half, dark-blue in lower half only 1. Flowers with upper lobe entire at distal half, light-blue in the center

Eichhornia azurea Eichhornia crassipes

208. Eichhornia azurea (Sw.) Kunth, Enum. Pl. 4: 129 (1843). Plate 38: C. Bas.:

Pontederia azurea Sw.

Syn.:

Piaropus azureus (Mart.) Raf.

Pontederia aquatica Vell.

Eichhornia aquatica (Vell.) Schltdl.

Vernacular names: lechuga, lirio, taruya. Floating herbs or rooting in mud if stranded, perennial. Rhizomatous or stoloniferous; roots feathery, dense. Leaves arranged in a rosette, somewhat ascending and widely spreading; petioles terete, elongated, to 50 cm long, inflated, spongy and aerenchymatous within, glabrous without. Blades glossy-green, coriaceous, suborbicular to ovate, 7–15 x 6–11 cm, widely obtuse to truncate at base, widely obtuse at apex, the veins inconspicuous, glabrous. Inflorescence an erect raceme, central, terminal, 25–40 cm long. Flowers showy, light-blue, the upper lobe bearing a small yellow blotch and dark-blue towards the petal base. Perianth lobes 6, 20–40 cm long. Stamens 6, 3 elongated, 3 shorter. Capsule coriaceous, ca.1.5 cm long; seeds many. COLOMBIA. CH: (S). V: (E). Cauca: (E). N: (E). ECUADOR. G: (S). Distribution: Mexico to Argentina. Ecology: It is an aquatic weed that mainly inhabits fresh-water inland habitats, and it is carried downstream by river water currents. This floating plant, whether solitary or forming colonies, is a mangrove visitor rather than a permanent resident. Due to higher levels of salinity, Eichhornia azurea eventually dies in the brackish estuarine waters. It occurs in dry, moist, wet forest, and rainforest life zones. Biological interactions: The densely fasciculate, hair-like black roots provide shelter to many species of alevins, small fish, snails, and other small invertebrates. The fish feed on the distal portion of the roots. Dispersal: Whether vegetatively or by seeds, this species is water dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the blue color of the flowers.

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209. Eichhornia crassipes (Mart.) Solms in A. DC. & C. DC., Monogr. Phan. 4: 527 (1883). Plate 38: B. Bas.: Syn.:

Pontederia crassipes Mart. Piaropus crassipes (Mart.) Raf.

Vernacular names: buchón, lechuga, lechuga de bototo, lirio, marimacha. Floating herbs or rooting in mud if stranded, perennial. Rhizomatous or stoloniferous; roots feathery, dense. Leaves arranged in a rosette, somewhat ascending and widely spreading; petioles terete, elongated, to 50 cm long, inflated or bulbous, especially towards base, spongy and aerenchymatous within, glabrous without. Blades glossy-green, coriaceous, suborbicular to ovate, 7–15 x 6–11 cm, widely obtuse to truncate at base, widely obtuse at apex, the veins inconspicuous, glabrous. Inflorescence an erect raceme, central, terminal, 25–40 cm long. Flowers showy, lavander-purple or light-blue, the upper lobe bluish and bearing a yellow blotch. Perianth lobes 6, 16–37 cm long. Stamens 6, 3 elongated, 3 shorter. Capsule coriaceous, ca. 1.2–1.5 cm long; seeds many. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: (S). M: (S). G: (S). Distribution: A widely distributed invasive weed, native to northern South America and introduced in the tropics and subtropics around the world. Ecology: Same as in E. azurea. Biological interactions: Snakes have been observed to be transported on the bed-like floating colonies of E. crassipes. The densely fasciculate, hair-like black roots provide shelter to many species of alevins, small fish, snails, and other small invertebrates. The fish feed on the distal portion of the roots. Dispersal: Whether vegetatively or by seeds, this species is water dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet is derived from the Latin word crassus, meaning thick, referring to the inflated or bulbous petioles. Note: During the rainy season, the salinity levels downstream in estuaries and mangrove areas decrease, sometimes allowing large floating populations of E. crassipes to be transported out to sea, hampering the boat movement and work of fishermen (Cornejo, 2005). POTAMOGETONACEAE (Pondweed Family) Plant herbaceous, perennial or rarely annual, glabrous, of fresh or brackish waters, growing entirely submersed or with both floating and submersed leaves. Roots fibrous. Stem slender, often dimorphic, the lower stems rhizomatous, the upper ones erect, leafy, elongate, the tips often modified into turions, without tubers. Stipules forming a tubular sheath around stem. Leaves alternate or subopposite, entire to serrulate, venation parallel; submersed leaves thin, linear to orbicular; floating leaves often leathery, lanceolate, elliptic to ovate. Inflorescences axillary or terminal spikes or panicle of spikes. Flowers perfect, hypogynous, actinomorphic; perianth absent or of 4 free, rounded, short-clawed, segments in one series. Styles short; stigma capitate. Fruit drupaceous; seeds solitary.

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210. Ruppia maritima L., Sp. Pl. 1: 127 (1753). Syn.:

Ruppia rostellata Koch ex Rchb.

Vernacular name: rupia. Submersed herbs, perennial. Stems much branched, to 50 cm x 1 mm, terete, rooting at lower internodes. Stipular sheats 6–7.5 x ca. 0.5 mm. Leaves alternate to subopposite, sessile; blades linear, 4–7.5 cm x 0.5–1 mm, acute at apex, minutely serrulate at margins below apex. Inflorescences few-flowered, capitate spikes, at first enclosed by sheating leaf bases; peduncles 0.3–17 cm long. Gynoecium of 4–8 stipitate carpels. Fruit beaked, stipitate, 1.8–2 x 0.7–1.5 mm, the gynophore 1–2 cm long, the beak terminal, slightly recurved, 0.6–1 mm long. COLOMBIA. CH: (E). V: (E). C: (E). N: (S). ECUADOR. E: Bonifaz & Cornejo 4016 (GUAY). M: (S). G: Cornejo & Bonifaz 122 (GUAY). O: (S). PERU. T: (S). P: (E). Distribution: A nearly cosmopolitan species, occuring throughout the world. Ecology: Ruppia maritima is a submersed weed, inhabiting brackish waters with little to no current. It occurs in dry, moist, wet forest, and rainforest life zones. Biological interactions: In shrimp ponds, the high density of R. maritima populations affect shrimp commercial productivity because they consume oxygen in the water. The richly branched, intertangled stems and abundant linear leaves provide shelter to several shrimp larva predators, such as the small fish called “millonarios” (Spanish) and the voracious odonata larvae of dragonflies and damselflies. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that refers to the marine habitat. RHIZOPHORACEAE (Red Mangrove Family) Trees or shrubs; often with conspicuous prop or stilt roots. Branches often swollen at nodes. Stipules large, conspicuous, caducous, leaving an annular scar. Leaves opposite, decussate, simple, persistent, with gland-like corky protuberances sometimes present as small black dots beneath. Flowers bisexual or rarely unisexual, usually basally bracteolate; in axillary, simple or branched, usually cymose lax or condensed cluster-like racemose, or fascicled inflorescences, rarely solitary. Calyx lobes 3–6, valvate, persistent. Petals same in number as and alternate with calyx lobes, sometimes embracing 1–2 stamens, caducous or rarely persistent. Stamens usually twice as many as petals, the filaments free. Disk fleshy, annular, rarely absent. Ovary inferior or rarely semi-inferior or superior; ovules (1–)2(-many) per locule, pendulous; style 1, persistent; stigmas simple or more or less lobed. Fruit a coriaceous berry or drupe, or rarely a capsule, crowned by persistent calyx; seed 1, pendulous. This is the emblematic plant family of mangroves, it is represented by two genera: Rhizophora, a typical true mangrove genus, and Cassipourea.

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Key to species 1. Trees with arching prop roots; inflorescences cyme or cymose, with an obvious peduncle; petals 4, erose with a dense marginal pubescence; stamens 8; style conical; stigma bifid, occasionally trifid; fruits berry, with a linear-obclavate hypocotyl, 11–52 cm long 2. Inflorescence pauciflorous, simple or 1(–2)-branched, cymose with 2 or 4 flowers and/or cyme with 3 flowers; ovules cream; fully developed hypocotyls 13–30 cm long Rhizophora mangle 2. Inflorescence multiflorous, (3–)4–5(–6?)–branched, cymose, with (4–)8–64 (or more?) flowers; ovules often crimson-red to reddish; fully developed hypocotyls 11–52 cm long 3. Inflorescence slightly to not contracted, the pedicels 1.5–7 mm length; fully developed hypocotyls 23–52 cm length Rhizophora racemosa 3. Inflorescence loosely arranged, the pedicels 3–11 mm length; fully developed hypocotyls 11–25 cm length Rhizophora x harrisonii 1. Trees without prop roots; inflorescences fasciculate, without peduncle; petals 5, fimbriate; stamens numerous; style filiform; stigma inconspicuous, undivided; fruits capsular, elliptical, ca. 1 cm long Cassipourea killipii

211. Cassipourea killipii Cuatrec., Fieldiana Bot. 27(1): 103 (1950). Plate 38: D. Vernacular name: ajo. Trees to 10 m tall. Leaves opposite; blades coriaceous, ovate to ovate-elliptic, 8–12 x 4–6 cm, obtuse at base, acuminate at apex, the lateral veins 5–7 per side; petioles 0.8–1 cm long. Inflorescences axillary, fasciculate, pedicels ca. 1 mm long. Calyx cupular, gamosepalous, 4–5-lobed, 5–6 x 4 mm, green, glabrous without, the sepals triangular, 1–2 mm long. Petals ca. 1 cm long, fimbriate, white. Stamens ca. 24. Ovary superior; style 3–4 mm long. Fruits capsular. COLOMBIA. CH: Killip & Cuatrecasas 39141 (COL, NY). V: Killip 5218 (NY). N: (E). ECUADOR. E: Játiva & Epling 2185 (MO, NY [2]). Distribution: Western Colombia and northwestern Ecuador. Ecology: Often growing as a fringe species in mangroves and in the transitional areas to the adjacent terra firme wet forests and rainforests. Phenology: Flowering and fruiting throughout the year. IUCN Red list category: Due to factors such as a narrow range of distribution and a loss of habitat, this woody species is herein categorized as Vulnerable (VU B2abiii). Etymology: The epithet honors Ellsworth P. Killip (1890–1968), a North American botanist from the (US) National Herbarium of the Smithsonian Institution, and an energetic collector of vascular plants who gathered dozens of species new to science from the forests of Colombia and Peru.

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Notes: The type was collected in 1946 by José Cuatrecasas, near sea level, in Buenaventura Bay, Department of Valle del Cauca, Colombia. The cited collection represents the first record of this species for Ecuador. 212. Rhizophora mangle L., Sp. Pl. 1: 443 (1753). Plate 39: A–F. Syn.:

Rhizophora americana Nutt.

Vernacular names: mangle de concha, mangle colorado, mangle dulce, mangle gateado, mangle injerto, mangle macho, mangle rojo, mangle zapatero. Trees or shrubs 0.4–40 m tall, to 1 m dbh. Trunk erect to decumbent. Prop roots arching, to 10 m tall in well-developed individuals. Adventitious roots growing from branches, pendulous, ropelike when young. Terminal stipules 2, enfolding each other in a narrowly-conical arrangement, each narrowly-lanceolate, 4–8 cm long, glossy and finely striate with basal nectaries within, deciduous, leaving annular scars on branchlets. Leaves simple, decussate, grouped towards the end of branches; blades olive-green and glossy above, greenish-yellow and abundantly blackdotted beneath, ovate to broadly-elliptic, obovate or shortly-lanceolate, 6–14 x 3–8(–10) cm, cuneate to broadly-obtuse at base, obtuse to retuse and occasionally somewhat acuminate at apex, the lateral veins 9–13 per side, the intersecondaries present, brochidodromous, glabrous; petioles 0.4–1.5 cm long. Inflorescences supra-axillary, geminate racemes, cymes or cymose, 2.5–8 cm long, bearing 2–4(–5) flowers; primary peduncle 1–7 cm long; pedicels articulate, 5–12 mm long; bracteolar cup truncate to 2-dentate (even in a same inflorescence), often 4-dentate in cymes. Flower buds usually lanceolate to ovate, truncate and sometimes flattened to acute at apex. Hypanthium 2–4 mm long. Sepals 4, alternipetalous, free, thickly-coriaceous, lanceolate, 7–11 mm long, yellow-banana at maturity, persistent, glabrous. Petals 4, free, linearlanceolate, 6–9(–11) mm long, white, erose and with a dense marginal pubescence clasping and protecting anthers until maturity. Stamens 8, (sub)sessile (4 epipetalous, 4 alternipetalous), the anthers multilocellate, ca. 4 mm long, dehiscent by lateral flaps, complanate in crosssection, the connective apiculate at apex. Ovary semi-inferior, developing to superior in fruit, the ovules 4, creamish-white; style conical, 2–3 mm long, greenish, glabrous; stigma bifid, 0.5–1 mm long, yellow. Fruit brown, 2–3 cm long; hypocotyl 1, linear-obclavate, 15–30 cm long, pendulous, lenticellate, deep-green, brown at base, glabrous. COLOMBIA. CH: Fuchs & Zanella 22082 (NY). V: Gentry & Juncosa 40702 (NY). C: (E). N: RomeroCastañeda 5308 (NY). ECUADOR. E: Cornejo & Bonifaz 7092 (GUAY, QCNE). M: Cornejo & Bonifaz 7087 (GUAY, QCNE). SE: Cornejo & Bonifaz 2879 (AAU, GUAY). G: Bonifaz 94 (GUAY). O: Cornejo & Bonifaz 5985 (AAU, GB, GUAY, QCNE, S). GAL: Illescas 90 (GUAY). PERU. T: Vargas 35 (NY, US, WIS). P: (E).

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Distribution: Widely distributed along the tropical shores of West Africa and the Caribbean region, and from southern Florida to Brazil, as well as along the west coast of the Americas from Baja California to northwestern Peru, also in the Galapagos islands. Ecology: Rhizophora mangle is a typical mangrove and pioneer species. It is often found growing in very loose and periodically submersed soils in the intertidal zone, on muddy estuarine riverbanks, and occasionally on sandy beaches. R. mangle usually forms large stands, and very often grows intermixed with the vegetatively undistinguishable R. racemosa, R. x harrisonii, and other major mangroves such as Avicennia germinans, Laguncularia racemosa, and remaining mangrove flora and associate vegetation. The former species inhabits dry thorn scrub, very dry, dry, moist, wet forest, and rainforest life zones. A pure stand of Rhizophora mangle or intermixed with R. racemosa has been called a Rhizophoretum (Acosta-SolĂs, 1959). A video of a Rhizophoretum can be seen on this link: http://www.youtube.com/watch?v=z9qtytgpwaE Phenology: The sepals, which enclose the entire flower bud (closed calyx aestivation), are externally deepgreen when immature. Before the flower opens, the sepals become a banana-yellow color, indicating the anthers have reached maturity and the pollen grains have been released into the interior of the still-closed flower bud. An inner whorl of white petals, shorter and more delicate in appearance than the sepals, are covered with abundant marginal hairs, which clasp and tightly hold the epipetal subsessile anthers until the flower opens. Two days before fully opening, the flower bud sutures located between the sepals slowly begin to dehisce, longitudinally splitting upwards until the sepals separate almost entirely lengthwise, while still remaining joined at the base. The flowers open slowly, through day and night, producing no scent or nectar. The first day, the flower opens at a speed of about 1 mm each five hours. The just-opened flower has an urceolate shape that is retained for at least seven hours. At the early blooming, the initially tight arrangement of petals hold the by then detached and dehisced anthers, preventing them from falling. The anthers, at this stage, still keep abundant pollen grains attached to the marginal hairs and to the interior of the flower. During the subsequent hours the calyx sepals slowly progress into a more divergent position. The stamens and petals are deciduous; the stamens fall off aproximately one day after the flower begins opening, and the petals fall off approximately one day later. The flowering occurs in succession, first one flower out of each pair in the inflorescence, followed by the second flower one to two days afterward (observations made in Rio Chone estuarium, Province of Manabi, Ecuador, see Cornejo and Bonifaz, 2006). Biological interactions: The newly opened flowers are visited by minuscule, nearly invisible to naked eye, whitish flower mites of Hattena rhizophorae (Acari, Ameroseiidae, Faraji and Cornejo, 2006; Plate E:A). No flower mites of H. rhizophorae have been observed in the flower buds before sepal dehiscence begins (observations made on mature closed flower buds dissected in situ). The mites are eager to access the flower interior immediately after the dehiscense of the inter-sepal lateral sutures. Small populations of H. rhizophorae are sometimes found on the tip of entirely closed mature flower buds, waiting for the suture dehiscence. The flower mites have been observed (under 10 x) to transport pollen grains attached to their bodies and legs while moving throughout the entire flower interior, even between the stigma lobes. As the flowers do not produce nectar or scent, it is most likely that H. rhizophorae feeds on the Rhizophora pollen grains. It is still unknown whether H. rhizophorae plays a roll in the pollination of this species. These mysterious flower mites are presumed to be transported by birds, but their entire life cycle is yet undocumented.

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The opened flowers of R. mangle are also occasionally visited by wasps and small ants, which also transport pollen grains attached to their bodies, but these species are not regarded as pollinators. The stout prop roots of well-developed individuals of R. mangle may be inhabited by epiphytic plant species (mainly Orchidaceae, Bromeliaceae, Araceae, and ferns) that often display a luxuriant appearance, especially in the Chocoan mangroves. Several species of Lepidoptera larvae feed on the leaves of R. mangle, and they occasionally become a plague. Dense forests of R. mangle provide shelter for a multitude of vertebrate and invertebrate species. Coastal birds such as Egrets (e.g. Ardea alba), Brown pelicans (Pelecanus occidentalis), and Cormorants (Phalacrocorax brasiliensis) build their nests on stout developed branches of R. mangle. Sometimes snakes such as the â&#x20AC;&#x153;matacaballoâ&#x20AC;? (which means horse killer, though it does not kill horses) Boa constrictor may reach the bird nests to feed on incubating females, eggs and nestlings. Bats such as the Greater Bulldog Bat or murciĂŠlago pescador mayor (Noctilio leporinus subsp. mastivus) are occasionally found inhabiting the interior of holes in R. mangle trunks (Salas, 2010). Some mammals as deer sometimes feed on fallen inflorescences and flowers of red mangroves. Individuals of R. mangle of higher development, with prop roots of at least 6 m in diameter above ground, are of vital importance for the Red crab or guariche (Ucides occidentalis) of commercial sizes which built burrows underground. In the Guarichal de Salinas, Rio Chone estuarium (Province of Manabi, Ecuador), ca. 300 mature comercial size Red crab individuals were recorded within 1000 m2 (in a transect of 10 x 100 m) in a mature Rhizophora forest (Cornejo, 2005). The Red crab is the largest crab inhabitting the muddy soils at the mangrove interior, and when they reach commercial size they may find it difficult otherwise impossible to build underground settlements under narrow diametrically developed young Rhizophora trees. In the intertidal zones, the periodically submersed prop roots are the substrate for Oysters such as Crassostrea columbiensis (Ostreidae), as well as dozens of microscopic and macroscopic epiphytic algae species, and provide shelter and refuge to many fish species, several of those of commercial size such as corvina (Cynoscion albus), lisa (Mugil cephalus), pargo (Lutjanus sp.), etc. Fishermen surround the submersed prop roots with nets when the tidal level is high; the nets are removed at the lowest tide with the captured fish inside. Finally, the leaves and prop roots provide an essential nursery habitat for larvae, myriad juveniles fish species, and marine invertebrates, many of them commercially valuable and nutritionally significant for human coastal populations. The complex ecological interactions and benefits associated with R. mangle make it one of the most valuable species in the Neotropics and the world. Rhizophora mangle hypocotyls that are still attached to the mother plant have been observed to be infested by insect larvae of Coccotrypes sp. (Coleoptera). Coccotrypes sp. is a small bettle (ca. 3 mm long) that carves holes and lays eggs in the surface of hypocotyls. After eclosion, the Coccotrypes sp. larvae penetrate into the hypocotyl, building irregular short labyrinths in the direction of the subwoody interior, thus destroying the viability (see selecting hypocotyls for reforestation). Once Coccotrypes sp. reaches maturity, it abandons the infested hypocotyl, mates, and lay eggs in other hypocotyl, hence closing the cycle. Coccotrypes sp. may heavily infest a whole population of R. mangle, becoming a plague.

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Dispersal and settlement: This is a water dispersed species. Once shed from the mother plant, the hypocotyls float horizontally in the estuary water and are carried out by currents. While the brackish water is slowly absorbed into the hypocotyls, air from the interior is released in micro-bubbles through the lenticels. The water accumulates in the subwoody and wider base of the hypocotyls, which become heavier and cause to float vertically, then to stand in the loose mud. Growth of leaves and/or roots is not achieved while the hypocotyls are still afloat; these develop only after mud implantation. Prop roots develop from very young saplings, of less than 80 cm tall. Prop roots provide this species a remarkable stability for successful settlement in periodically inundated very soft and loose muddy soils. Uses: The wood of this species is used in heavy construction, such as in pier pillars and in ship keels and ribs (Little, 1969; Madsen et al., 2001), because of its hardness and high resistance to marine and brackish waters. The trunk wood is commonly used as columns or planks in house construction. The columns of some of the old “barrio Las Peñas” houses, in the city of Guayaquil, are made of straight thick trunks having an estimated age of 100 years old, and are still in perfect condition. In the past, the wood from straight trunks was used atop and underground muddy soils in the building of railroad tracks called “durmientes” (Spanish, Acosta-Solís, 1944). The combusted wood produces high quality charcoal. Tannins are occasionally extracted from the bark and used to tan leather. The bark from dry branches of less than 1 inch in diameter is used to elaborate hammocks, chairs, and crafts. Medicinally, a bark infusion is imbibed as a cough suppressant. The bark also can be boiled in water, and the leaves added to the infusion, to be inhaled as a “bajo al aire” to cure colds. Native people from the coast of Ecuador used the terminal tender roots of Rhizophora to soothe stings and bites from poisonous animals. This species can concentrate heavy metals as: Cadmium (Cd), cupper (Cu), lead (Pb), and zinc (Zn) (Campos and Gallo, 1997; Villamil, 2010). IUCN Red list category: According to the Registro Oficial No. 148, issued on March 16 1993, Rhizophora mangle is categorized as an endangered species in Ecuador (INEFAN/GEF, 1998). However, due to factors such as a widespread range of distribution, commonly healthy populations almost always comprised of abundant individuals, a high rate of hypocotyl viability, and abundant seedlings and juvenile individuals, it is deemed its categorization as of Least Concern (LC) (Polidoro et al., 2010). Selecting hypocotyls for reforestation: The mature hypocotyls can be harvested from the mother plant and from the ground up to a few days after they have fallen, and be directly planted in the ground; pots or bags should not be used for reforestation. Only hypocotyls of healthy appearance should be collected. Dehydrated hypocotyls that are shrunken, desiccated, opaque or that exhibit physical damage must be rejected (Duke and Allen, 2006). Also, the Coccotrypes infestation -if present- must be observed. It can be detected by the presence of small holes ca. 2 mm in diameter on the hypocotyl surface. When the Coccotrypes holes are located at the base, below the broadest part, it is likely that the hypocotyl is still viable. However, if those holes are located in or above the broadest part of the hypocotyl, then viability is lost; the hypocotyls are useless for reforestation. Based on traditional knowledge, local people recommend harvesting the hypocotyls during the full moon to ensure more vigorous seedlings and faster growth.

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Etymology and pre-Linnean names: The generic epithet is formed from the Greek prefix rhizo (root), and the Greek suffix phoros (to bear). This may refer to the very distinctive prop roots of this taxon (e.g., Tomlinson, 1986). However, it has been argued that W. Piso (=G. Pisone, from Piso and Marcgrave, 1648) “considered the rhizophore to be the exposed hypocotyl of the viviparous propagule itself” (=the embryo; Menezes, 2006); see also the fruit description provided by Plukenet (1696). It is stated that the generic name Rhizophora was coined in 1747 by Everardus Jacobus van Wachendorff (1702–1758), who studied the cultivated plants in Holland (Austin, 2004). Thus Rhizophora is a pre-Linnean name that was subsequently adopted and formally proposed in Species Plantarum (Linnaeus, 1753). In Historia Naturalis Brasiliae (Piso and Marcgrave, 1648), one of the earliest publications of plants in America that deals with R. mangle, the species was given the pre-Linnean name of “Mangue guaparaiba”, “mangue” being the vernacular name for R. mangle in Brazil. R. mangle was also named and described as “Mangle arbor Pyrifoliis salsas and uliginosis locis in America proveniens; fructu oblongo tereti, summis ramis radicola” (Plukenet, 1696). Subsequently, “Mangles”, an “American name” that means thicket, was formally proposed as a generic name for this species, and its authorship was attributed to Piso (Plumier, 1703). Linnaeus (1753), however, used “mangle” as the specific epithet, presumably because it was considered to be of barbaric origin. Notes: The flowers of R. mangle usually have calyces of four sepals, but once in a while can produce a flower of five or even six sepals (Cornejo 7890, GUAY), which is regarded as abnormal. Variability of primary peduncle length (between 1.5–5 cm long) has also been observed, even in a same tree; therefore, the primary peduncle length it is not considered of taxonomical significance. The trunk position can vary from erect and straigth to fully decumbent. In the later case the trees develop thick lateral arching prop roots arranged along the trunk, hence “mangle gateado”, one of the vernacular names, which means the catlike walking mangrove. Sometimes native people in the field differentiate the two as different types of mangrove trees, and undoubtely the wood of each is used for different purposes; however, it is most likely that the trunk disposition (erect vs. decumbent) is a result of the different consistency of soils, and does not have any taxonomical significance. 213. Rhizophora racemosa G. Mey., Prim. Fl. Esseq. 185 (1818). Plates 40: A–F; 41: A. Syn.:

Rhizophora mangle var. racemosa (G. Mey.) Engl.

Vernacular names: mangle, mangle caballero, mangle cholo, mangle colorado, mangle gateado, mangle patucho, mangle pava, mangle pecho de pavo, mangle rojo, manglillo, palo salado. Trees or shrubs 1–40(–50?) m tall, to 1 m dbh. Trunk erect to decumbent. Prop roots arching, to 10 m tall in well-developed individuals. Adventitious roots growing from branches, pendulous, rope-like when young. Terminal stipules 2, enfolding each other in a narrowlyconical arrangement, each narrowly-lanceolate, 4–8 cm long, glossy and finely striate with basal nectaries within, deciduous, leaving annular scars on branchlets. Leaves simple,

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decussate, grouped towards the end of branches; blades olive-green and glossy above, greenish-yellow and abundantly black-dotted beneath, lanceolate or ovate to elliptic-oblong or obovate, 5.5–18 x 2–7(–10) cm, cuneate to broadly-obtuse and somewhat decurrent onto petiole at base, acute to truncate at apex, the lateral veins 11–15 per side, the intersecondaries present, brochidodromous, glabrous; petioles 0.7–2 cm long. Inflorescences supra-axillary, cymose, 4–12 cm long, bearing 8–64 flowers; primary peduncle 1.5–7 cm long; pedicels articulate, 1.5–7 mm long; bracteolar cup 2–4-dentate (even in a same inflorescence). Flower buds usually ovate to lanceolate, rarely elliptical, acute to obtuse at apex. Hypanthium 2–4 mm long. Sepals 4, alternipetalous, free, thickly-coriaceous, lanceolate, 6–11 mm long, yellowbanana at maturity, persistent, glabrous. Petals 4, free, linear-lanceolate, 6–8 mm long, white, erose and with a dense marginal pubescence clasping and protecting anthers until maturity. Stamens 8, (sub)sessile (4 epipetalous, 4 alternipetalous), the anthers multilocellate, ca. 5 mm long, deshicent by lateral flaps, triangular in cross-section, at maturity with 2 longitudinalmarginal granulose ribs dorsally arranged on each side of connective, the connective apiculate at apex. Ovary semi-inferior, developing to superior in fruit, the ovules 4–5, crimson-red to reddish; style conical, 2–3 mm long, greenish, glabrous; stigma bifid, occasionally trifid, 1–2 mm long, yellow. Fruit brown, 2.5–4 cm long; hypocotyl 1, linear-obclavate, 11–52 cm long, pendulous, lenticellate, deep-green, brown at base, glabrous. COLOMBIA. CH: (E). V: Gentry et al. 40429 (NY). C: Cabrera 4136 (CUVC). N: (S). ECUADOR. E: Cornejo & Bonifaz 1885 (AAU, GUAY, QCNE [2]). M: Cornejo & Bonifaz 7084 (GUAY). SE: (E). G: Cornejo & Bonifaz 6760 (COL, GB, GUAY). O: Cornejo & Bonifaz 5982 (GB, GUAY). PERU. T: Smith 10085 (MO). Distribution: Widely distributed along the tropical shores of West Africa and the southern Caribbean region (Trinidad and Tobago), and from Venezuela to Brazil, as well as along the west coast of the Americas from Honduras to northwestern Peru. Ecology: Rhizophora racemosa is a truly mangrove species that inhabits loose muddy saturate and saline to brackish soils. This species usually forms large stands, often growing intermixed with populations of R. mangle and R. x harrisonii. R. racemosa is occasionally dominant seaward in the hyposaline mangroves, and occurs in the dry thorn scrub, very dry, dry, moist, wet forest, and rainforest life zones. Phenology: Similar as in R. mangle. Once the flower buds progress from immature deep-green to a mature banana-yellow color, the longitudinal sutures located between the sepals begin splitting basally, upward from the base. The process takes several hours, until the sepals are entirely separated from each other. Initially, the newly opened inter-sepal lateral grooves remain partially open at the base, while abundant flower mites of Hattena rhizophorae enter the flower bud. The flower mites presumably feed on the pollen grains, and remain in the flower for approximately one day. The inter-sepal splitting process takes several hours, during which the sutures slowly open longitudinally towards the bud apex. Finally,

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the flower often opens abruptly, and the calyx suddenly takes the shape of a cup. At this point the already detached anthers are usually still held by the marginal pubescence of the petals, but fall off few hours later. The petals remain in the flower interior approximately one day after the flower opens, depending chiefly on the force of wind. The flowers open at any time during the day or night, and do not produce any scent or nectar (observations made in Rio Chone estuary, Province of Manabi, Ecuador, see Cornejo and Bonifaz, 2006). Biological interactions: Same as in R. mangle. Dispersal: Same as in R. mangle. Uses: Same as in R. mangle. IUCN Red list category: Due to factors such as a widespread range of distribution, a common presence of healthy populations comprised of abundant individuals, a high rate of hypocotyl viability, and abundant seedlings and juvenile individuals, it is deemed the categorization of this species as of Least Concern (LC) (Polidoro et al., 2010). Selecting hypocotyls for reforestation: Same as in R. mangle. Etymology: The epithet, which means raceme-like, refers to the characteristic many-flowered inflorescences. Notes: The flowers of R. racemosa are highly similar and often undistinguishable from those of R. mangle, the latter being an indisputable sister species with a tendency to produce somewhat larger flowers than in R. racemosa. The size of the flowers of both species intergrades, and therefore cannot be used as a consistent character to separate the American species of Rhizophora. It is not possible to distinguish R. racemosa from R. mangle based on vegetative characteristics only. As described in the key, the main character separating these species is the number of flowers per inflorescence.The position of the trunk in R. racemosa, like in R. mangle, may vary from fully decumbent to entirely erect, even in a same locality. Finally, the fruits of R. racemosa rarely are 2-embryoned, producing twin hypocotyls (Cornejo & Bonifaz 6751, GUAY). Discussions: Rhizophora racemosa has been commonly treated as R. harrisonii (Dinaf-Clirsen, 1991; Jørgensen and León, 1999; Valverde, 1988; West, 1956, 1977), and occasionally has been regarded in a wide sense under R. mangle (Acosta-Solís, 1966; Little, 1969). Despite a previous report of R. racemosa in Ecuador (Prance et al., 1975, based on Little & Dixon 21219, NY; and Játiva & Epling 810, NY), this species was not recorded or discussed for the country; rather R. harrisonii was cited (Jørgensen and León, 1999).

The anthers of R. racemosa have been described as having a multiporicidal dehiscence

(Barbosa, 1988, cited as R. harrisonii). Nevertheless, the anthers are indistinguishable from those of R. mangle, which are deshicent by lateral flaps that shed at maturity to reveal the multilocellate pits containing pollen grains.

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Rhizophora x harrisonii Leechm., Harvard Pap. Bot. 18(1): 37 (2013). Bas.: Rhizophora harrisonii Leechm. Syn.: Rhizophora brevistyla Salvoza Rhizophora x harrisonii is characterized by the many-flowered and loosely arranged inflorescences, often bearing somewhat longer pedicels (3–11 mm length), and shorter hypocotyls (11–25 cm length) than in R. racemosa. Distribution: Rhizophora x harrisonii is found on both coasts of tropical America and in West Africa, along the same range of distribution than R. mangle and R. racemosa. Ecology: Same as in R. racemosa. Phenology: Same as in R. racemosa. Biological interactions: Same as in R. mangle and R. racemosa. Dispersal: Same as in R. mangle and R. racemosa. Uses: Same as in R. mangle and R. racemosa. IUCN Red list category: Same as in R. racemosa. Selecting hypocotyls for reforestation: Same as in R. mangle and R. racemosa. Etymology: The epithet honors Mr. John Burchmore Harrison (1856–1928), Director of the Department of Science and Agriculture and goverment geologist of the British Guiana. Notes: It has been observed in the field that the more flowered inflorescences have a weaker appearance and a tendency to produce somewhat smaller flowers (Cornejo & Bonifaz 7091, GUAY). On the other hand, the lesser flowered inflorescences may produce somewhat larger flowers. The variability of floral sizes in R. x harrisonii may be exhibited in the same individual.

Rhizophora x harrisonii individuals represent morphotypes produced by ongoing hybridization

and introgression between the often-sympatric R. mangle and R. racemosa. Breteler (1969, 1977) also supports R. x harrisonii as hybrids with low pollen fertility compared with parental R. mangle and R. racemosa. However, he also observed pollen malformation in individuals that morphologically resemble either R. mangle or R. racemosa suggesting that they could be advanced generations of hybrids (Breteler, 1969). Thus, both molecular evidence and pollen morphology support that introgressive hybridization is a common process in the New World Rhizophora (Cerón-Souza et al., 2010). RUBIACEAE (Coffee Family) Usually trees or shrubs, less often herbs or vines, sometimes epiphytic. Stipules present but often caducous, interpetiolar, intrapetiolar, or sheating. Leaves opposite or whorled; blades simple, usually entire. Inflorescence usually a dichasial cyme, sometimes capitate, umbellate, thyrsoid, or spicate, or reduced to a solitary flower. Flowers usually bisexual. Calyx usually gamosepalous. Corolla gamopetalous. Stamens

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of same number as and alternating with corolla lobes, inserted on corolla, the filaments usually free. Ovary inferior. Fruit a capsule, schizocarp, drupe, or berry. Key to the species 1. Shrubs or trees; stipules 1–3.2 cm long; corollas 1.1–9 cm long; fruits cylindric or obovoid, (sub) terete, 0.7–10 cm long 2. Leaf blades with 11–16 lateral veins per side; corolla 1.1–1.4 cm long, reddish to purple; fruits obovoid, 0.7–1.2 cm long

Rustia occidentalis

2. Leaf blades with 4–6 lateral veins per side; corolla 4–9 cm long, white; fruits cylindric, 3–10 cm long 3. Leaf blades 4–10 cm wide; inflorescences terminal racemes; corolla 6–9 cm long; fruits ca. 8–10 cm long Cosmibuena macrocarpa 3. Leaf blades 1.5–3.5 cm wide; inflorescences solitary flowers; corolla 4–5.5 cm long; fruits 3–6 cm long

Hillia maxonii

1. Lianas, epiphytic vines or shrubby herbs; stipules 0.1–0.4 cm long; corollas 0.3–0.8 cm long; fruits orbicular or subglobose, 0.1.5–0.7 cm long 4. Shrubby herbs; inflorescences subsessile heads, less than 1 cm long

Tobagoa maleolens

4. Lianas or epiphytic vines; inflorescences racemose to/or paniculate, 2–10 cm long 5. Lianas, several to many meters long; inflorescences axillary; fruits laterally compressed, white at maturity Chiococca alba 5. Epiphytic vines, to 1 m long; inflorescences terminal; fruits subglobose, red turning black at maturity

Notopleura epiphytica

214. Chiococca alba (L.) Hitchc., Annual Rep. Missouri Bot. Gard. 4: 94 (1893). Plate 41: B.

Bas.:

Lonicera alba L.

Syn.:

Chiococca racemosa L.

Chiococca brachiata Ruiz & Pav.

Chiococca trisperma Hook f. Lianas (also shrubs or trees in other localities). Stipules often broader than long, 1–2 mm long, abruptly acuminate or mucronate. Leaf blades coriaceous, ovate, lanceolate or elliptic, 2–10 x 1–5 cm, obtuse to rounded and short decurrent at base, acute to short acuminate at apex, dark-green and lustrous above, paler beneath, glabrous; petioles 2–12 mm long. Inflorescences axillary, racemose to paniculate. Calyx and hypanthium 2–2.5 mm long. Corolla 6–8 mm long, white, the lobes triangular. Stamens included. Berry orbicular, 4–8 mm long, laterally compressed, white at maturity, glabrous.

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COLOMBIA. CH: (E). ECUADOR. SE: Cornejo & Bonifaz 7161 (GUAY, MO). Distribution: Southern United States to Paraguay. Ecology: This widespread species mostly inhabits terra firme ecosystems; in moist forest, wet forest, and rainforest life zones. Ecological interactions and dispersal: No observations have been recorded, but it is most likely that birds feed on the small fleshy berries, digest the mesocarp, and disperse the seeds endozoochoreously. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word that means white, referring to the white color of the corollas and fruits at maturity. Discussion: The populations of this species that inhabit from Mesoamerica to Bolivia (including western Ecuador) exhibit laterally compressed fruits. However, populations of this species in the Galapagos islands, as well as in other countries as Puerto Rico have subglobose fruits. 215. Cosmibuena macrocarpa (Benth.) Klotzsch ex Walp., Repert. Bot. Syst. 6: 69 (1846). Plate 41: C. Bas.: Syn.:

Buena macrocarpa Benth. Cosmibuena gorgonensis Wernham Cosmibuena paludicola Standl. Trees to 15 m tall. Terminal branchlets with annular scars. Terminal stipules oblanceolate to elliptic, 1.8–2 x 0.6–0.8 cm, rounded at apex, glabrous, soon deciduous. Leaves simple, opposite; blades coriaceous, obovate to elliptic, 7.5–20 x 4–10 cm, entire, the lateral veins 5–6 per side, ascendent, inconspicuosly pilose; petioles 1–2 cm long. Inflorescences terminal racemes; peduncles 5–30 mm long; bracts obovate to ligulate, 6–11 mm long; pedicels 9–16 mm long. Flowers 3–8, 5-merous, white to light-yellow. Calyx 2–9 mm long, lobed. Corolla tube 6–9 cm long, the lobes elliptic to oblong, 2–3 cm long, obtuse to rounded at apex. Ovary 6–13 mm long; style 1–2 mm, slightly longer than corolla tube; stigma 6–9 mm long. Capsules erect, cylindric, ca. 8–10 x 0.7 cm, greenish; seeds 7–9 mm long, winged.

COLOMBIA. CH: (E). V: Gentry 40715 (MO). C: (E). N: (S). ECUADOR. E: Cornejo & Cangá 8134 (GUAY). Distribution: Costa Rica to Peru. Ecology: In wet forests and rainforests. Biological interactions: The elongate narrow corollas produce a strong scent at night, suggesting pollination by moths. Dispersal: The seeds are wind dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet is formed by the Greek words makros (=large) and carpos (=fruits), referring to the relatively large fruit size. Note: The type was collected by the Englishman Richard Brinsley Hinds (1812–1847) on Gorgona Island, Department of Nariño, southwestern Colombia, during the voyage of H.M.S. Sulphur.

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216. Hillia maxonii Standl., J. Wash. Acad. Sci. 18: 163 (1928). Syn.:

Cosmibuena rhizophorae Standl. Suffrutescents herbs or shrubs to 5(–15) m tall. Stipules 1.2–3.2 x 0.8–1 cm. Leaf blades coriaceous, elliptic to oblanceolate or suborbicular, 2.5–10 x 1.5–3.5 cm, cuneate to attenuate at base, obtuse to rounded at apex, the lateral veins 4–5 per side, glabrous; petioles 3–8(–15) mm long. Flowers solitary, 4-merous; peduncles 1–2(–4) mm long; bracts lacking or 1–3 mm long. Calyx 6–7 mm long, lobed. Corolla white, the tube 4–5.5 cm long, the lobes elliptic, 1.5–2.7 cm long, rounded at apex. Ovary 3–10 mm long; style ca. 15 mm long, shorter than corolla tube; stigma ca. 7 mm long. Capsules erect, cylindric, 3–6 x 0.5–0.9 cm, smooth to slightly ridged longitudinally, green; seeds 2–4 mm long, with filaments 6–13 mm long.

COLOMBIA. V: Killip 34972 (COL, F, the type of Cosmibuena rhizophorae). Distribution: Nicaragua to northwestern Ecuador. Ecology: In wet forests and rainforests. Dispersal: The seeds are wind dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet honors William R. Maxon (1877–1948), a North American botanist and taxonomist of ferns, who collected the type from Nicaragua in 1923. Note: The Cosmibuena rhizophorae type was collected in 1939, from the mangroves of Buenaventura, Department of Valle del Cauca, Colombia. 217. Notopleura epiphytica (K. Krause) C. M. Taylor, Ann. Missouri Bot. Gard. 88(3): 512 (2001). Bas.: Syn.:

Psychotria epiphytica K. Krause Psychotria orchidearum Standl. Holoepiphytic vines, to 1 m long. Stipules forming a truncate sheath around stem, 0.5–1.5 mm long, deciduous. Leaf blades coriaceous, elliptic to elliptic-oblong or oblanceolate-elliptic, 4.5–10 x 1–3 cm, obtuse to rounded at base, acute to acuminate at apex, the lateral veins 2–5 per side, rather inconspicuous, glabrous; petioles 1–6 mm long. Inflorescences short terminal panicles; peduncles 0.5–2 cm long; bracts reduced. Calyx ca. 2 mm long, dentate. Corolla white, the tube 4–8 mm long, the lobes 1.5–2 mm long. Fruits subglobose, 4–7 x 3–6 mm, red turning black at maturity; seeds 2–4.

COLOMBIA. V: Killip 34988 (COL). Distribution: Nicaragua to Bolivia. Ecology: In wet forests and rainforests. Biological interactions and dispersal: The fleshy fruit is most likely eaten by birds, while the small seeds, embedded in a juicy mesocarp, are endozoochorously dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet is a Latin word, referring to the epiphytic habit.

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218. Rustia occidentalis (Benth.) Hemsl., Biol. Centr.-Am., Bot. 2: 14 (1881). Plate 41: D. Bas.:

Exostemma occidentale Benth. Trees or shrubs 2–8 m tall. Stipules narrowly-triangular, 1–3 x 2.5–3 cm, present only in bud. Leaf blades chartaceous, narrowly-elliptic, lanceolate to oblanceolate, 17–28(–40) x 4–12 cm, cuneate to acute at base, acute-caudate at apex, the lateral veins 11–16 per side, glabrous; petioles 2–4 mm long. Inflorescences reduced, paniculate to racemoid, 4–15 cm long. Flower buds clavate. Calyx up to 3 mm long, persistent. Corolla 5-petala, 11–14 mm long, reddish to purple. Stamens with yellow anthers, the anthers 5–7 mm long, opening by 2 apical pores. Style 10–14 mm long. Capsules obovoid, 7–12 x 7–9 mm, with minute lenticels, yellow at maturity; seeds numerous.

COLOMBIA. CH: Fuchs & Zanella 21783 (COL, F, MO, S, US). V: Killip 38680 (MO). C: (E). N: (S). ECUADOR. E: Cornejo & Bonifaz 7257 (GUAY). Distribution: Nicaragua to northwestern Ecuador. Ecology: Rustia occidentalis is mostly distributed through fresh water swampy coastal areas, such as the Natales, and occurs as a fringe species in more compact, yet still wet-muddy-hyposaline soils in mangroves. It occurs in the wet forest and rainforest life zones. Biological interactions: The flowers are mainly visited by bees such as Euglossa sp. and Melipona sp. (Delprete, 1999). IUCN Red list category: Least Concern (LC). Dispersal: The seeds are wind dispersed. Etymology: The epithet refers to the first known distribution of this species. The type was collected by the Englishman Richard Brinsley Hinds (1812–1847) on Gorgona Island, Department of Nariño, southwestern Colombia, during the voyage of H.M.S. Sulphur. Discussion: Rustia occidentalis has been referred to as a minor mangrove (Chapman, 1975). But its pattern of distribution, mostly through fresh water swampy coastal areas, and primarily its wind dispersed seeds, indicate that R. occidentalis should not be regarded as a minor mangrove, but as a swamp species that has the ability to settle in hyposaline environments. The fact of it having wind dispersed seeds unquestionably excludes it from any of the mangrove categories herein proposed. 219. Tobagoa maleolens Urb., Repert. Spec. Nov. Regni Veg. 14 (405/408): 343 (1916). Shrubby herb to 1 m tall. Stipules 3-linear-digitate, 1.5–4 mm long. Leaf blades thinly chartaceous, very narrowly oblong-elliptic, 3–10 x 2–3 cm, cuneate at base, tapering at apex, the lateral veins ca. 5–6 per side, shortly pilose beneath; petioles 3–10 mm long. Inflorescences axillary heads, 10–20-flowered. Calyx lobes triangular to lanceolate, ca. 0.5–0.6 mm long, persistent. Corolla ca. 3 mm long, white. Stamens inserted just below sinuses within the corolla tube. Style 2–4 mm long. Capsules subglobose, ca. 1.5 x 1 mm, green; seeds numerous.

246

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

COLOMBIA. V: Gentry & Juncosa 40723 (COL, MO). Distribution: Mesoamerica to Colombia and Venezuela, and in the West Indies. Ecology: In wet forests and rainforests. IUCN Red list category: Least Concern (LC). Etymology: The generic epithet refers to Tobago, where the species was initially known to occur. The specific epithet is formed by the Latin words malus (=bad) and olens (=smell), referring to the plant’s unpleasant odor. SCROPHULARIACEAE (Figwort or Snapdragon Family) Herbs, vines or shrubs, sometimes epiphytic. Stipules absent or small pseudostipules present. Leaves opposite, alternate, whorled or sometimes all basal; blades entire to pinnately or palmately lobed. Inflorescence a spike, raceme, thyrsoid panicle, fascicle or solitary flower; bracts foliaceous to reduced. Flowers bisexual, 5- to 4-merous. Calyx persistent, the sepals distinct or variously united. Corolla bilabiate to nearly regular. Stamens 5, 4 or 2, inserted on corolla. Ovary superior. Fruit a capsule or berry. 220. Schlegelia darienensis Sandw., Kew Bull. 1930: 212 (1930). Hemiepiphytic woody lianas. Branchlets subterete, glabrous. Leaves simple, opposite, (sub) sessile; blades coriaceous, ovate-elliptic, 7–16 x 4–10 cm, rounded at base, obtuse to apiculate at apex, minutely scattered–lepidote beneath, otherwise glabrous, with a glandular field near base of blade, drying olive to olive-brown. Inflorescence a many-flowered terminal elongateracemose panicle, the lateral braches contracted and almost fasciculate, each inflorescence node subtended by a pair of oppossite foliaceous bracts. Calyx cupular. Corolla white to pink, tubular with reflexed lobes, 1.5–2 cm long. Stamens didynamous. Pistil 3–4 mm long; ovary spherical, 2 mm diam., glabrous. Fruit a globose fleshy berry, ca. 1 cm diam., glabrous. ECUADOR. E: Cornejo & Bonifaz 6936 (GUAY). Distribution: Southeastern Panama to western Ecuador. Ecology: In wet forests and rainforests. Biological interactions: No information is recorded, but it is most likely that the fleshy berries are eaten by birds and bats, and the seeds are endozoochoreously dispersed. IUCN Red list category: Least Concern (LC). Etymology: The epithet refers to the Province of Darien in Panama, where the species was first known to occur.

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TETRAMERISTACEAE Trees. Leaves alternate, sessile, grouped at apex of branchlets; blades asymmetrical, the lateral veins inconspicuous. Flowers large, axillary (apparently terminal), bisexual, solitary, subtended by large, foliaceous bracts. Sepals 5, distinct, unequal, deciduous. Petals 5, alternisepalous, free, caducous. Stamens 5. Gynoecium narrowly-conical, the ovary superior; stigma punctate. Fruits indehiscent; seed 1. This is a pantropical family that comprises three genera. In our area it is represented by Pelliciera, a monotypic genus, distributed in American mangrove swamps (Kobuski, 1951; Pool, 2001). Pelliciera has been commonly placed in Theaceae (Tea family) or even regarded to its own family, Pellicieraceae (Piñuelo mangrove family). 221. Pelliciera rhizophorae Planch. & Triana, Ann. Sci. Nat., Bot., sér. 4, 17: 381 (1862). Plate 42: A–C. Vernacular names: mangle piñuelo, piñuela, piñuelo. Trees to 30 m tall, and 50 cm dbh. Buttresses narrow, irregularly conical, to 1.5 m tall. Branches with petiole scars. Stipules absent. Terminal buds narrowly-conical, ca. 8 cm long. Leaves simple, alternate, sessile, grouped at apex of branchlets; blades coriaceous, narrowlyelliptic, asymmetric, 7–17 x 2–4 cm, attenuate at base, acute at apex, when young glandular on distal margin along wider side of blade, becoming apparently entire with age, the lateral veins 8–12 per side, inconspicuous, glabrous. Flowers solitary, sessile, lateral and subterminal, showy, 8–10 cm diam., white. Floral bracts 2, patent at anthesis, oblong-lanceolate, ca. 5 cm long, light green, denticulate at margins. Sepals 5, unequal, ovate to obovate, 1.5–2 cm long, shortly-acuminate at apex. Petals 5, lanceolate, 5–7 x 0.7–1 cm, glandular at base, acute at apex, widely spreading at anthesis. Stamens 5, alternipetalous, the filaments filiform, arranged for most of their length closely appressed in and along longitudinal grooves of ovary, the anthers linear-oblong, pendulous. Gynoecium narrowly-conical, divided almost equally into a longitudinal 10-sulcate ovary and a smooth style, the ovary superior; stigma punctate. Fruit napiform, thickly coriaceous (apparently woody or subwoody when dry), deltate to subcordate, 9–12 x 6–8 cm, longitudinally sulcate, somewhat dorsoventrally complanate, acuminate-rostrate at apex, dark-brown to reddish-brown, lenticellate, glabrous, on a short and thick peduncle; seed 1, consisting of 2 large fleshy obcordate red cotyledons, the radicle firm, pointed, the plumule large, curved. COLOMBIA. CH: Fuchs & Zanella 21841 (MO, NY [2]). V: Killip 5222 (NY). C: (E). N: RomeroCastañeda 5298 (NY). ECUADOR. E: Bonifaz & Cornejo 4011 (AAU, GB, GUAY, QCNE). Distribution: In scattered patches located along the Pacific coast from Costa Rica to the Muisne estuary in northwestern Ecuador, and on the Atlantic side from the vicinity of Austara, in the northeast coast of

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

the Atlantico Norte Department in Nicaragua to the Bolivar Department in Colombia. Regarding the southernmost distribution range on the Pacific coast, Pelliciera rhizophorae has been reported to occur in the Río Chone estuary in the province of Manabi (Bodero, 1993; PMRC, 1993), and has been mapped even farther, to the south of the Guayaquil Gulf (West, 1977), in the very dry and dry forest life zones of southwestern Ecuador. Based on numerous field observations, and the absence of herbarium collections from those localities, it is herein stated that the southernmost distribution range of P. rhizophorae is the Muisne river estuary in the Province of Esmeraldas in northwestern Ecuador, and not southwards in the very dry and dry forest life zones at the Equatorial-Pacific mangroves. Pelliciera rhizophorae has a long fossil record, that proves it once occurred as far north as Chiapas (southwestern Mexico) on the Pacific and throughout the Caribbean region (Graham, 1977). As the climatic conditions deteriorated during the Pleistocene, Pelliciera became restricted to the wettest Neotropical coastal region in the Choco, being the sole survivor of Eocene mangroves (approximately, 50 million years ago; Rull, 1998). This instance is one of the best supports for the refugium theory (Gentry, 1982; Tomlinson, 1986). Ecology: Pelliciera rhizophorae usually inhabits water saturated and poorly aired hyposaline or brackish environments, on periodically shallowly inundated, slightly elevated soft-muddy soils at high tides; it occasionally inhabits sandy beaches. It occurs in the wet forest and rain forest life zones. Phenology: This species bears flowers and fruits throughout the year. Biological interactions: The flowers of Pelliciera rhizophorae segregate nectar at the inner base. They have been reported to be frequently visited by hummingbirds of the genus Amazilia (Prahl, 1984), which has been confirmed by local residents in El Viento (REMACAM), in northwestern Ecuador. However, because of the large size of the terminal white flowers and the pendulous arrangement of the elongated linear-oblong stamens, it cannot be discarded that P. rhizophorae could be also pollinated by bats. This assumption needs to be verified by field observations. Outside of our area, in the mangroves of the Peninsula de Osa, Costa Rica, the inner base of P. rhizophorae flowers has been observed to be visited by domestic bees (Cornejo, field obs.). Finally, the nectar of the flowers is occasionally enjoyed by children, who call it “agua dulce” (sweet water). The function of the marginal foliar glands is not firmly known, but the presence of salty fluid droplets along the leaf margin suggests that they may operate as salt secretory glands (Luna and Ochoterena, 2004). Uses: The wood is solid and incorruptible, used for construction of floor frames known as “cuerdas”, or columns called “puntales” (Spanish) (Acosta-Solís, 1944). IUCN Red list category: Due to factors such as a narrow and fragmented distribution range of less than 2,000 km2 and a loss of habitat, it is agreed with the previous categorization of Pelliciera rhizophorae as Vulnerable (VU B2abiii) (Polidoro et al., 2010). It has been recommended to enrich the local genetic diversity of this species by artificial seed translocation (Castillo-Cárdenas and Toro-Perea, 2012). Discussions: Pelliciera rhizophorae var. rhizophorae presents “floribus candidis”, that means flowers of a pure white color, as originally described by Triana & Planchon (1862) based on the type collected in

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1852 by Triana, in Buenaventura, Valle del Cauca Department, western Colombia. The P. rhizophorae var. benthamii Triana & Planchon type collected by Sutton Hayes on the Mesoamerican Pacific coast, probably in Panama, presents “sepalis primum albis, demum sicut petala roseis”, sepals that are at first white and later become rose-colored like the petals. Subsequent collections gathered from the Pacific coast of eastern Panama document a P. rhizophorae that has flowers with rose to bright-red floral bracts and pale to medium pink petals (e.g., Gentry 3961, Ducke 10948, MO); however, these have not been seen in the field by the author, and they don’t appear to have been recorded on the collection labels of this species in our area. It is suspected that P. rhizophorae var. benthamii might be a valid variety based on the flower colors (as originally proposed by Triana and Planchon, 1862), but as such differences remain undetected in dried herbarium specimens the author is reluctant to place P. rhizophorae var. benthamii in the synonymy of P. rhizophorae until more field and molecular work is done. Acosta-Solís (1966) defined this species merely as an “associated tree” in the mangroves, but its narrow pattern of distribution, combined with its highly specialized adaptations to inhabit estuarine environments, such as fluted buttresses at the trunk base and water dispersed fruits/seeds, indicate that Pelliciera rhizophorae is a minor mangrove. Etymology: The genus Pelliciera was dedicated to the naturalist Bishop Pellicier of Montpellier. The genus has been spelled Pelliceria, a lapsus calamus of Triana and Planchon (Kobuski, 1951). The specific epithet refers to the common association of this species with Rhizophora. VISCACEAE (Christmas Mistletoe Family) Leafy to squamate plants; lacking epicortal roots, and implanted by means of primary haustoria only, these often expanding by means of endophytic strands. Stems often articulated at nodes, brittle. Leaves decussate, entire. Monoecious or dioecious. Flowers monochlamydeous, unisexual, 2–4-parted, at least females without aborted organs of opposite sex; male flowers with perianth segments around small central disk prominence; anthers isomorphic, sessile. 222. Phoradendron chrysocladon A. Gray, U.S. Expl. Exped., Phan. 743–744 (1854). Syn.:

Viscum flavens Sw. Viscum macrophyllum Macfad. Phoradendron flavens (Sw.) Griseb. Phoradendron flavum Eichler Phoradendron reticulatum Urb. Phoradendron urbanianum Ule Phoradendron lindavianum Pacz. Phoradendron quinquenervium Krause Phoradendron knoopii Trel. Phoradendron membranaceum Trel. Phoradendron pachyphyllum Trel.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Phoradendron supravenulosum Trel. Phoradendron trisulcatum Trel. Phoradendron glyptoneuron Diels Parasitic shrubs, rather large, yellowish-green. Stems terete to slightly 2-keeled. Pairs of leaves rather widely spaced, to 12 cm or more apart; 3–15 mm above each pair is one pair of acute intercalary cataphylls. Blades broadly-lanceolate, to 15 x 9 cm, with usually 5 conspicuously palmate veins; petioles stout, ca. 5 mm long. Monoecious, the sex distribution on spikes extremely variable. Spikes golden yellow, without sterile cataphylls; peduncle to 1 cm long, followed by 3–5 fertile internodes becoming to 5 cm long in fruit; flower series usually very regular, 3 series above each bract with often 12 or more flowers per series; fertile internodes commonly regularly tapering at base; rarely 5 flower series above each bract, producing two ± confluent flowers areas ensheathing axis. Fruit an ovoid to spherical berry, ca. 3 mm long.

ECUADOR. E: Játiva & Epling 750 (S, US). Distribution: Southern Mexico to Bolivia. Ecology: In wet forests and rainforests. Biological interactions: No information is recorded, but the small berries are most likely are eaten by birds, and the seeds endozoochoreously dispersed. IUCN Red list category: Least Concern (LC). Etymology: The generic epithet is formed by the Greek words phoros (bearing) and dendron (tree), referring to the characteristic parasitic habit of this taxon, which grows on trees. The specific epithet is formed by the Greek prefix chryso (golden) and the Greek word clados (branch), which refers to the golden-yellow color of the branches. Note: This species has been treated as P. flavens (Sw.) Griseb. (Kuijt, 1986).

Table 8. Number of plant species (incl. expected) per country in the mangroves on the Pacific coast of South America. Country Number of species

Colombia 184

Ecuador 179

Peru 17

Xavier Cornejo, editor - 2014

251

TABLE 9. IUCN CATEGORY AND HABITS OF MANGROVE SPECIES AND ASSOCIATE FLORA ON THE PACIFIC COAST OF SOUTH AMERICA (Plant habits: T: tree. S: shrub. H: herb. L: liana. V: vine. E: epiphyte. s: strangler. P: parasite.) Family

Scientific Name

Asple Asple Blechn Dryopt Dryopt Hymen Hymen Hymen Lomar Polyp Polyp Polyp Polyp Polyp Polyp Pteri Pteri Pteri Zamia Acanth Aizo Amaran Amaran Amaran Amaryl Anno Apoc Arac Arac Arac Arac Arac Arac Arac Arac Arac Arac Arac Arac Arac Arac Arac

Asplenium serratum Asplenium sp. Blechnum serrulatum Elaphoglossum latifolium Elaphoglossum productum Hymenophyllum polyanthus Trichomanes ankersii Trichomanes crinitum Nephrolepis biserrata Cochlidium serrulatum Microgramma percussa Microgramma reptans Niphidium crassifolium Pleopeltis bombycina Serpocaulon triseriale Acrostichum aureum Acrostichum danaeifolium Vittaria lineata Zamia roezlii Avicennia germinans Sesuvium portulacastrum Chamissoa altissima Salicornia fruticosa Saueda sp. Crinum kunthianum Annona glabra Rhabdadenia biflora Anthurium asplundii Anthurium cordobense Anthurium friedrichsthalii Anthurium guayaquilense Anthurium hacumense Anthurium incomptum Anthurium littorale Anthurium obtusum Anthurium oxyanthum Anthurium paludosum Anthurium trilobum Anthurium verrucosum Monstera pinnatipartita Philodendron chrysocarpum Philodendron musifolium

Habit

IUCN Category

Reference

E E H E E E E E H E E E E E E H H E T/S T/S H V H H H T L E E E E E E E E E E E E E E E

LC DD LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC VU B1abiii LC LC LC LC DD LC LC LC LC NT LC NT LC LC LC LC NT LC LC LC LC LC NT

This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication Polidoro et al., 2010 Polidoro et al., 2010 This publication Galeano et al., 2005 Polidoro et al., 2010 This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication This publication Croat et al., 2008 This publication

252

Family Arac Arac Arac Arac Arec Arec Aster Aster Batac Bigno Bigno Bigno Bigno Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Brom Cact Cact Cact Chrys Chrys

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Scientific Name Philodendron sparreorum Philodendron tripartitum Stenospermation angustifolium Stenospermation robustum Cocos nucifera Euterpe oleracea Tuberostylis axillaris Tuberostylis rhizophorae Batis maritima Amphitecna latifolia Anemopaegma chrysanthum Phryganocydia phellosperma Tabebuia palustris Aechmea angustifolia Aechmea dactylina Aechmea nudicaulis Aechmea pyramidalis Billbergia macrolepis Catopsis sessiliflora Guzmania hitchcockiana Guzmania monostachia Guzmania musaica Guzmania scherzeriana Pitcairnia heterophylla Racinaea multiflora Tillandsia barclayana var. minor Tillandsia bulbosa Tillandsia complanata Tillandsia disticha Tillandsia dyeriana Tillandsia flagellata Tillandsia latifolia var. divaricata Tillandsia narthecioides Tillandsia subulifera Tillandsia triglochinoides Tillandsia usneoides Tillandsia venusta Werauhia gladioliflora Werauhia cf. kupperiana Werauhia ringens Werauhia sanguinolenta Epiphyllum rubrocoronatum Hylocereus polyrhizus Rhipsalis micrantha Chrysobalanus icaco Hirtella carbonaria

Habit

IUCN Category

Reference

E E E E T T T/S/L E H T L L T/S E E E E E E E E E E E E E E E E E E E E E E E E E E E E E E E T T

NT LC LC LC LC LC VU B2abiii VU B2abiii LC LC LC VUB2abiii VU B2abiii LC LC LC LC LC LC NT LC LC NT LC LC NT LC LC LC CR A4c NT LC LC LC VU B1abiii LC LC LC DD LC LC LC LC LC LC VU B2abiii

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Xavier Cornejo, editor - 2014

Family Clusi Clusi Comb Comb Comb Cycla Cycla Cype Cype Cype Cype Cype Cype Cype Cype Cype Cype Erica Erica Erica Erica Erica Erica Erica Faba Faba Faba Faba Faba Gesn Gesn Hippo Lora Lora Lora Lora Lora Lora Lythr Malph Malph Malph Malph Malph Malph

253

Scientific Name

Habit

IUCN Category

Reference

Clusia fructiangusta Clusia leptanthera Conocarpus erectus Laguncularia racemosa var. racemosa Laguncularia racemosa var. glabriflora Sphaeradenia acutitepala Sphaeradenia killipii Cyperus ligularis Cyperus odoratus Eleocharis elegans Fimbristylis dichotoma Fimbristylis spadicea Fuirena umbellata Rhynchospora corymbosa Schoenoplectus californicus Scleria eggersiana Scleria melaleuca Anthopterus wardii Cavendishia callista Cavendishia urophylla Macleania smithiana Psammisia occidentalis Sphyrospermum buxifolium Thibaudia pachypoda Dalbergia brownei Entada polystachya Mora oleifera Muellera chocoensis Pterocarpus officinalis Codonanthe crassifolia Drymonia serrulata Hippocratea volubilis Oryctanthus florulentus Oryctanthus occidentalis Phthirusa pyrifolia Phthirusa stelis Psittacanthus divaricatus Struthanthus orbicularis Crenea patentinervis Banisteriopsis elegans Banisteriopsis martiniana subsp. subenervia Bunchosia plowmanii Heteropterys leona Hiraea brachyptera Hiraea fagifolia

E E T/S T/S T/S E E H H H H H H H H H H E E E E E E E L L T T T E E L P P P P P P H L L

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T/S L L L

NT LC VU B2abiii LC

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254

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Family

Scientific Name

Habit

IUCN Category

Reference

Malph Malph Malv Malv Malv Marcgr Marcgr Marcgr Marcgr Melast Melast Melast Melast Morac Morac Morac Morac Morac Myrsi Nyct Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch

Stigmaphyllon ellipticum Tetrapterys subaptera Pavonia kearneyi Pavonia rhizophorae Talipariti tiliaceum var. pernambucense Pseudosarcopera diaz-piedrahitae Schwartzia chocoensis Souroubea guianensis Souroubea intermedia Blakea glabrescens Conostegia polyandra Miconia reducens Tococa guianensis Ficus brevibracteata Ficus jacobii subsp. mantana Ficus maxima Ficus nymphaeifolia Ficus obtusifolia Ardisia granatensis Cryptocarpus pyriformis Brassavola grandiflora Camaridium atrovinaceum Campylocentrum ecuadorense Catasetum macroglossum Caularthron bilamellatum Dichaea panamensis Dichaea trulla Dimerandra rimbachii Elleanthus discolor Epidendrum bracteolatum Epidendrum buenaventurae Epidendrum cocornocturnum Epidendrum littorale Epidendrum luckei Epidendrum macroophorum Epidendrum microphyllum Epidendrum pseudonocturnum Epidendrum rigidum Epidendrum sculptum Epidendrum stangeanum Epidendrum sp. Maxillaria hennisiana Notylia replicata Ornithidium fulgens Ornithocephalus bryostachys Platystele cornejoi

L L S/H S T/S E E E E E S T/S T/S s s s s s T/S L E E E E E E E E H E E E E E E E E E E E E E E E E E

LC VU B2abiii VU B2abiii VU B2abiii LC NT NT LC LC LC LC LC LC LC NT LC LC LC NT LC LC NT NT NT LC LC LC NT LC NT LC NT LC LC NT LC DD LC DD DD DD NT NT LC LC EN B1abiii

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Xavier Cornejo, editor - 2014

Family Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Orch Piper Piper Piper Piper Piper Piper Piper Piper Piper Piper Piper Piper Poac Poac Poac Poac Pont Pont Potam Rhizo Rhizo Rhizo Rhizo Rubia Rubia Rubia Rubia Rubia Rubia Scroph Tetram Visca

255

Scientific Name Polystachya concreta Prosthechea fragrans Prosthechea pseudopygmaea Sarcinula sp. Scaphyglottis crurigera Scaphyglottis graminifolia Scaphyglottis prolifera Scaphyglottis sp. nov. Sobralia madisonii Sobralia rhizophorae Soterosanthus shepheardii Trichosalpinx orbicularis Trigonidium seemannii Peperomia blephariphylla Peperomia ciliaris Peperomia elongata Peperomia emarginella Peperomia gorgonillana Peperomia jamesoniana Peperomia obovalis Peperomia obtusifolia Peperomia pernambucensis Peperomia portobellensis Peperomia rotundifolia Peperomia serpens Jouvea straminea Paspalum vaginatum Uniola pittieri Urochloa mutica Eichhornia azurea Eichhornia crassipes Ruppia maritima Cassipourea killipii Rhizophora mangle Rhizophora racemosa Rhizophora x harrisonii Chiococca alba Cosmibuena macrocarpa Hillia maxonii Notopleura epiphytica Rustia occidentalis Tobagoa maleolens Schlegelia darienensis Pelliciera rhizophorae Phoradendron chrysocladon

Habit

IUCN Category

Reference

E E E E E E E E E E E E E E E E E E E E E E E E E H H H H H H H T T/S T/S T/S L T S E T/S H E T P

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256

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 1: A. Blechnum serrulatum. B. Elaphoglossum latifolium. C. E. productum. D. Hymenophyllum polyanthos.

Xavier Cornejo, editor - 2014

Plate 2: A. Nephrolepis biserrata. B. Cochlidium serrulatum. C. Microgramma percussa. D. M. reptans.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 3: A. Niphidium crassifolium. B. Pleopeltis bombycina. C. Serpocaulon triseriale. D. Vittaria lineata.

Xavier Cornejo, editor - 2014

Plate 4: Aâ&#x20AC;&#x201C;B. Acrostichum aureum. Câ&#x20AC;&#x201C;D. A. danaeifolium.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 5: Aâ&#x20AC;&#x201C;E. Avicennia germinans. A. Inflorescences. B. Infructescences. C. Terminal leafy branch. D. Roots and pneumatophores. E. Best developed individual of this species found.

Xavier Cornejo, editor - 2014

Plate 6: A. Sesuvium portulacastrum. B. Salicornia fruticosa. C. Saueda sp. D. Crinum kunthianum. Photo A courtesy of S. Ferreiros; photos B, C courtesy of S. Torres-Bรกez

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262

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 7: A. Annona glabra. B. Rhabdadenia biflora. C. Anthurium asplundii. D. A. incomptum.

Xavier Cornejo, editor - 2014

Plate 8: A. Anthurium littorale. B. A. obtusum. C. Philodendron musifolium. D. P. sparreorum.

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264

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 9: A. Cocos nucifera. B. Euterpe oleracea. C. Tuberostylis rhizophorae. D. Batis maritima. Photo D courtesy of S. Torres-Bรกez.

Xavier Cornejo, editor - 2014

265

Plate 10: A. Amphitecna latifolia. B. Anemopaegma chrysanthum. C. Phryganocydia phellosperma. D. Tabebuia palustris.

266

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 11: A. Aechmea angustifolia. B. A. dactylina. C. A. nudicaulis. D. A. pyramidalis.

Xavier Cornejo, editor - 2014

Plate 12: A. Billbergia macrolepis. B. Catopsis sessiliflora. C. Guzmania hitchcockiana. D. G. monostachia.

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268

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 13: A. Guzmania musaica. B. Racinaea multiflora var. multiflora. C. R. multiflora var. decipiens. D. Tillandsia barclayana var. minor. Photo A courtesy of P. Nelson.

Xavier Cornejo, editor - 2014

Plate 14: A. Tillandsia bulbosa. B. T. complanata. C. T. disticha. D. T. dyeriana. Photo D courtesy of P. Nelson.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 15: A. Tillandsia flagellata. B. T. latifolia var. divaricata. C. T. narthecioides. D. T. subulifera. Photo A courtesy of P. Nelson.

Xavier Cornejo, editor - 2014

Plate 16: A. Tillandsia triglochinoides. B. T. usneoides. C. T. venusta. D. Werauhia gladioliflora. Photo D courtesy of P. Nelson.

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272

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 17: A. Werauhia cf. kupperiana. B. W. ringens. C. W. sanguinolenta. D. Epiphyllum rubrocoronatum.

Xavier Cornejo, editor - 2014

Plate 18: A. Chrysobalanus icaco. B. Hirtella carbonaria. C. Clusia fructiangusta. D. C. leptanthera.

273

274

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 19: Aâ&#x20AC;&#x201C;E. Conocarpus erectus. A. Inflorescences. B. Infructescences. C. Terminal leafy branch. D. Leaf blade beneath. E. Developed trunk.

Xavier Cornejo, editor - 2014

275

Plate 20: Aâ&#x20AC;&#x201C;E. Laguncularia racemosa. A, C, E, F. L. racemosa var. racemosa: A. Inflorescence. C. Infructescence. E. Terminal young leaves. F. Roots and lower trunk. B, D. L. racemosa var. glabriflora, from the type locality: B. Inflorescence. D. Infructescence.

276

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 21: A. Sphaeradenia killipii. B. Cyperus ligularis. C. C.odoratus. D. Anthopterus wardii. Photo D courtesy of P. Pedraza-PeÃ±aloza.

Xavier Cornejo, editor - 2014

Plate 22: A. Cavendishia callista. B. C. urophylla. C. Macleania smithiana. D. Sphyrospermum buxifolium. Photos A, B courtesy of J. Luteyn; photo D courtesy of P. Pedraza-PeĂąaloza.

277

278

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 23: A. Thibaudia pachypoda. B. Dalbergia brownei. C. Entada polystachya. D. Muellera chocoensis. Photo A courtesy of J. Luteyn.

Xavier Cornejo, editor - 2014

Plate 24: Aâ&#x20AC;&#x201C;D. Mora oleifera. A. Inflorescence. B. Fruit. C. Paripinnate leaf blade, above. D. Buttresses and lower trunk of a well-developed individual.

279

280

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 25: Aâ&#x20AC;&#x201C;D. Pterocarpus officinalis. A. Partial view of inflorescence. B. Imparipinnate leaf blade, above. C. Bark. D. Buttresses and lower trunk.

Xavier Cornejo, editor - 2014

Plate 26: A. Codonanthe crassifolia. B. Psittacanthus divaricatus. C. Struthanthus orbicularis. D. Crenea patentinervis.

281

282

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 27: A. Banisteriopsis martiniana subsp. subenervia. B. Bunchosia plowmanii. C. Stigmaphyllon ellipticum. D. Tetrapterys subaptera.

Xavier Cornejo, editor - 2014

Plate 28: A. Pavonia rhizophorae. B. Talipariti tiliaceum var. pernambucense. C. Blakea glabrescens. D. Conostegia polyandra.

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284

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 29: A. Miconia reducens. B. Ficus nymphaeifolia. C. F. obtusifolia. D. Cryptocarpus pyriformis.

Xavier Cornejo, editor - 2014

Plate 30: A. Camaridium atrovinaceum. B. Catasetum macroglossum. C. Caularthron bilamellatum. D. Dichaea panamensis.

285

286

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 31: A. Dichaea trulla. B. Dimerandra rimbachii. C. Epidendrum buenaventurae. D. E. cocornocturnum. Photo C courtesy of E. Hรกgsater.

Xavier Cornejo, editor - 2014

Plate 32: A. Epidendrum luckei. B. E. macroophorum. C. E. pseudonocturnum. D. E. rigidum. Photo B courtesy of E. Hรกgsater.

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288

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 33: A. Epidendrum sculptum. B. E. stangeanum. C. Epidendrum sp. D. Maxillaria hennisiana. Photo B courtesy of McCullough.

Xavier Cornejo, editor - 2014

Plate 34: A. Notylia replicata. B. Ornithidium fulgens. C. Platystele cornejoi. D. Polystachya concreta. Photo A courtesy of N. Pyrooz.

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290

PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 35: A. Sarcinula sp. B. Scaphyglottis crurigera. C. Scaphyglottis prolifera. D. Scaphyglottis sp. nov.

Xavier Cornejo, editor - 2014

Plate 36: A. Sobralia madisonii. B. Sobralia rhizophorae. C. Soterosanthus shepheardii. D. Trichosalpinx orbicularis.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 37: A. Peperomia obovalis. B. Peperomia obtusifolia. C. Peperomia pernambucensis. D. Peperomia portobellensis.

Xavier Cornejo, editor - 2014

Plate 38: A. Uniola pittieri. B. Eichhornia crassipes. C. E. azurea. D. Cassipourea killipii.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 39: Aâ&#x20AC;&#x201C;F. Rhizophora mangle. A. Inflorescences bearing flower buds. B. Flower. C. Fruit and developed hypocotyl. D. Stipules. E. Leaf blades, beneath. F. Prop roots and lower trunk of a well-developed individual.

Xavier Cornejo, editor - 2014

Plate 40: Aâ&#x20AC;&#x201C;F. Rhizophora racemosa. A. Terminal leaves and conical stipules. B. Stipules and new leaves C. Inflorescence. D. Fruits and developed hypocotyls. E. Mature trees. F. Adventitious roots.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 41: Holotype of Rhizophora racemosa (GOET). B. Chiococca alba. C. Cosmibuena macrocarpa. D. Rustia occidentalis. Photo A courtesy of Jochen Heinrichs.

Xavier Cornejo, editor - 2014

297

Plate 42: Aâ&#x20AC;&#x201C;C. Pelliciera rhizophorae. A. Flower. B. Fruit (left) and seed with a removed cotyledon displaying embryo (right). C. Buttresses and lower trunks during low tide, the dark-brown color of buttresses indicates the local intertidal range.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

Plate 43: A. Arrangement of leaves: 1. Alternate. 2. Opposite. 3. Verticillate. B. Types of leaves: 4. Simple. 5. Trifoliolate. 6. Paripinnate. 7. Imparipinnate. C. Outlines of leaves: 8. Elliptic. 9. Ovate. 10. Lanceolate. 11. Oblong. 12. Linear. 13. Oblanceolate. 14. Obovate. 15. Cordate. 16. Orbicular.

Xavier Cornejo, editor - 2014

299

LITERATURE CITED

Acosta-Solís, M. 1944. Nuevas contribuciones al conocimiento de La Provincia de Esmeraldas, Tomo I. Publicaciones científicas MAS. Quito, Ecuador. Acosta-Solís, M. 1959. Los Manglares el Ecuador. Contribución Nº 29 del Instituto Ecuatoriano de Ciencias Naturales. Quito, Ecuador. Acosta-Solís, M. 1966. Las Divisiones Fitogeográficas y las Formaciones Geobotánicas del Ecuador. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 12: 401–117. ACTMANG. 1995. Landform and vegetation around the investigated area. Vegetation, topography and geological profile across the Majagual mangrove forest. Tokyo, Japan. Alvarez-León, A. 1993. Mangrove Ecosystems of Colombia. In: L. D. Lacerda (ed.), Conservation and Sustainable utilization of mangrove forest in Latin America and Africa Regions. Part I–Latin America. International Society for Mangrove Ecosystems. Yokohama, Japan. Anderson, C. 2001. The identity of two water-dispersed species of Heteropterys (Malpighiaceae): H. leona and H. platyptera. Contributions from the University of Michigan Herbarium 23: 35–47. Anderson, W. R. 1987. Notes on Neotropical Malpighiaceae II. Contributions from the University of Michigan Herbarium 16: 55–108. Arbeláez, A. L. 1996. La tribu Pteridae (Pteridaceae). Flora de Colombia 18: 1–104. Austin, D. F. 2004. Florida Ethnobotany. CRC Press. Boca Raton, Florida. Barbosa, C. 1988. Rhizophoraceae en el Herbario Nacional Colombiano. Trianaea (Acta Científica y Tecnológica Inderena) 2: 251–253. Barrionuevo, R. and R. Marcial. 2006. Ecología trófica de la fauna acuática en el Manglar de San Pedro - Sechura. Universalia 11: 44–56. Bentham, G. and J. D. Hooker. 1844. The botany of the voyage of the H.M.S. Sulfur, under the comand of Captain Sir Edward Belcher during the years 1836–42. Published under the authority of the Lords Commissioners of the Admiralty. Edited and superintended by Richard Brinsley Hinds, attached to the expedition. The botanical descriptions by George Bentham. Smith, Elder & Co. London, United Kingdom. Bernal, H. Y. and J. E. Correa. 1989. Especies Vegetales Promisorias de los países del convenio Andrés Bello, tomo II. Bogotá, Colombia. Bernal, H. Y. and J. E. Correa. 1990. Especies Vegetales Promisorias de los países del convenio Andrés Bello, tomo IV. Bogotá, Colombia. Bergoeing, J. P. 1998. Geomorfología de Costa Rica. San José, Instituto Geográfico Nacional. 409 p. Blake, S. F. 1943. Ten new species of Asteraceae. Journal of the Washington Academy of Sciences 33: 265–272. Bodero, A. 1993. Mangrove Ecosystems of Ecuador. In: L. D. Lacerda (ed.), Conservation and Sustainable utilization of mangrove forest in Latin America and Africa Regions. Part I–Latin America. International Society for Mangrove Ecosystems. Yokohama, Japan. Bonifaz, C. 1988. Vegetación de un manglar hiposalino de la Provincia del Guayas. Facultad de Ciencias Naturales, Departamento de Publicaciones, Universidad de Guayaquil. Guayaquil, Ecuador. Bravo, M. 2010. Interpretación del Estudio Multitemporal (Clirsen 1969–2006) de las coberturas de manglar, camaroneras y áreas salinas en la franja costera del Ecuador continental. Tesis de grado para la obtención del título de magister en Ciencias con énfasis en manejo sustentable en recursos bioacuáticos y medio Ambiente. Universidad de Guayaquil, Facultad de Ciencias Naturales, Ecuador. 56 p. Breteler, F. J. 1969. The Atlantic species of Rhizophora. Acta Botanica Neerlandica 18: 434–441. Breteler, F. J. 1977. America’s Pacific species of Rhizophora. Acta Botanica Neerlandica 26: 225–230.

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Plumier, C. 1703. Nova plantarum Americanarum genera. J. Boudot, Paris. PMRC. 1993. Plan de manejo de la ZEM Bahía-San Vicente-Canoa, 1ra ed. Guayaquil, Ecuador. Polidoro, B. A., K. E. Carpenter, L. Collins, N. C. Duke, A. M. Ellison, J. C. Ellison, E. J. Farnsworth, E. S. Fernando, K. Kathiresan, N. E. Koedam, S. R. Livingstone, T. Miyagi, G. E. Moore, V. N. Nam, J. E. Ong, J. H. Primavera, S. G. Salmo, J. C. Sanciangco, S. Sukardjo, Y. Wang and J. W. Hong Yong. 2010. The Loss of Species: Mangrove Extinction Risk and Geographic Areas of Global Concern. PLoS ONE 5(4): e10095. doi:10.1371/journal.pone.0010095 Pool, A. 2001. Pellicieraceae. Flora de Nicaragua, Angiospermas (Pandanaceae-Zygophyllaceae). Monographs in Systematic Botany from the Missouri Botanical Garden 85: 1924. Prahl, H. Von. 1984. Notas Sistemáticas de las Diferentes Especies de Mangle del Pacífico Colombiano. Cespedesia 13: 222–238. Prahl, H. Von. 1989. Manglares de Colombia. Villegas Editores, Bogotá, Colombia. 207 p. Prahl H. Von, J. R. Cantera, and R. Contreras. 1990. Manglares y hombres del Pacífico colombiano. FEN, Colombia/ COLCIENCIAS y Editorial Presencia, Bogotá, Colombia. 193 p. Prance, G. T. 1982. Forest Refuges: Evidence from Woody Angiosperms. In: G. T. Prance, Biological Diversification in the Tropics. Columbia University Press, New York. Prance, G. T. 1998. Chrysobalanaceae. Flora of the Venezuelan Guayana 4: 202–246. Missouri Botanical Garden Press, St. Louis. Prance, G. T., M. Freitas da Silva, B. W. Albuquerque, I. de J. da Silva Araújo, L. M. Medeiros Carreira, M. M. Nogueira Braga, M. Macedo, P. N. da Conceição, P. L. Braga Lisboa, P. I. Braga, R. C. Lobato Lisboa and R.C. Queiroz Vilhena. 1975. Revisão Taxonômica das espécies amazônicas de Rhizophoraceae. Acta Amazonica 5: 5–22. Presl, K. B. 1825–1835. Reliquiae Haenkeanae, seu descriptiones et icones plantarum, quas in America Meridionali et Boreali, in insulis Philippines et Marianis collegit Thaddaeus Haenke (reprint 1973). Amsterdam. Pupulin, F. 2007. Contributions toward a reassessment of Costa Rican Zygopetalinae (Orchidaceae). 3. A systematic revision of Dichaea in Costa Rica. Harvard Papers in Botany 12: 15–153. Restrepo J. D. and J. R. Cantera (eds.). 1995. Delta del Río San Juan-Bahías de Málaga y Buenaventura, Pacífico colombiano, Tomo I: 337 p; Tomo II: 344 p. Rodríguez, W., R. Marcial and R. Barrionuevo. 1997. Biodiversidad del Manglar de San Pedro entre mayo 1995–abril 1997. Sechura. UNIVERSALIA. Rull, V. 1998. Evolución de los manglares neotropicales: La crisis del Eoceno. Interciencia 23: 355–362. Saenger, P., E. J. Hegerl, and J. D. S. Davie (eds.). 1983. Global status of mangrove ecosystems. The Environmentalist 3 (Supplement): 1–88. Salas, J. 2010. Diversidad y Ecología de los Quirópteros como indicadores del estado de conservación de la reserva de producción de fauna “Manglares El Salado”. Tesis de grado para la obtención del título de magister en Ciencias con énfasis en manejo sustentable en recursos bioacuáticos y medio Ambiente. Universidad de Guayaquil, Facultad de Ciencias Naturales, Ecuador. 80 p. Sánchez-Páez, H., R. Álvarez, O. A. Guevara, A. Zamora, H. Rodríguez, and H. E. Bravo. 1997. Diagnóstico y zonificación preliminar de los manglares del Pacífico de Colombia. Ministerio del Medio Ambiente, Bogotá, Colombia. 343 p. Sánchez-Páez, H., G. A. Ulloa-Delgado, H. A. Tavera-Escobar, and W. O. Gil-Torres. 2003. Plan de manejo integral de los manglares de la zona de uso sostenible sector estuarino de la bahía de Cispatá departamento de Córdoba, Convenio No. 063 CVSCONIF. Ministerio de Ambiente, Vivienda y Desarrollo Territorial, CVS, CONIF, Montería, Córdoba, Colombia. 312 p. Sierra, S. 2001. Estudio de la relación entre los cambios en la zonación y estructura del ecosistema de manglar y la actividad antrópica que se ha ejercido sobre él, en la ensenada de Virudó (Bajo Baudó, Chocó), Pacífico colombiano. Tesis de pregrado. Universidad del Valle, Facultad de Ciencias, Cali, Colombia. 112 p. Sigrist, M. R. and M. Sazima. 2004. Pollination and reproductive biology of twelve species of Neotropical Malpighiaceae: Stigma morphology and its implications for the breeding system. Annals of Botany 94: 33–41.

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Smith A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, and P.G. Wolf. 2006. A classification for extant ferns. Taxon 55(3): 705–731. Souza, M. and S. Sotuyo. 2012. El género Muellera L.f. en Mesoamérica y norte de Sudamérica. Acta Botánica Mexicana 100: 15–40. Stace, C. A. 2007. Combretaceae. Flora of Ecuador 81: 1–63. Stace, C. A. 2010. Combretaceae. Flora Neotropica Monograph 107: 1–369. Stearn, W. T. 1995. Botanical Latin: History, Grammar, Syntax, Terminology and Vocabulary (4th ed.). Timber Press. Portland, Oregon. Steiner, K. E. 1985. The role of nectar and oil in the pollination of Drymonia serrulata (Gesneriaceae) by Epicharis Bees (Anthophoridae) in Panama. Biotropica 17: 217–229. Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 12, Jul 2012 [and more or less continuously updated since]. http://www.mobot.org/MOBOT/research/APweb/ Stevenson, D. W. 2004. Cycads of Colombia. The Botanical Review 70: 194–234. Stothert, K. E. 2011. Coastal resources and the Early Holocene Las Vegas Adaptation of Ecuador. In: Bicho, N. F. et al. (eds.), Trekking the shore: Changing coastlines and the Antiquity of Coastal Settlement: Interdisciplinary contributions to Archaeology. New York, 355–382. Takahashi, J. 2002. Los manglares del Perú, Revisión Histórica 1942–2002. Proyecto FANPE: INRENA–GTZ. Tansley, A. and F. Fritsch. 1905. The flora of Ceylon littoral. II. New Phytologist 4: 27–55. Tomlimson, P. B. 1986. The Botany of Mangroves. Cambridge University Press, Cambridge, United Kingdom. Triana, J. and J. E. Planchon. 1862. Prodromus Florae Novo-Granatensis ou Enumeration des plantes de la Nouvelle– Grenade, avec description des espècies nouvelles. Annales des Sciences Naturelles, Botanique, Série 4, 17: 319–382. Valverde, F. de M. 1988. Perspectiva Botánica de los Manglares de la Provincia del Guayas. Acta Científica Ecuatoriana Vol. 1. Ecuador. Villamil, C. A. 2010. Evaluación de ecosistemas de manglar con diferente cobertura en cuanto a los procesos de retención, absorción y acumulación de metales pesados (Cr, Cd, Pb, Zn y Cu). Tesis de grado para la obtención del título de magister en Ciencias en Biología–línea Biología Marina. Universidad Nacional de Colombia. 79 p. West, R. C. 1956. Mangrove swamps of the Pacific coast of Colombia. Annals of the Association of American Geographers 46: 98–121. West, R. C. 1977. Tidal salt-marsh and mangal formations of middle and south America. In: Chapman, V. J. (ed.), Ecosystems of the World 1. Wet coastal ecosystems. Amsterdam-Oxford-NewYork, 193–212. Wolf, T. 1892. Geografía y Geología del Ecuador. Brockhaus, Leipzig. [Reprinted, 1975, Casa de la Cultura Ecuatoriana, Quito, Ecuador]. Zambrano, C. H. and D. J. Rubiano. 1996. Mapas del bosque de manglar de la costa pacífica colombiana años 1969, 1996 y multitemporal. Ministerio del Medio Ambiente, Bogotá, Colombia. 36 p.

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INDEX TO NAMES OF PLANTS Scientific names and plate pages are written in italics, respectively. Vernacular names are indicated by (vern.). ACANTHACEAE: 77. Acanthus family: 77. achotillo (vern.): 173. Acrostichum aureum: 73, 259. Acrostichum danaeifolium: 75, 259. adorote (vern.): 159. Aechmea angustifolia: 112, 266. Aechmea dactylina: 113, 266. Aechmea nudicaulis: 114, 266. Aechmea pyramidalis: 114, 266. AIZOACEAE: 81. ajo (vern.): 233. alcornoque (vern.): 161. Amaranth family: 82. AMARANTHACEAE: 82. amargosa (vern.): 224. AMARYLLIDACEAE: 84. Amphitecna latifolia: 106, 265. Anemopaegma chrysanthum: 107, 265. Annona glabra: 85, 262. ANNONACEAE: 85. Anthopterus wardii: 153, 276. Anthurium asplundii: 90, 262. Anthurium cordobense: 91. Anthurium friedrichsthalii: 91. Anthurium guayaquilense: 92. Anthurium hacumense: 92. Anthurium incomptum: 93, 262. Anthurium littorale: 93, 263. Anthurium obtusum: 94, 263. Anthurium oxyanthum: 94. Anthurium paludosum: 95. Anthurium trilobum: 95. Anthurium verrucosum: 96. APOCYNACEAE: 86. ARACEAE: 88. Ardisia granatensis: 193. ARECACEAE: 100. Arum family: 88. ASPLENIACEAE : 63. Asplenium serratum: 63. Asplenium sp.: 64. Aster family: 103.

ASTERACEAE: 103. Avicennia bicolor: 78. Avicennia germinans: 43, 78, 260. Avicennia nitida: 78. Avicennia tonduzii: 78. Avicenniaceae: 77. bagata (vern.): 163. bambil (vern.): 102. bambudo (vern.): 164. bambulo (vern.): 164. Banisteriopsis elegans: 174. Banisteriopsis martiniana subsp. subenervia: 175, 282. barba (vern.): 127. barba de viejo (vern.): 127. Barbados Cherry family: 173. barbasco (vern.): 108, 163. barbasquillo (vern.): 163. BATACEAE: 104. Batis maritima: 105, 264. Bean family: 159. bejuco de agua (vern.): 160. bejuco de cerca (vern.): 87. bejuco gallinero (vern.): 107. BIGNONIACEAE: 105. Billbergia macrolepis: 115, 267. Blakea glabrescens: 186, 283. BLECHNACEAE: 64. Blechnum serrulatum: 65, 256. Blueberry family: 152. botoncillo (vern.): 139. Brassavola grandiflora: 198. BROMELIACEAE: 109. Bromeliad family: 109. Bunchosia plowmanii: 175, 282. buâ&#x20AC;&#x2122;machi (vern.): 190. buchon (vern.): 231. CACTACEAE: 131. Cactus family: 131. calabacillo (vern.): 106. Camaridium atrovinaceum: 199, 285. Campylocentrum ecuadorense: 199. carboncillo (vern.): 136.

carbonero (vern.): 136. Cassipourea killipii: 233, 293. Catasetum macroglossum: 200, 285. Catopsis sessiliflora: 116, 267. Caularthron bilamellatum: 201, 285. Cavendishia callista: 154, 277. Cavendishia palustris: 155. Cavendishia urophylla: 156, 277. Chamissoa altissima: 82. Chascarra: 74, 75. Chenopodiaceae: 84. chicharrĂłn (vern.): 74, 75. chichihua (vern.): 171. chigua (vern.): 76. chiguamacho (vern.): 74, 75. Chiococca alba: 242. chope (vern.): 194. Christmas Mistletoe family: 249. CHRYSOBALANACEAE: 134. Chrysobalanus icaco: 135, 273. Clusia fructiangusta: 137, 273. Clusia leptanthera: 138, 273. CLUSIACEAE: 137. Clustard-Apple family: 85. coca (vern.): 135. Cochlidium serrulatum: 70, 255. Cocoa-plum family: 134. Cocos nucifera: 101, 264. cocotero (vern.): 101. Codonanthe crassifolia: 165, 281. Coffee family: 241. COMBRETACEAE: 138. comedero (vern.): 79. Conocarpus erectus: 139, 274. Conocarpus erectus forma sericeous: 140. Conostegia polyandra: 187, 283. corrientoso (vern.): 95. cortadera (vern.): 150, 151, 152. Cosmibuena macrocarpa: 243, 296. Crenea patentinervis: 173, 281. Crinum kunthianum: 84, 261. Cryptocarpus pyriformis: 194, 284. CYCLANTHACEAE: 144.

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INDEX TO NAMES OF PLANTS (cont.) Scientific names and plate pages are written in italics, respectively. Vernacular names are indicated by (vern.). CYPERACEAE: 146. Cyperus ligularis: 147, 276. Cyperus odoratus: 147, 276. Daffodil family: 84. Dalbergia brownei: 159, 278. deshinchador (vern.): 95. Dichaea panamensis: 201, 285. Dichaea trulla: 202, 286. Dicotyledons: 47, 48. Dimerandra rimbachii: 203, 286. disciplina (vern.): 133. Dogbane family: 86. Drymonia serrulata: 166. DRYOPTERIDACEAE: 65. Eichhornia azurea: 230, 293. Eichhornia crassipes: 231, 293. Elaphoglossum latifolium: 66, 256. Elaphoglossum productum: 66, 256. Eleocharis elegans: 148. Eleocharis mutata: 148. Elleanthus discolor: 203. Entada polystachya: 160, 278. Epidendrum bracteolatum: 204. Epidendrum buenaventurae: 204, 286. Epidendrum cocornocturnum: 205, 286. Epidendrum littorale: 205. Epidendrum luckei: 206, 287. Epidendrum macroophorum: 206, 287. Epidendrum microphyllum: 207. Epidendrum pseudonocturnum: 207, 287. Epidendrum rigidum: 208, 287. Epidendrum sculptum: 208, 288. Epidendrum sp.: 209, 288. Epidendrum stangeanum: 209, 288. epiphytes: 40, 47, 48, 49. Epiphyllum rubrocoronatum: 132, 272. ERICACEAE: 152. Erythroxylum coca: 135. Euterpe oleracea: 102, 264. FABACEAE: 159. femu-piu-tapé (vern.): 156. ferns: 48, 63.

Ficus brevibracteata: 189. Ficus jacobii subsp. mantana: 190. Ficus maxima: 190. Ficus nymphaeifolia: 191, 284. Ficus obtusifolia: 192, 284. Figwort or Snapdragon family: 246. Filicopsida: 63. Filmy fern family: 67. Fimbristylis dichotoma: 149. Fimbristylis spadicea: 149. Flower mite: 235, 296. Four-o’clock family: 193. Fuirena umbellata: 150. gabazo (vern.): 127. garrapato (vern.): 136. garza (vern.): 87. GESNERIACEAE: 165. Gesneriad family: 165. Grass family: 226. guaca de mata (vern.): 95. guaco (vern.): 95. guanábana de pozo (vern.): 85. guanabanillo (vern.): 85. guaral (vern.): 99. Guzmania hitchcockiana: 116, 267. Guzmania monostachia: 117, 267. Guzmania musaica: 118, 268. Guzmania scherzeriana: 118. Gymnosperms: 47, 48, 76. Hattena rhizophorae: 235, 296. herbs: 39, 47, 48, 49. Heteropterys leona: 176. Hibiscus tiliaceus: 181. hierba de pajarito (vern.): 171. Hillia maxonii: 244. Hippocratea family: 166. Hippocratea volubilis: 167. HIPPOCRATEACEAE: 166. Hiraea brachyptera: 177. Hiraea fagifolia: 177. Hirtella carbonaria: 136, 273. Hylocereus polyrhizus: 133.

HYMENOPHYLLACEAE: 67. Hymenophyllum polyanthos: 67, 256. icacales: 135. icaco (vern.): 135. iguanero (vern.): 79. Indian Almond family: 138. jabalí (vern.): 139. jelí (vern.): 139. jelí de semilla (vern.): 139. jelí salado (vern.): 79, 139. jelicales: 140. Jouvea straminea: 227. Laguncularia racemosa: 43, 141, 275. Laguncularia racemosa var. glabriflora: 143, 275. Laguncularia racemosa var. racemosa: 142, 275. lechera (vern.): 87. lecho de mar (vern.): 87. lechuga (vern.): 230, 231. lechuga de bototo (vern.): 231. lengua de vaca (vern.): 74, 75. lianas: 40, 47, 48, 49. liga (vern.): 171. lirio (vern.): 230, 231. LOMARIOPSIDACEAE: 69. Loosestrife family: 172. LORANTHACEAE: 167. LYTHRACEAE: 172. Macleania smithiana: 156, 277. Maidenhair family: 73. majado (vern.): 181. majagua (vern.): 181. majagual: 182. Mallow family: 179. MALPIGHIACEAE: 173. MALVACEAE: 179. mangle (vern.): 238. mangle amarillo (vern.): 141. mangle blanco (vern.): 141. mangle bobo (vern.): 141. mangle botón (vern.): 139.

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INDEX TO NAMES OF PLANTS (cont.) Scientific names and plate pages are written in italics, respectively. Vernacular names are indicated by (vern.). mangle caballero (vern.): 238. mangle cholo (vern.): 238. mangle colorado (vern.): 234, 238. mangle de concha (vern.): 234. mangle de humo (vern.): 79. mangle dulce (vern.): 234. mangle gateado (vern.): 234, 238. mangle hembra (vern.): 141. mangle iguanero (vern.): 79. mangle injerto (vern.): 234. mangle jelí (vern.): 139. mangle llorón (vern.): 79. mangle macho (vern.): 234. mangle nato (vern.): 161. mangle negro (vern.): 78, 79. mangle patucho (vern.): 238. mangle pava (vern.): 238. mangle pecho de pavo (vern.): 238. mangle piñuelo (vern.): 247. mangle prieto (vern.): 79. mangle rojo (vern.): 234, 238. mangle salado (vern.): 79, 139. mangle zapatero (vern.): 234. manglillo (vern.): 141, 238. manglillo salado (vern.): 194. mangosté (vern.): 184. Mangosteen family: 137. manguté (vern.): 184. mano (vern.): 95. manoa (vern.): 141. MARCGRAVIACEAE: 183. marimacha (vern.): 231. mata de coca arisca (vern.): 135. matapalillo (vern.): 172. matapalillo de barrial (vern.): 173. matapalo (vern.): 137, 138, 154, 169, 170, 189, 190, 191, 192. Maxillaria hennisiana: 210, 288. MELASTOMATACEAE: 186. Melastome family: 186. Miconia reducens: 187, 284. Microgramma percussa: 71, 257. Microgramma reptans: 71, 257.

Monocotyledons: 47, 48. Monstera pinnatipartita: 96. monte salado (vern.): 194. mora (vern.): 187. Mora megistosperma: 161. Mora oleifera: 161, 279. MORACEAE: 189. Muellera chocoensis: 163, 278. mulata (vern.): 133. Mulberry family: 189. murrapo (vern.): 102. musgo (vern.): 127. MYRSINACEAE: 193. naidí (vern.): 102. naidizales: 102. nato (vern.): 161. nato rojo (vern.): 161. Nephrolepis biserrata: 69, 257. Niphidium crassifolium: 71, 258. Notopleura epiphytica: 244. Notylia replicata: 210, 289. NYCTAGINACEAE: 193. Orchid family: 194. ORCHIDACEAE: 194. Ornithidium fulgens: 211, 289. Ornithocephalus bryostachys: 211. Oryctanthus florulentus: 168. Oryctanthus occidentalis: 169. pajonal (vern.): 228. Palm family: 100. palma (vern.): 101. palma de coco (vern.): 101. palma naidí (vern.): 102. palmicha (vern.): 102. palmiche (vern.): 102. palo de sal (vern.): 78. palo salado (vern.): 79, 139, 141, 194, 238. Panama hat family: 144. parachique (vern.): 83. parasites: 42, 47, 48, 49. Paspalum vaginatum: 227. Pavonia kearneyi: 180. Pavonia rhizophorae: 180, 283.

Pelliciera rhizophorae: 247, 297. Pellicieraceae: 247. Peperomia blephariphylla: 220. Peperomia ciliaris: 221. Peperomia elongata: 221. Peperomia emarginella: 222. Peperomia gorgonillana: 222. Peperomia jamesoniana: 223. Peperomia obovalis: 223, 292. Peperomia obtusifolia: 223, 292. Peperomia pernambucensis: 224, 292. Peperomia portobellensis: 225, 292. Peperomia rotundifolia: 225. Peperomia serpens: 226. Pepper family: 219. Philodendron chrysocarpum: 97. Philodendron musifolium: 98, 263. Philodendron sparreorum: 98, 263. Philodendron tripartitum: 99. Phoradendron chrysocladon: 249. Phryganocydia phellosperma: 108, 265. Phthirusa pyrifolia: 169. Phthirusa stelis: 170. pija de perro (vern.): 117. pikuyo (vern.): 126. piña de monte (vern.): 76. piñuela (vern.): 247. piñuela de monte (vern.): 114. piñuelo (vern.): 247. Piñuelo mangrove family: 247. pipa trees (vern.): 101. PIPERACEAE: 219. pitahaya (vern.): 132, 133. pitahayita de monte (vern.): 133. pitajaya (vern.): 133. Pitcairnia heterophylla: 119. Platystele cornejoi: 212, 289. Pleopeltis bombycina: 72, 258. POACEAE: 226. Polypodiaceae: 69. Polystachya concreta: 212, 289. Pondweed family: 231.

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PLANTS OF THE SOUTH AMERICAN PACIFIC MANGROVE SWAMPS (Colombia, Ecuador, Peru)

INDEX TO NAMES OF PLANTS (cont.) Scientific names and plate pages are written in italics, respectively. Vernacular names are indicated by (vern.). PONTEDERIACEAE: 229. POTAMOGETONACEAE: 231. Prosthechea fragrans: 213. Prosthechea pseudopygmaea: 213. Psammisia occidentalis: 157. Pseudosarcopera diaz-piedrahitae: 184. Psittacanthus divaricatus: 171, 281. PTERIDACEAE: 73. Pteridophyte: 47, 48. Pterocarpus officinalis: 164, 280. quereme (vern.): 156. racancha (vern.): 74, 75. Racinaea multiflora: 120, 268. Racinaea multiflora var. decipiens: 120, 268. Racinaea multiflora var. multiflora: 120, 268. ranconcha (vern.): 74, 75. Red Mangrove family: 232. Rhabdadenia biflora: 87, 262. Rhipsalis micrantha: 133. Rhizophora brevistyla: 241. Rhizophora mangle: 43, 234, 294. Rhizophora racemosa: 238, 295, 296. Rhizophora x harrisonii: 241. RHIZOPHORACEAE: 232. Rhynchospora corymbosa: 150. romero (vern.): 105. RUBIACEAE: 241. rupia (vern.): 232. Ruppia maritima: 232. Rustia occidentalis: 245, 296. salado (vern.): 79, 194. Salicornia fruticosa: 83, 261. Saltwort family: 104. salvaje (vern.): 127. sangre de dragรณn (vern.): 165. sangre de gallo (vern.): 164. Sarcinula sp.: 214, 290. Saueda sp.: 84, 261. Scaphyglottis crurigera: 214, 290. Scaphyglottis graminifolia: 215. Scaphyglottis prolifera: 215, 290.

Scaphyglottis sp. nov.: 216, 290. Schlegelia darienensis: 246. Schoenoplectus californicus:151. Schwartzia chocoensis: 184. Scleria eggersiana: 151. Scleria melaleuca: 152. Scleria mitis: 152. SCROPHULARIACEAE: 246. Sedge family: 146. seed plants: 77. Serpocaulon triseriale: 72, 258. Sesuvium portulacastrum: 81, 261. Showy Mistletoe family: 167. shrubs: 38, 47, 48, 49. Sobralia madisonii: 216, 291. Sobralia rhizophorae: 217, 291. Soterosanthus shepheardii: 217, 291. Souroubea guianensis: 185. Souroubea intermedia: 185. Sphaeradenia acutitepala: 145. Sphaeradenia killipii: 145, 276. Sphyrospermum buxifolium: 157, 277. Stenospermation angustifolium: 99. Stenospermation robustum: 100. Stigmaphyllon ellipticum: 178, 282. stranglers: 42, 47, 48, 49. Struthanthus orbicularis: 172, 281. suela (vern.): 164. suela de playa (vern.): 164. suelda con suelda (vern.): 171, 200. Tabebuia palustris: 108, 265. Talipariti tiliaceum var. pernambucense: 181, 283. taruya (vern.): 230. Tea family: 247. TETRAMERISTACEAE: 247. Tetrapterys subaptera: 178, 282. Theaceae: 247. Thibaudia pachypoda: 158, 278. Tillandsia barclayana var. minor: 121, 268. Tillandsia bulbosa: 121, 269. Tillandsia complanata: 122, 269. Tillandsia disticha: 123, 269.

Tillandsia dyeriana: 123, 269. Tillandsia flagellata: 124, 270. Tillandsia latifolia var. divaricata: 124, 270. Tillandsia narthecioides: 125, 270. Tillandsia subulifera: 126, 270. Tillandsia triglochinoides: 126, 271. Tillandsia usneoides: 127, 271. Tillandsia venusta: 128, 271. Tobagoa maleolens: 245. Tococa guianensis: 188. tortuguero (vern.): 107. totora (vern.): 151. totumillo (vern.): 106. trees: 38, 47, 48, 49. tres dedos (vern.): 95. Trichomanes ankersii: 68. Trichomanes crinitum: 68. Trichosalpinx orbicularis: 218, 291. Trigonidium seemannii: 218. Trumpet-creeper family: 105. Tuberostylis axillaris: 103. Tuberostylis rhizophorae: 104, 264. tumbagaspar (vern.): 228. Uniola pittieri: 228, 293. Urochloa mutica: 229. Verbenaceae: 77. verdolaga (vern.): 81. vidrillo (vern.): 83, 105. vidrillo salado (vern.): 83. vidrio (vern.): 105. VISCACEAE: 249. Vittaria lineata: 75, 258. wam bio tape (vern.): 152. Water hyacinth family: 229. Werauhia cf. kupperiana: 129, 272. Werauhia gladioliflora: 129, 271. Werauhia ringens: 130, 272. Werauhia sanguinolenta: 130, 272. zamia de los manglares (vern.): 76. Zamia roezlii: 76. Zamiaceae: 76. zeneida (vern.): 229.

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At the same time Xavier served as curator at

GUAY, he taught Botany at the University of Guayaquil and the Pontifical Catholic University of Manabí (PUCE-M). He was also involved in floristic inventories across the country and was cited among the most important collectors of Ecuadorean plants by Jørgensen and León-Yánez in The Catalogue of Vascular Plants of Ecuador (1999). Based on his field experience he authored revisions of several families and served as the scientific Xavier Cornejo, during field work in mangroves of Ecuador, 2010. Photo by Máximo Cangá.

advisor to The Red Book of Endemic Plants of Ecuador (R. Valencia et al. eds., 2000; S. León-Yánez et al. eds., 2011). In addition, he has contributed taxonomic treatments of Proteaceae (2002, junior author), Nymphaeaceae (2003) for Flora of Ecuador, and the Flora del Bosque de Garúa de la comuna Loma Alta, Prov. Guayas (2004, junior author). Between 2004–2006, he was an honorary

The senior author

fellow at the Department of Botany of the University of Wisconsin-Madison where he carried out research on

Xavier Cornejo (1968) is an Ecuadorian-

American botanist.

He was born in Guayaquil, the

largest city of Ecuador. In 1991, he started as a field

Capparaceae (the caper family) with Hugh H. Iltis for the Flora of Ecuador and Flora Mesoamericana (both submitted).

assistant in a project dedicated to floristic inventories and environmental studies of the mangroves of Ecuador.

In 1994, he graduated with a degree in Biology from the

Botanical Garden staff as a curatorial assistant of imaging

Faculty of Natural Sciences of the University of Guayaquil

vascular plant types, and was the manager of the Vascular

and, in 1993, started working as a plant collector. Later he

Plants of the Osa Peninsula, Costa Rica. Currently, he

became a curator for the herbarium of the University of

collaborates with Scott A. Mori on research of the Brazil

Guayaquil (GUAY), a position held up to October, 2004.

nut family (Lecythidaceae), especially those of Ecuador

During that time Xavier traveled extensively to explore

(http://sweetgum.nybg.org/lp/ecuador.php),

the forests in the small but heterogeneous and biologically

a specialist on the genus Grias. Xavier has described

rich Ecuador, collecting 8,000 numbers of vascular plants,

60 species of vascular plants new to science in several

including many new species and new records for the

families from tropical America and also discovered a new

country. The main set of his collections is in GUAY, and

species of flower mite on red mangroves. He also works

duplicates have been distributed to specialists at AAU,

restoring disturbed forest with native plants and planting

COL, GB, K, MO, NY, QCA, QCNE, SEL, US, and WIS.

native species as ornamental in the country.

Between 2007–2012 he joined The New York

and