The vegetation of Panama: A review - Porter 1973

VEGETATION-AND VEG-ETATIONAL HISTORY OF NORTH ERN LATIN AMERICA PAPERS PRESENTED AS PART OF A SYM POSIUM, " VEGETATION AND VEGETAT IONAL HISTORY IN NORTH ERN LAT IN AMERICA", AT TH E AMERI CAN INSTIT UTE OF BIOLOGICAL SCIENCES MEETJN G,S. BLOOMINGTON, IN D. (U.S.A.), 1970

edited by

ALAN GRAHAM Departmelll of Biological Sciences, Ken/ State University, Kent, Oflio (U.S .A.)

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ELSEVIER SCIENTIFIC PUBLISHING COMPANY Amsterdam - London- New York

. 1973

Contents Preface . . . . . . . . . . . . . . . . . .

v

Chapter 1. The Vegetation of the Antilles RICHARD A. HOWARD Summary . . . . . . . . l ntroduct iOJl . . . . . . .

Vegetation types in the Antilles Coastal formations .

Lowland formations Montane forma tions Qrigins of the existing vegetation of the Antilles Relationships and distribution of the Antillean vegetation Disjunct distribution patterns in the Antilles . . . . .

Distribut ion patterns wit hin the islands . .

Long-distance 1elationships and disjunct distribution patterns Conclusions References . .

4

5 8 12 16 17

29 31 34 37 37

Chapter 2. A-Paleoclimatic Interpretat ion of the Eocene Floras of Southeastern

North America DAVID

L.

D ILCHER

Summary . . . . .

Introduction . . .

Individual components of the fl ora Identification to modern taxa Evolution of the fossil taxa .

Relation of range and ecological tolerance.

Community structure . . . . . . . . . . . .

Stability through time . . . . . . . . .

Separation into equivalent modern communities Foliar physiognomy . . . .

Leaf-margin analysis . . . . . . . .

Leaf-size analysis . . . . . . . . .

Interpretation of foliar physiognomy .

Application to this nora Additional evidence Wood . . . . . . .

Pollen . . . . . . .

Paleoclimatic interpretations References . .

39

40 41 4t 42

43 44 44 45

46 48

50 53 54 55 55 55 56 57

Chapter 3. G eographical Relationships ofthe Flora of Mexican Dry Regions J. RZEDOWSKI Suillmary .

Introduction

61 61

f X

CONTENTS

CONT ENTS

Relationships between diOcrent arid zones of Mexico Relationships with other arid zones of the world Relationships with zones of non-a rid climate .

Conclusions References . . . . . . . . . . . : . • .

63 1;6'

68

70 71

Clrapter 4. Ecology of the Vegeta tion of Veracruz ARTURO G6MEZ-POMI'A

Summary . . . .

Introduction . . .

Physical condi lions Topography Climate Soils .

Vegetation . .

Forests Selvas .

High sclvas .

Low selvas .

Grasslands .

Arid or semi-arid vegetation Secondary vegetation References . . . . . . . . . . .

73 73 73 73 75 85

89 94 105 106 118

.-

128 135

145 147

Chapter 5. The Phytogeography and Vegetation ofChiapas (Mexico)

149

149 150 154

155 155 158 161 163 164

Chapter 6. The Vegetation of Panama: a Review D UNCAN M. PORTER

Summary .

Introduction Climate Soils Man Vegetation .

Vegetation maps . . .

Koppen macroclima tes Holdridge Life Zones Human innucnccs . . .

Darien forest types . _ .

A summary of plant comnumiti<.,-s Incipient Forest . . . _ Early Secondary Forest Late Secondary Forest Liltoral Forest . . . .

Thorn Forest _ . . . .

Deciduous Seasonal Forest .

Savanna .. . . . . _ .

S<tltwatcr Swamp Forest .

Saltwater Riparian Forest.

Brackish Swamp Forest _ llrackish Riparian Forest Freshwater Marsh . . .

Freshwater Swamp Forest Frcshw<tter Riparian Forest .

Monsoon Forest . . . .

Evergreen Seasonal Forest Premontane Rain Forest Cloud Forest Elfin Forest Acknowledgements ~ References . . . .

17 1

171 172 173 174 175

175 177 179

182 183 184 185 187 187 187

189 189 190 192

193 197

197 198 199 199 200

Chapter 7. Phytogeographic History of the Isthmus of Panama during the Past

DENNIS E. B REEDLOVE

Summary . . . . . . .

Introduction . . . . . .

Phytogeography of Chiapas .

Vegetational formations ofChiapas Description of the formations .

Optimum forma tions. . . .

Seasonal format ions . . . .

Wet-land forest formations .

Treeless formations References . . . . . . . . . . .

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IX

167 167 169 170

170 171

12,000 Years (A History of Vegetation, Clima te, and Sea-level Change) ALEXANDRA S. B ARTLETT AN D ELSO S. BARGHOORN Summary . . . . . . . . . .

Introduction . . . . . . . . .

Northern South America .

Southern Central America Panama . . . . . . .

Geological history of Panama ..

Pre-Quaternary history . .

Late Quaternary and Recent history Physiography' and climate of Panama .

Climate of the Canal Zone The present vegetation of Panama Preparation ofsamples . . . .

Preparation techniques .

Preservation of samples The pollen diagrams . . . . .

Theoretical considerations The Gatun Basin diagrams The modern pollen rain .

Climatic change in Panama . . .

Identification and nomenclature of the fossil pollen .

203 204 .-

204

206 207 208

208 209

211 211

212 216 219

219 2 19 2 19

222 225 228

229

Chapter 6

The Vegetation of Panama: a Review DU NCAN M . .PORTER Smithsonian lustitllliou, Washington D. C. (U.S.A.)

Summary The physiograph y, climate, soils, and t he influence of man on the vegetation of the Isthmus of P anama are outli ned. Followi ng a brief disc ussion of the vegetation maps of the isthmus that heretofore have been produced , more detailed critiques are offered of Bennett's (1968) and Myers' (1969) maps of Koppen M acroclimates, Holdridge and B ud owski's (1956, 1959) map of H oldridge Life Zones, Bennett's ( 1968) map indicating human d isturbance, and Duke's (Duke and Porter, 1970) map of Darien forest types. The last section of the paper p resents lists of species and their relative abundance, when-given, that investigators have reported from various plant commun ities in the Republic of Panama and the Canal Zone. ~

Introduction The Isthmus of Panama stretches sinuously between Costa Rica on the west and Colo mbia o n t he east; north is the Car ibbean Sea, so uth the Pacific Ocean (Fig. I)

Fig.!. Map of Panama, showing political subdivisions.

168

D. M. PORTER

THE VEG ETATION OF PANAMA : A REV IEW

169

The Continental Di vid e lies approximately along the center of the western part of the isthmus. In the low central par!, ius much closer to the Pacific side, while in the eastern half it lies closest to the Caribbea n shore.

Flat and gently sloping low land plains arc not common on the isthmus. Most lowland areas are dissected by rivers and low ridges and hi lls. The most extensive lowlands occur on the Pacific side. These are in the provinces of Darien, Panama, Code, VcragLias, and Chi riq ui, and on the Azuero Peninsula. Lowland areas on the Caribbean side are mai nly west of the Canal Zone in Colon and Panama provinces and in the vicinity of Chiriqui lagoon in the province of Bocas del Toro.

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Geograph ically and biologically, this geologically rather recen! isth mus is a bridge between Cent ral and South America. I t has the shape of a sigmoid curve, lying between 7• 10' and 9°40'N, with its axis runn ing essentially east and west from 77° 15' to 83°30'W. In the center of the isthmus, bisecting the Republic of Panama, lie the Panama Canal and the Canal Zone. The Republic of Panama and the Canal Zone together cove r about 48,000 km 2 . It is 676 km from border to border, and the ist hmus varies from 50 to !93 km in width.

Elevations range from sea level to 3,427 m, but most of the ist hmus has an elevation of less than 500 m, and nea rly 90% is less than I,000 m high (Fig.2). The western half of the isthmu s is traversed by two ~;;o n ti n uou s volcanic and igneous mountai n ranges with Panama's highest elevations. The Cordillera de Talamanca in the west is conti nuous with the eastern highlands of Costa Rica. The eastern Serrania de Tabusara and its assoc iated high lands are separated from the mountainous areas or eastern Panama by about 65 km of lowland s in the center of the isthmus. The Panama Canal was built across this break in the isthmus' backbone, which here is a little over 60 m above sea leveL South of t he Serrania de Tabasara, the Azuero Peninsula juts out into the Pacific Ocean. Much of the Azuero Peninsula is low, but near its southwestern end it rises to the isolated Cerro Hoya, with an elevation of about I ,500 m.

Elevations in the eastern half of. the isthmus are much lower than those in the west, few areas rising over 1,000 m. The long Serrania de Darien, which parallels the northeastern coastline, in the main averages between 300 and 600 m high, with higher elevations in its southernmost parts. The highest spots in eastern Panama are isolated peaks, the highest of which is Cerro Tacarcuna, on the Colombian border, with an elevation of 1,859 m.

Rainfall patterns and temperatu re are the most impor tant climatic factors for plant distribution in the humid tropics. Elevation and temperature are closely interrelated; temperature decreases as elevation increases. Variatio ns in monthly mean temperatures appear to be slight at all elevations in the tropics. Diurnal temperature variation, however, becomes more important as elevat ion increases, the range becoming wider as one goes higher.

The tropical year is not divided into seasons by variations in temperature, as is true for the temperate zones. fn the tropics the year is divided into wet and dry seasons on the basis of the relative amounts or rainfall falling at different times. L stress relatil•e amounts of rainfall, because a month with 12.5 em of rain at Balboa Heights, on the Pacific side of the Canal Zone, will be part of the wet regime, while ~t Changuinola, Bocas del Toro, it will fall into the dry season (Panama Canal Company, 1967).

The climate of Panama is greatly influenced by the Intert ropical Convergence Zone (lTC), that area where the trade winds from the Northern and Southern Hemispheres meet. The lTC roughly parallels the equ ator. Heavy precipi tation .and unstable rising air masses are associated with the lTC throughout much of the humid tropics. The ITC shi fts north and south with the seasonal shifting of the sun. In the eastern Pacific, it dips southward to reach th e equator in January, rarely reaching 2° or 3°S. In July the lTC has moved northward to lie along the fsthmus of Panama, but it apparently does not enter into the Caribbean Bas in (Trewartha, 1961 ).

This latitudinal shifting of the lTC has a profound effect on the weather of Panama. In the Pacific lowlands, the wet season is essentially from May through December, and the dry season, usually with less than 5 em of rain per month, from January through Apt iL In some localities periods as long as two months without measurable precipitation have been reported. The dry season on the Pacific side is less pronounced at higher elevations, with some areas reported to be almost continually covered by low-lying cloud ~.

The dry season in most of lowland Caribbean Panama is not so pronounced as

170

D. M.l'ORT ER

that of the Pacific side. When the lTC is at the equator in January, and the dry season has begun in Paci lk Panama , the Caribbean side of the isthmus is influenced by moisture-laden clouds that have formed over the Caribbea n Sea and arc now moving · southwa rd. Most of their precipitation falls on the Caribbean side. This results in a few lowland locali ties having no apparent dry season at all. Altitudinal effects on precipitation are even more pronounced on the Caribbean slopes than on the Pacific. As a generalization, it may be said that the vegetation on the Caribbean side is more mesophyt ic and less seasonal than that on the Pacific side.

Soils Tempera tu re and precipitat·ion clearly are not the only parameters affecting plant distribution in Panama. Edaphic factors may be just as important.

However, few vegetation stud ies mention edaphic factors, and fewer studies still have been made on Panamanian soils. In· the soils maps that I have examined (Bennett , 1912, 1929; Martini et al., 1960), there appears to be little or no correlation. between soil types and vegetat ion types. However, those plant communities dominated by a single species, such as Prioria copaifera in the Freshwater Swamp Forest of Darien, have specialized edaphic conditions. As those familiar with tropical vegetatio n are well aware, there are few communities in the humid tropics with a single do minant species. Usually there arc a large number of common species, with some of the lusher Panamanian forest communities having upwards of 50 species of large and conspicuous trees scattered throughout.

- Man Probably the most importa nt influence on Panama's vegetation since pre-Colu mbian times is man. Man's fires, axes, and domestic animals have had a profoun d effect on the plants of the isthmus. This effect is by no means limited to the drier lowland commun ities. I have seen hundreds of hectares of blackened hillsides where once rose forests with over 2,500 mm of average annual precipitation (Porter, 1970).

Such areas were cut.and burned to provide grazing grounds for cattle. It is only after these forests are cut and burned that the cattlema n finds that the resulting growth will not support a significant amount of grazing.

Even the mangrove communities are not immu ne to the hand of man. Mangrov es long have been exploited for charcoal, but now they have fallen victim to another offspring of technology, the oil-spill (Riitzler and Sterrer, 1970).

TH E VEGETA TION O F PANAMA : A REVIEW

171

Vegetation

Vegewtion maps There have been a number of attempts to produce a map indicating the vegetatio n of Panama. These have been either as a part of a larger map of Latin America (Smith and Johnston, 1945), Central America (Trejos and Archer, 1953; Holdridg e, 1957; Lauer, 1960; Shelfo rd, 1963; Wagner, 1964; Stuart, 1966), southern Central America (Sapper, 1900; James, 1950), or a map of Panama itself (Goldman, 1920; Goldman and Zelek, 1926; Platt, 1938; Schery, 1942; Garver, 1947; Holdridge and Budowski, 1956, 1959; Bennett, 1968; Myers, 1969). The vegetation of parts of the isthmus also has been mapped, incl uding eastern Pana.ma (Lamb, 1953; Gollcy et al., 1969; Duke and Porter, 1970; Viksne et al., 1970), San Jose Island (Erlanson, 1946; Johnston, 1949), Barro Colorado fsland (Park eta!., 1940; Bennett, 1963), and Fort Sherman and Fort Gulick (Wiley et al., 1955). The vegetation of Darien Province has been mapped in detail (Duke and Porter, 1970), but few detailed vegetat ion maps are available for most of the isthmus. Almost all maps that have been produced show only the gross plant formations, and even these maps may vary marked ly from one another. T he only maps of which I am aware that attempt to deli mit plant communities on a large scale are those of Johnston (1949) for San Jose Island and Duke (Duke and Porter, 1970) for Darien.

Koppen macroclimates Bennett (1968) and Myers (1969) recently have produced maps of Panama delimitin g the Koppen macroclimates. Fig.3 combines their results. Many investiga tors feel that

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(After Bennett, 1968, and Myers, 1969.)

172

D. M. PORT ER

Koppen macroclimatcs, which arc based on tem perat ure and precipitat ion, arc broadly equiva lent to major vegetation types.

Koppen's A and C cli mates occu r in Panama. A cl imates arc tropical rainy climates with no cool season, the average tempera ture of the cooles t month being above 18°C. c cli mates arc warm temperate climates with mild win ters, the average temperat ure of the coldest month being between 18 and - 3°C, and that for the warmest month over 10°C. .

Climates reported for Panama are Tropical Wet, with no dry season and all months hav ing over 6 em precipitation, equated with L owland Tropical Rain Forest; Tropical Monsoon, with a short dry season, but rainfall during the wet season sufficient to compensate for it, and all months having over 6 em precipitation, equated with Monsoon Forest; Tropical Wet and Dry, wi th a longer dry season having at least one month with less than 6 em precipitation and insufficient rain during the wet season to compensate for it, equated with Tropical Deciduous Forest; Temperate Wet, with high rainfall and no distinct dry season, the lowest month receiving more than 3 em precipitation, eq uated with Montane and Lower Montane Rai11 Forests; and Temperate Wet and Dry , with high rainfa ll and a distinct dry season, haviJJg at least ten times as much rain in the wettest mont h as in the driest, equated with Et·ergreen and Semiet·ergreen Seasonal Forest.

The Koppen ~ystem perhaps is useful fo r a large-scale generalized map, but it is not very useful for small areas or for detailed vegetation mapping. Also, one cannot equate vegetation types with Koppen macrocli mates on a one-to-one basis.

Holdridge Life Zones

Holdridge and Budowski (1 956, 1959) have produced a Life Zone map of Panama (Fig.4). Holdridge's life zones are based on the ratio between potential evapotranspiration and average total annual precipitation, and they are equated with gross vegetation formations. Their map gives six life zones and two transition zones for Panama: Tropical Moist Forest, Tropical Dry Forest, Subtropical Moist Forest, Lower Montane Wet Forest, Mon tane Wet Forest, Tropical Moist Forest (Transition), and Tropical Dry Forest (Transition).

According to Holdridge and Budowski, the areas of transition between two life zones generally are not extensive enough to be mapped. Bu t in those areas where there are gradual shifts in temperature or precipitation, broad transition zones can be recognized, as they have done in eastern Panama.

Both the Koppen and Holdridge systems ignore edaphic, anthropogenic, physiognomic, and flo ristic factors. However, it must be recalled that Koppen's system primarily attempts to delimit climatic zones, not vegetation per se. Like the previous map (Fig.3), this map (Fig.4) does not show the vegetation actually occurring in each area, but it gives the gross plant formations that ideally should occur in a given area.

TH E VEGETATION OF I' ANA MA: A REVIEW

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Human influences

Unfortunately, the information on which the Koppen and Holdridge systems are based is lacking for much of Panama. Be that as it may, vegetation cannot be mapped from environmental parameters alone, no mal!er how much cl imatic informat ion is avai lable. The way to map vegetation is to go into the field and look at it.

One investigator who has done just thi s is Bennett (1968), who is noteworthy for assessing the impact of man on the vegetation of the Isthmus. Bennett's thesis is that man has had a profound effect on the vegetation as long as he has been in Panama.1 In the past this effect was greatest at the time of the Spanish Conquest, when the population was approximately what it is now. The Indian population, drastically reduced by disease and slavery, has slowly been replaced by various ethnic groups until Panama's population again approximates what it was 500 years ago. H uman pressures on the environment also have varied with the population.

Bennett recognizes a number of vegetation types for Panama (Fig.5). Mature Forest is forest that has had only mino r or intermittent disturbance by man ; Old Forest is now growing where records indicate previous important disturbance, but it is now little used; and Disturbed Forest is presently being disturbed by various forms of exploitation. Low Trees andfor Shrubs occur where there are disturbed conditions due to fire, overgrazing, and associated edaphic changes. Thorn-scrub and Cacti a re not due to man, climatic and edaphic factors apparently being of primary importance, although fire also may be involved. Sat>anna is found in areas where t here is marked disturbance; such areas would normally support forest. Grassy Plains would support scrub if left unburned. Grass and/or Sedges on Steep Slopes apparently are due to fire 1

For further information on this subject, see the excellent book by Sauer (1966).

174

D. M. PORTER

T HE VEGETA TION OF PANA M A: A REV IEW

175

A s ummary of plant commun ities

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In the absence of a detai led vegetati on map for Panama , the fo llowing is a step toward such a goal. Studies on the biology or the isthmus often have mentioned species and thei r abunda nce in given planf comm unities. The lists that fo llow at tempt to use contem porary nomenclature for the species ci ted and to p rovide equivalents for the comm unity na mes used in the original p ublications. H owever, no attempt has b.een made to provide equivalents for the original terms or relative abunda nce.

The commu ni tic~ arc listed in order o r: (/) decreas ing disturba nce (Incipie nt F o rest, Early Second ary f o rest, La te Second ary F o rest); (2) decreas ing xericity (Li ttoral f orest, Thorn Forc"St, Deciduo us Seasonal F o rest, Savann a); (3) decreasing salinity (Saltwater Swamp forest, Saltwat er Riparian F o rest, Brackish Swamp Forest, Brack ish Riparia n F orest, Freshw ater Ma rsh, F reshwa ter Swamp Forest, freshwa ter R iparian Forest) ; a nd (4) increasing rainfall a nd increasing altitude (Monsoon Forest, Evergreen Seasona l fore st, Premon tane Rain forest, Cloud Forest, Elfi n Forest) .

Incipient .forest

F ig.5. Phytophysiognomic map of Panama. (After Bennett, 1968.)

and would support fo rest or scrub if unburned . Galeria Forest is simply forest a long rivers tl1at has not been cut, unlike the su rrounding area.

Banana Plantations are self-explanato ry.

Bennett 's map is t he best yet produced that represen ts the actual vegetati on of the entire isthmus on a large scale. Unfortu nately, however, it is no t precise enough to be used on the local level.

Darien forest types The most detailed vegetati on map on a large scale yet p rod uced for Panama is D uke's map of t he forest types of D arien (Duke and Porter, 1970).

This map resulted from studies initia ted by the Interoceanic Sea-Level Canal Feasibi lity Study. Duke in the main has followed Beard's terminology in delimiti ng forest types (Beard, 1944, 1953, 1955a,b). Unlike most o ther vegetati on maps of Panam a, much of this map has been verified by intensive field studies on the ground . Unfortu nately, sufficien t information still is not availab le to produce such a detailed map of the plant commu nities of the entire Isthmu s ofPana ma.1 1 Duke's map originall y appeared in a publication of limited circulati o n. It will be published in the Annals of the Missouri Botanical Garden.

[Duke and Porter (1970), Darien]: frequent trees:

Carim papaya L. Coch/ospel'llwm ••itifolillm (Willd.) Spreng.

Cocos 1/llcifera L. locally dominan t shrubs:

Mull/ ingia ca/abura L. frequent shrubs :

Acalypha macrostachya Jacq.

Cestmmlati/olillm Lam. C/ibadiumleiocarpum Stectz.

Lantana camara L. L. tri/olia L. Leandro dichotoma ( D. Don) Cogn.

Manihot esculenta Crantz Neurolaena Iobato (L.) R. Br. Piper aduncum L. Pothomorphe peltata (L.) Miq. Rolandrafruticosa (L.) 0 . Ktze.

Solanwnllirttmt Vahl Struchium sparganophorum (L.) 0. Ktze.

Vemonia canescens H .B.K. V. patens H.B.K. locally dominan t banana-like o r small palmlike plants: Heliconia latispatha Benth.

frequent banana-like o r small pa lm-like plants: Carludovica palmata Ruiz and Pav. Helicouia mariae Hook. f.

Zea mays L. freq uentlianas:

Elc11sine indica (L.) Gaertn.

Solanllmjanwicense Mill.

Wulffia baccata (L. f.) 0. Ktze. locally dominan t vines:

Dioscorea spp. frequent vines :

Cissus rhombi/olio Vahl C. sicyoides L. Dalechampia tiliifolia Lam. Desmodium adscendens (Sw.) D C. D. canwn (Gmel.) Schinz and Thellung Eupatorium ir·esineoides H .B.K. E. macrophyllum L. E. microstemon Cass.

E. odoratum L. Gurania coccinea Cogn.

Mik ania guaco H umb. and Bon pl.

Momordica clrarantia L.

176

D. M. PORTER frequent herbs :

1'1t:alyplw an·en.\'i.v l)oc

Al!emtllllheraficoidea (L.) R. Br. Asclepias Cl/ras.wivicll L. Bo11rreria/aet•is (Lam.) Grise b.

Bro wal/ia wuericmw L. Cllpsicmu ji-11tesceus L. Ceurhms brownii Rocm. and Schult.

C. eciliuattls L. Chamacsyce hirta (L.) Millsp.

C.l1ypericifolia (L.) Small C. hyssopifolia (L.) Millsp.

C. 1hymifo/ia (L.) M illsp.

Coix /achryma-jobi L. Cypems dijfusus Vahl Dichromeua cilia to Vahl Digitaria sauguiualis (L.) Scop.

Echiuoch/oa co/onm11 (L.) Link Eclipta alba ( L.) Haask.

Eleutherauthera mdera/is (Sw.) Sch. Bip. Erechtites hieracifo/ia (L.) Raf. Erlcerou bonarieusis L. Fimbristylis miliaceo (L.) Vahl liyptis capita/a Jacq.

H. savwmarum Briq .

H..<uat•eoltms (L.) l'oit.

H . t•erticil/ata Jacq.

Just ida pectoralis J acq.

Lmlwigiu spp. (as Ju.uiaca ;pf1.)

Oi'imwumicrantlwm Willd.

Oryza sati•·a L. Pauimmmaximmu Jacq.

P. pi/osum Sw. Po.rpalmu p/imtalum Michx.

P. ••irgotmu L. Phj•l/outlms urinaria L. Polygala pouiculata L. Porophyl/um ruderale (Jacq .) Cass.

Portulaca oleracea L. Pseudclwhantopus spicat11s (Juss.) Rohr.

Smtvage:da erect(l L. Scoporio d11/d;· L. Sida glomertua Cav. S.pymmidata Cav. S. rlwmbifo/ia L. So/auumnigrum L. (s. 1.)

Spilamhe.< ocymifolia (Lam.) Moore Synedrella nodi/fora (L.) Gaertn.

Waltheria indica L.

[Johnston (1949), Bale tlill, San Jose Island, Panama (as "Grassland")]: dominant grasses:

Axonopus aureus Beauv.

Trachypogon secundus (Pres!) Scribn. the most conspicuous secondary grass :

Panicum mdgci Roem. and Schul t. other herbaceous plants:

A eschynomene brasiliana (Poir.) DC. Andropogon bicomis L. A. brevifolius Sw. A.leucostachyus H.B.K. Aristida capillacea Lam. Axonopus centra/is Chase Blepharodon mucroJwlatum Decne.

Bourreria /atifolia (Aubl.) Schum.

B. parvifolia Meyer B. pumilia Standi.

Bulbostylisjunciformis(H.B.K.) Lindman Calopogonium mucwtoides Desv.

Centrosema pubescens Ben th.

Chamacsyce hyssopifo/ia (L.) Small (as Euphorbia hyssopifo/ia L.) Clitoria rabigbzosa Juss.

Coutoubea spicata A ubi.

Crotalaria pilosa Mill.

Curculigo scorzoneraefo/ia (La m.) Baker Curtia tcnel/a (Mart.) Ch am.

Cypems/favus (Vahl) Necs Desmodium augu.wifolium (H.B.K.) DC. D. barbatum (L.) Bcnth. and Ocrst.

Dichromena ciliato Vahl Dioscorea sapindoides J>resl Eleplwntopus mol/is H.B.K. Eleutherrmthera mdera/is (Sw.) Sch. Bip. Habenaria paucif/ora (Lind!.) Re ichb. f.

Hyptis lantauaefo /ia Poit.

Ipomoea miuutif/oro (Mart. an d Gal.) House Kylliugia odorala Vahl Oplismenus burmannii (Retz.) Beauv.

Paspa/ummulticaule Poir.

P.plicatulum Michx.

Phaseolus longipedunculatus Mart.

Pterolepispumila (Bon pl.) Cogn.

Sauvagesia tene//a Lam. Sc/eria micrococca (Liebm.) Steud.

S . pterota Pres I Setaria lenox (L. R ich.) Desv.

Sida linifolia Cav. S . spinosa L. Spirantlies orchioides (Sw.) L. Rich.

Waltheria indica L. (as W. americana L.)

TH E VEGETATIO N OF PANAMA: A R EV IEW

177

[Standley ( 1928), a bandoned vegetable gardens about Balboa, Canal Zone] : weeds:

persisting garden plants :

Cordwms spp.

Desnwdimn spp. (as Meibomia) Sidaspp.

Dio.,·corc'a spp.

DtJiic/ros lab/ub L. /1ibi.1·ms esculemus L. Manihot esw lenta Crantz

Early Secondary Forest [Duke and Porter (1970), Darien]: locally dominant trees: Oc/n·oma pyrtu11ida/e (Lam.) Urb. Trema micrantlw (L.) Blume frequent trees:

Carica papaya L. Casearia niti<la (L.) Jacq.

Cochlo>1Jertm/ln vitifolium (Willd.) Spreng.

Cordia al/iodora (Ruiz. and Pav.) Roem. an d

Stmdlium sparganophomm (L.) 0 . Ktzc.

Tollmcfortiu CllspidatCJ H .llK. Triam/iiiiCJ lrrpfJU/a L. Vemo11i11 conescens H.B.K. V. patcm H.B.K. locally dominant banana-like or sma ll palmlike plants: Heliconia latispatha Benth.

Schull.

Didymopcmax moroto/oni (Aubl.) Dec. and Planch.

Guawma ulmifolia Lam. He/iocarpus popaycu1ensi;· H .B.K .

Luehea ;peciosa Willd.

P.l'idium guajm•a L. Triplaris cumingitlllll F isch. and Mey. Vi;mia baccifera (L.) T r. and Planch.

V.latifolia Choisy Zanthoxy/11m panamemc P. Wilson Z . setulos11m P. Wilson frequent shrubs:

Acalypha macrostachya Jacq.

Cestmm latifolillm Lam. Clibadiw11 appressipilum S. F. Blake C.leiocarpum Steetz.

Clidemia octona (Bonpl.) L. 0. Wms. Gonzalag1111ia mdis (Standi.) Stand i.

Hamelia axillaris Sw. Hasseltiaf/oribmula H. B.K. lsertia haenkeana D C. Lantana camara L. L. trifolia L. Leandro dic/wtoma (D. Don) Cogn.

Manilwt esculenta C rantz Miconia /acera (Boo pl.) Naud.

Muntingia ca/ab11ra L. Neuro/aena Iobato (L.) R. Br. Piper adwzcam L. P. marginatum Jacq.

Pothomorphe pe/ta/a (L.) Miq. Rolandrafruticosa (L.) 0. Ktze.

Solanum hirtum Yah! S. rugosum Dunal

frequent banana-like or small palm-like plants: Car/udovica palma/a RLtiz and Pav. Hypan'henia nr/tt (Nees) Stapf Heliconia imbricotCJ (0. Ktzc.) Baker H . u1ariae Hook. f.

Renealmio cerm111 (Sw.) Macbr. frequent lianas :

Byllneria oculeata Jacq.

EleuJ·ine indica (L.) G acrtn .

G11rania coccinea Cogn.

Mucuna bracteata D wyer Pluyganocydia corymbo>·a (Vent.) K. Sch. Solmtwnjamaicense Mill.

Wulffta baccata (L. f.) 0. Ktze. frequent vines :

Cissus rhombifo/ia Ya hI C. sicyoides L. Dalechampia tiliifolia La m.

Desmodium camrm (Gmel.) Schinz and Thellung Eupatorium iresineoides H.B. K. E. macrophyllum L. E. microstemon Cass.

E. odoratum L. E. sinclairii Bent h.

Mikania guaco H um b. and Bon pl.

M. micrantlza H.B.K. Momordica charantia L. Rlzynclzosia ca/ycosa Standi.

Sabicea villosa Roem. and Schult. frequen t herbs:

Asclepias carassavica L.

D. M. PORTER THE VEG ETATION OF PANAMA : A REVIEW

/Jourreria lae.,is (Lam.) Grise b.

Browallia amerit·mw L .

Cyperus dij}imts Vahl Diciii'OIIIel/11ciliaw Yah! J:.clitJta alba (L.) Haask.

Eleatlwrwnlwra mderalis (Sw.) Sch. Bip.

Eri.~teron bmwriensi~·

L.

Fimbrist)•lis milia,·ea (L.) Vahl Hyplis :>'ffl'ilunanun Briq.

Justieia peetora/is Jacq.

Ocimummicralllfllmt Willd.

Panicmumoxinmm Jacq.

P. pi/os11m Sw. Pa1palum plicatulum Michx.

P. l'irgalflm L.

l'oruf'lryllllm ruderale (Jacq.) Cass. l'ril'lllappulucca (L.) Pcrs. p,,·cudelcplumtopus .1picatus (Juss.) Rohr.

Scleria melalctl<'a Schlcchl. and Cham.

S. ptemw Pres!

Scoparia dulcis L Sida ~:lomemw Cav. S. pymmidura Cav. S. rlwmbifo/ia L. SpilaullrcsoC)'nll[o/ia (La m.) Moore Synedrella nudiflora (L.) Gaertn.

Wa/tlreria indica L.

(Johnsto n (1949), San Jose Isla nd, Pan;un!t as " High Green Thicket"]: principal trees:

Cecropia spiuesccus L. (as C. araclmoidca Pittier) Cordia al/iudora (Ruiz a nd Pav.) Roem. and Schult.

C. bicolor DC. Hirtel/a americana L. Nectandra ge111/ei Lundell Pltoebejo/ms/oniiC.K. Allen Pilltccellobium rufcsceus (Bent h.) Pitticr Protium remufolium ssp. mcleodii (I. M. J_ohnsl.)

Porter (as P. mcleodii I. M. Johnst.)

Xylopiafmlescens Aubl.

Zuelauia guidouia (Sw.) Britt. and Millsp. secondary trees:

_

Casearia commersoniana Camb. (as C. myrirmtfw T urcz.)

C. syfl•eslris Sw. Cordia panamcnsis Riley Cupauia fulvida Tr. and Planch.

Didymopauax morolotoni(Aubl.) Dec. and Planch .

Ficus citrifolia P. Mill. (as F. staud/eymia Dugand) Hasseltiaf/ oribuuda H .B.K. lserlia hae11keaua DC. Mico11ia argelllea (Sw.) D C. M. hyperprasiua Naud.

M. insufaris G Ieason

ill. impetio/aris (Sw.) DC. kf. pteropoda Bent h.

emergent trees:

Schult.

Tabebuia pentaphylla (L.) Hems!.

Zuelania guido11ia (Sw.) Britt. and Millsp. principal shrubs:

Ca/ea prunifolia H. B.K.

~ommon

179

D""ill" uspem (Aubl.) Benoist (as D . lucie/a Pres!)

vines:

Plrl')'gnuoc)'diacorymbosa (Vent.) K. Sch. Serj£mia ntmliueata Sauv. and Wright T~trm:era t•ulubilis L.

Amphilophitmt pauiculatum(L.) Kunth Anemopaegma urbimlalum (J:tcq.) DC. Armbidaea chim (Humb. and Bo n pl.) Veri.

[Kenoyer (1929), Barro

Color~do

lslan!l, Canal Zone]:

pio neer

forest trees: Didymoptmax morotoloni (Aubl.) D ec. and A11eiba tibuurbou A ubi.

Planch.

Bombacopsis quinata (Jacq.) Dugand (as B . . Heliocarpuspopayaue11 sis H.B. K. fendleri Seem.)

Luehea seemcmnii Tr. antll'htnch .

B. sessi!i.1· ( Benth.) Pitticr Micouia argeurea (Sw.) DC. Cecropia /augipes Pinier Oclii'0/1/a pymmida/e (Lam.) Urb. (as 0. limoC. ablusifolia Bcrtol. (as C. mexicaua Hems!.) nemis Rowlce) Cordia al/iodom (Ruiz a nd Pav.) Rocm. and Sclteclea gomphococca (Ma rt.) Burrell (as AtSchult. raleagomphococca Ma rt.)

Trema micralllha (L.) Blume

common vines:

Amphilophimn paniculaltm';'( L.) Kunth Auemopaegma orbicttlatum (Jacq.) DC. Bauhinia excisa (Griseb.) Hems!. (as B. rhomp.wnii I.M. Johnst.)

Chiococca alba (L.) 1-l itchc.

Cissus eros(l L. Rich.

C. rlwmbifolia Yah! Cliroria porrobellensis Buerling Cnestidiummfescens Planch.

Cunnams fJ"namensis G riscb.

Dm·illa aspcra (A ubi.) Benoist (as D. Iucida Pres!)

Dioclea altissima var. megacarpa (Ro lfe) Maxwell (as D. megacarpa Rolfe) D. rej/exa Hook .f.

Gouania polygama (Jacq.) Urb. 1/ipf!Ocrarca volubilis L fliraea rec/inara Jacq .

Maclwerium purptwascens Pittier Philodeudron erlansonii l. M. Johns!.

Plwyganocydia corymbosa (Vent.) K. Sch. Se1jauia alrolineata Sauv. and Wright S. nesiles LM. J ohns!.

Tetracera volubilis L.

[Johnston (1949), San J ose Island, Panama (as "Low Gray Thicket")) :

Cecropiapeltata L. (as C. araclmoidea Pittier) Cordia alliodora (Ruiz and Pav.) Roem. and

Lycoseris latifolia (Don) Uenth.

Callicarpa acwninala H .B.K. Cordia spi11esceus L. (as C. fermginea (Lam.)

[Standley ( 1928), Canal Zone]: land o n Atlantic side recently under cultivation, but now abando ned, trees and shrubs: Adeuariaf!oribmrda H.B.K. Ca/athea spp.

Cecropia sp p.

CIJ/wslcgia xalapensis (Bon pl.) D. Don Guazunra ulmifolia Lam.

Helicouia spp.

Hclicteres guazunwefolia H. B. K. Iudigofera sujji'uticosa Mill. f.Aulana spp.

thickets on low hills between Colon and Catival: Solanum hayesii Fernald Vismia spp.

Late Secondary Forest [Duke and Porter (1970), Darien): frequent canopy trees:

Cm•auillesia platallifolia (Humb. a nd Bonpl.)

H .B.K.

Ceiba pe111a11dra (L.) Gaerln.

Temtillalia amazonia (J. F. Gmel.) Exell locally dominant trees:

Trema micrautlui (L.) Blume frequent trees:

Roem. and Schult.)

Solanum exlensum Bitter

Casearia 11ilida (L.) Jacq.

Didymoparrax moroto/oui (Aubl.) Dec. and

secondary shrubs: Dalbergia brownei (Jacq.) Urb. Gynerium sagiuatum (Aubi.) Beauv.

Guazuma ulmifolia Lam. Heliocarpus popayauensis H.B.K. Luelrea :.pcciosa Willd.

Margarilaria nobilis L f.

Psidium guaja"a L.

Hibiscus tiliaceus L.

Ocltroma pyramidale (Lam.) Urb. (as 0. limoneu-l'is Rowlce) Piperaurittmt H.B.K. Se1jauia spp.

Trema micrarrtfra (L.) Blume Triumfctta spp.

Planch.

Vismia baccifera (L.) Tr. and Planch.

V. Ia!ifolia Cho isy Zalltlrox_vlwrr pauamense P. Wilson Z. se/ulosum P. WiiSO!l frequent shrubs:

Clibadiwn appressipilum S. F. Blake C. /eiocarpum Steetz.

Gonza!Cigwria rudis (Standi.) Standi.

Hamelia axil/oris Sw. Hasseltiafioribunda H .B.K. l sertia lraenkeana DC. umtaua camara L. Manihot esculellla Crantz Miconia /accra (Bonpl.) Naud.

Neurolaena looora (L.) R. Br. Piper aduncum L P. margiuatum Jacq.

180

D. M. PORTE R

Potlwmmplte pdtata (L .) Miq. Solanum rugusum D unal Tourm:fortiacuspidma 1-I.IJ. K. Triumfella /appula L.

l:upatorium bi//bergillllll/11 Lie uri.

E. iresim:uides 1-I.LI.K. E. /llti!TU[Ih)'ll/1111 L. E. micwstemuu C.tss.

E. odomtwn L. locally dominant banana-like or small E ..riuclairii 13cnth. palm-like pla nts: Gumnia cocciuea Cogn.

Hclicouia latispatlta Llcnth.

Mikaniaguaco Humb. and llo npl.

M.mit:rCIIIIhaH.l3.K. frequent banana-like or smajl palm-like · Rlty11tho.ria C11lycosa Standi. plants: S(tbicea vil/oso Rocm. and Schull.

Helicouia i111bricma (0. Kt zc.) Llaker H .mariae Hook. f. frequent herbs: }fyparrhenia rnf(• (Nccs) Stapf Asclepias cur(W'aviw L. Reuealmia cermw (Sw.) M acbr.

Baurreria laevis (Lam.) G riseb.

Cyperus diffitsus Vahl Dichromeua cilia/a Vahl frequent lianas : Byttueria acu/eata Jaeq.

Eclipltl alba (L.) Haask.

Eleusiuc indicct (L.) Gaertn.

Eleutheralllltera mderalis (Sw.) Seh. Bip. Mucuna bractea((l Dwyer Erigeron bouariensis L. P/u·ygcmocydia cory111bosa (Vent.) K. Sclt.

Panicum maximum Jacq.

Wulflia batCl/((1 (L. f.) 0. Ktze.

Paspaltmt plicatulum M ichx.

P. virga/11111 L. Prit•a lappulacea (L.) Pers. freq uen t vines: Cissu.1· rho111bi[olia Vahl Sclcria111elalcuca Schlecht. and Chanl.

C. sicyoides L. S. plerola Prest Desmodiu111 cauu111 (Gmel.) Schinz and Thellung [Johnston (1949), San Jose Island, Panama (as ' 'North Forest")]: principal canopy trees:

Andira inermis (Wright) D C. Byrsaninw crassifolia (L.) H .B.K. Cordia alliodom (Ruiz and Pav.) Roem. and Schult.

C. biro/or DC. Hirtcl/a americana L. Ilexguianensis(Aubl.) 0. Ktze.

Miconia argentea (Sw.) DC. Pem arborea Mulis Pltoebejo/mstonii C. K. Allen Pithcccllobiwu rufescens (Bent h.) Pittier Prolium /cnuifolillm ssp. mcleodii (I. M. Johnst.)

Porter (asP. mcleodii L M. Johnst.)

Temstroemia tepezapa/e Schlecht. an d Cham. (as T. semamtii Tr. and Planch.)

Xylopiafrutescens A ubi. secondary canppy trees:

Annona purpurea DC. A. spraguei Saff.

Bombacopsis sessilis (Benth.) Pittier (as Bombax sessile (Benth .) Bakh.)

Coussapoa panamensis Pittier Didymopanax moro/otoni (Aubi.)Dec. and Planch.

Ficas bullenei I. M. Johns!.

F. popcnoei Standi.

Ocotea rubrinerl'is Mez Spondias mombi11 L. understory trees and shrubs:

Amaio11a corymbosa H .B.K. Calyeo/pus warszewicziamts llenl,.

E11genia origanoides Berg Hirlella mcemosa Lam . (as H. oblongifolia D C.) Myrciaria jloribwufa (Willd.) Berg (as M. oucillii (Lundell) I. M. Johns!.)

Ouratea lncens (H.B.K.) Mart. (as 0. gaalemalensis Engl.) common vines:

Amphilophiam paniculatum (L.) Kunth Anemopaegma arbiculatum (Jacq.) DC. Arrabidaca pachycalyx Sprague Bauhinia excisa (Griseb .) Hems!. (as B. tlwmpsonii I. M . Johns!.)

Chiococca alba (L.) Hitchc.

Cissus rhombifo/ia Vahl Clitoria portobellensis Beurl.

Cnestidium panamensis G riseb.

Connarus rufescens Planch.

Davilla aspera (Aubl.) Benoist (as D. Iucida P rest) Dioc/ea rej/exa Hook. f.

THE VEGETATION OF PANAMA: A REVIEW

DoliowqJIIs den/1111/.1" (Aubl.) Standi.

Forl·temnia o•iridt•.\·cen.\· lllakc Gouania palygtmut (Jacq.) U rb.

Hiraea redinaw J:1cq . klaclwerinm purpm·a.I'I'CI/.1'Pitticr Philodendron erlcmsunii I. M . J ohnst.

P. gumemalense Engl.

lS I

Phr)'J:t/1/ocydia corymbo.m (Vent.) K . Sch. Securitlaca tlirer.•ifolia (L.) Ll l;~ ke Setjtmianesites I. M. Johnst.

Tetl'tlcera volubilis L. Tetmpteris tlh·colot· (Meyer) DC. · T . macromrpa I. M. Johnst.

[Johnston (1949), San Jose Island , Panama (as "South Forest")]: principal canopy t rees:

Andira iuennis(Wright) DC. Byrsonit;ta cra.uifolia (L) H .B.K. Cassipourea el/iptica (Sw.) Poir.

H irtella americana L /lex !llliauensis(Aubl. ) 0. Ktze.

Pem arburea M utis PhvcbejolmstouiiC. K. Allen · Pouleria calllpcdtiaua (H.B.K .) Bachni Protium teuuifolium ssp . mc/eodii (1. M. Johnst.)

Porler(as P. mc/eotlii I. M. Johns!.)

Roupa/a monlana A ubi. (as R. complicala H.LI.K.) Temstroemia seemmmiiTr. and Planch.

Xylopiafrutesceus A ubi. secondary canopy trees :

Bambacopsis sessilis (Bent h .) Pittier (as Bomba.\· sessile (llcnth.) Bakh.)

Cordia bicolor DC. Coussapoa panamensis Pittier Didymopauax 111orototoni (Aubl.) Dec. a n d Pla nch.

Ficus bullenei l. M. Jo hns!.

Maui/kara cltic/c (Pit tier) Gilly Micauia argenlea (Sw.) DC. Ocotea rubrinervis Mez

Zuelrmia guitlonia (Sw.) Brill. and Millsp. understory trc<.-s and shn1bs:

Anwioua corymbosa H.B.K. Calycolpus warszewiczimms Dcrg Eugenia ariganoides Berg Hirtella mcemosa Lam. (as H. ob/ongifolia DC.) il'ly•·ciaria flvribwula (Willd .) Berg (as M. oneil/ii (Lundell) l. M. J ohnst.)

Oumtea lucens (H.D.K.) Mart. (as 0. gnlltemalen.l·is Engl.) common vines:

Cltiococca alba (L.) Hilchc.

Cnestidium rufescem Planch.

Comtams panamensis Griseb .

Davilla aspera (Aubl.) 13enoist (as D. Iucida Pres I)

Doliocarpus delllatus (Aubl.) Standi.

Gouaniapo/ygm1w (Jacq.) Urb. lfi{Jpocratea t'olubilis L. Philodendron erlansonii I. M . Johnst.

P. gualemaleuse Engl.

Se('llridaca dio•ersifolia (L.) Blake Tetracera I'Oiubilis L.

[Kenoyer (1929), Barro Colorado Island, Canal Zone (as "Older Pioneer Forest" and "Mixed Secondary Forest")] trees and shrubs :

Castilla elastica Cerv. (as C. panamensis Cook) Coccoloba spp.

Ficusspp.

Gus/a via superba (H.B.K.) Berg Ingaspp.

lianas:

Bauhinia exci.ra (Griseb.) H em s!. climbing bamboos:

Artltrostylidium racemiflorum Steud.

Clmsquea simplicifolia Munro

Melastomataceae

0/media aspera Ruiz and Pav. Pal mae

Poulsenia anna/a (Miq.) Standi. (as lnophloem armatum (Miq.) Pittier Protium spp.

Sapium s pp.

Zanthoxylum spp:

plan ts of forest floor: Commelinaceae Filicinae M arantaccae Se/aginella spp.

Xiphidium caeruleum Aubl.

Z ingiberaceae

182

D. M. PORTER

THE VEGETATION OF PANAMA: A REVIEW

IS3

Littoral Forest

Conliaptmamellsis Riley

[Duke and Porter (1970). Darien ):

Eugenia origauoides Berg Gcnipa mucrica11a L .

Bromeli11 karatas L.

G11aZ1111ia lllmilola Lam. Guettarda odumta Lam. L11ehea candida (DC.) Mart. and Zucc.

Matlt)•ba f!/aberrima Radlk.

Plumcritl mbm L. Psidium guineensc Sw. Stemmadenia grandiflora (Jacq .) Miers Tabebuia pelltaplty/la ( L.) Hcmsl.

Cissus .•icyorifes L. Dim·lea altissima var. megacarpa (Rolfe) Maxwell (as D.megacarpa Rolfe) Marsdenia margaritaria R. C. Foster Odontocaryanitida Riley

locally do111inant 1rces:

Ena/lagmalaflfolia (Mill.) Small Hibisws tilit~ccus L. Tecoma stans (L.) H .B.K. Termiua/ia cawppa L. Xylo.mw pmwmeusis Turcz. frequent trees: JJucida bucera.1· L. Coccoloba uvifera (L.) L. Cocos nucifera L. Hippomnne ninucinelln L. Plumeria mbrn L. locally dominant shrubs:

Cluysoba/auus icaco L. Morindn pmromensis Seem.

l ttCtJIIinia sp. frequent lianas:

Combrelum decmulrum Jacq. loca lly dominant vines:

lt>omoea pex-caprae (L.) Roth.

Wetle/ia trilobata (L.) Hitchc.

I

·I

I

frequent vines:

Ctmavnlit1maritimn (A ubi.) Thouars Me/authera aspera (Jaeq .) L. Rich.

Vigna luteola (Jacq.) Bent h. locally dominant herbs:

S tachylllrphetajamaicensis (L.) Yahl

in shelter of thicket:

large vines:

[Standley (1928), C aribbean side of the Canal Zone: as "Strand Vegetation" ): shrubs:

herbs:

Caesalpinia crista L. Coccoloba uvifera (L.) L. Do/bergin ecastophy/la (L.) Tau b.

Omplwlen diandrtt L. Xy losma prmamensis Turcz.

Croton puncta/us Jacq.

Ky/lingia peruvitmn Lam. K.ptmgensVahl Pancratium litlom/e Jacq . (as Hyme11ocallis americana (L.) Stand i.)

Philoxerus vermiculat"is (L.) R. Br. Vigna h1teo/a (Jacq.) llcnth. (as V. repcns (L.)

frequent herbs: frequent s hrubs:

Capparis cynoplwlloplwra L. Chiococcn alba (L.) Hitchc.

Cenchru.r ecliinatlls L. Eragrostis amabilis (L.)"Wright and Arn. Philoxerus 1•ermicularis (L.) R. BJ-.

vines:

Cnllltl•alia maritima (A ubi.) T houars Ipomoea pes-caprac (L.) Roth.

O.Ktze.)

[Johnston (1949), San Jose Island, Panama (as "Beach Vegetation")): cha racteristic trees of sandy beaches: Cocos mtcifera L. Couocat·pus erecta L. Hippomane nwnciue/la L. much less common trees on upper parts of sandy beaches : Elaeodemlrou xylocarpum (Vent.) D C. Ficus trigona/a L. (as F.leaveusii I. M. Johnst.)

Lonchocarpus peutapltyl/us (Pa ir.) DC. Ximeuia americana L. most common herbs confined to stabilized beach sand: Canavalin maritima (Aubl.) Thouars (as C. rosea (Sw.) DC.) Ipomoea pes-cnprne (L.) Roth.

Pancratiumlittorale Jacq.

PhaseolusndennntlwsG. F. W. Mey. Uttiola pittieri Hack.

Vigna luteola (Jacq.) Benth.

dense thickety growth at head s of many sandy beaches, (!)principal shrubs: Dalberf{io brownei(Jacq.) Urb. Hibiscus tiliaceus L. Tecoma stan.< (L.) H.B.K. (2)vines and scramblers:

Caesalpinia crista L. (as C. bondnc (L.) Roxb.)

Entada gigas (L.) Fa we. and Rendle Heteropterys mncrostacltya Juss.

Mucuna mutisiatta (H.D.K.) DC. M. sloanei Fa we. and Rendle Omphalen diaudra L. (as 0. pauamemis (Beurl.

I. M . Jo hnst.) sheltered shingle beaches:

Avicenuia germinatts (L.) L. (as A.ttitida J acq.)

Cottocarpus erecta L. Lagunculnria racemosa (L.) Gaert n. f.

Sporobolus virginicus (L.) Kunth

[Johnston (1949), San Jose Island, Panama (as "East Harbor Thicket")): trees and shrubs:

Apei!Ja ti!Jourbou A ubi.

Aulomyrcia tomenlosa (Aubl.) Amsho ff (as Myrcia tomentosa (Aubl.) DC.) Bombacopsis quinata (Jacq.) D ugand (as Bombax· quinatum Jacq.)

Buncltosia cornifolia H .B. K. Bursera simaruba (L.) Sarg.

Cnesalpinia eriostac/rys Benth.

Casearia uitida (L.) Jacq. (as C. bauquitana Krause)

Cochlospermum vitifolium (Willd.) Spreng.

Thorn Forest [Duke and Porter (1970), Darien]: locally dominantlianas:

Caesalpinin crista L.

[Bennett (1968): as " Thorn-bush and Cactus"]: some of the many taxa present:

Act1cia spp.

Acamltocereus pemagonus (L.) Britt. and Rose Hamelin spp.

Oenocarpus spp.

Prosopis sp.

[Holdridge and Budowski (1956): as "Tropical Dry Forest", association on poorer soils] :

Acaciafamesiana (L.) Willd.

Prosopisjulijiora (Sw.) DC.

[Johnston (1949), San Jose Island, Panama: as "Rocks and Islets Close to Shore"]: trees:

Bombacopsis quina/a (Jacq.) Dugand (as Bombax quinatum Jacq.)

Bw·sera simarubn (L.) Sarg.

Coclt/ospermum vitifolium (Willd.) Spreng. most frequent shrubs:

Colen prunifolia H.B.KGuettarda odorata (L.) Lam. Plumeria rubra L. succulents:

Acanthocereus pentagonus (L.) Britt. and Rose (as Cereus pentngonus L.)

Agave panamena T rel.

Bromelia karatas L. Hylocereus polyrhizus (Weber) Brill. and Rose (as Cereus costaricensis (Weber) Berger) Opunlin elatior Mill. herbs:

Aeschynomene americana L. A_ brasiliaua (Poir.) DC. Andropogon angustatus (Presl) Steud: A_ brevifoliusSw. Anemia pastiuacaria Prantl

Til E VEGET ATION OF PANAM A: A REVIEW

D. M. PORTE R

I H4

Axolloprr.,· wpillal'is (Lam.) Chase 8/etio prrrpurc•a (Lam.) DC. Borrrrl'l'ia de11sij/om DC. B. lorijiJ/io (Aubl.) Schum. /J . ocymoides (Uurm .) DC. Bro.l·.,·m·olo 11odu.\'O(L.) Lincll.

Colapogo11ium tmtculloide., Dcsv.

Clwmaesycc ltypericifolia (L .) M illsp. (as E,pltorbio g/omifem (M illsp.) Wheeler) Clitoriombigino.1·o Juss.

Corrtorrbea.lpicata Aubl.

Cypems dif}ir.Wt.\' Vahl C./igrrlari,,· L. De.\'luodium barbatum (L.) Benth. and Ocrsl.

Diclrromena ciliara Vahl Dioscorea sapindoides Prcsl El•oh•ulusfilipe.\· Mart.

Ht1benaria panciflo•·(l (Lind I.) Rcichb.

Hypti.1· pu/egoiodes Bentli.

H .rectlrl•ata Poit .

Jacquemontia tattmifolia (L.) Griscb.

Kohlel'ia trrbif/ora (Cav.) Hemsl.

Mamula amndinocea L.

Mitm cor[/11.\ /n·c·l·ij/llmsGmy Op/ismcnu,, lmrmwmii (Rctz.) Ucauv.

J>onicum llwlleSw. Pectiselongoto H.II.K. Pepcromia pcllllcido (l.} l·l.U.K. Plwscol11s lollgipedmu·nlttttls Marl.

Phyllcmtlws stipulotus (Ra L) Webster (as P. . dijJII.\'1/S Kl.) Polygololonxicoulis H.O.K .

Polypodimn kulmii Fourn .

Prerolcpis pumi/11 (flonpl.) Cogn.

Rhyuchmpora micmmlw Va hl Sau1•agesia pulche/la Sc~m .

S. tcue/la Lam. Schll'ackaea cuplwiode.1· (Ocnth.) Durand Scleria intermpra L. Rich.

S. mict·ococc(t (Liebm.) Stcud.

Setaria tenax (L. R ich.) Dcsv.

Sit/a linifo/ia Cav. Spigelia tmthelmia L. Spiracamlw comifolia H.B.K. Sporobolus ci/iatn.r PresI Walt heria indica L. (as· W. americana L.)

large trees usually pr~scnt also :

o rher lrccs present in varying abundance:

' numcinel/a L. Hippoiumre Pisonia aculeata L. Prosopis julif/ora (Sw.) DC. (as P. chi/eusis

shrubs and small trees:

Acacia t'O.\'turicensis Schenck Cacraceae

Celtis iguanaeus (J acq.) Sarg.

Er;•throxylou me.ricauwu H.B.K.

(Mol.) Stu ntz)

Deciduous Seasonal Forest [Holdridge and Budowski (1 956): as "Tropica l Dry Forest", association on good soils]: characteristic trees: Acromia patwmeusis Bailey (as A. sc!CI'ocmpa Marl.)

Albizzia caribaea (Urb.) Brill. and Rose A. guachapcle (H.B.K.) Dugand (as Pseudosauumea guacltapele (H.B.K .) Harms) Apeiba tibourbou A ubi.

Bombacopsisquinai a (Jacq.) D ugand Bume/ia sp. {?)

Bursera simaruba (L.) Sa rg.

Calycophyllum candidissinwm (Yah!) DC. Ceiba pemandra (L.} Gaerln.

Cltloropl10ra tirrctoria (L.) G aud.

Cocltlospemwm vitifolium (Willd.) Spreng.

vines:

Caescdpiuiu crios/achys Bc nrh.

CochlosftCI'IIWIIIl'itifuliwu (Willd.) Spreng.

Guazuma ulmifo/ia L. Luelteo candida (DC.) Mart. and Zucc.

Cis.ws siqoides L. most conspkuous low plant:

JJw me/ia kamtas L.

o ther woody planls frequenlly fo und:

herbs

Al/ophyllus occideutali.l' (Sw.) Radlk.

Ca.l'earia 11itidt1 (L.) Jacq. (as C. bauquitmw Krause)

Cocco/oba corouara Jacq. (as C. waittii l. M. Johnst.)

Desmodiumll/l.lfiiSii[oliwn (H.D .K.) DC. Jacquemontia ta11mi[olia (L.) G riscb.

Oplismena.\' brtrmmmii (Rctz.) Dcauv.

Selaginclla diffusa (Prcsl) Spring Sic/a linifolia Cav. Tri{>sacwnlmrceolatwn Fourn.

Walrlreria indica L. (as W. americana L.)

Savanna [Duke and Po rter ( 1970), Darien]: locally dominant lrces:

[Stan d ley ( 1928), rocks along I he Pacific shore, Panam{l : as ''Coastal T hicket"] :

Puuteria stipitata Cronq uis t Spmulitt.r JIIII'JIIIrea L. Su•tnmadcuiu gm udiflora (Jacq.) Miers

Bur.Hnt.riuwrulm (L.) Sarg.

Spouclia., mombiu L.

Guettardaodomta Lam. Matayba glaberrima Radlk.

frequent banana-like or small palm-like plants: Hypt~rrhenia rufa (Nccs) Stapf

Anacarditwt occidentale L. Xylo{Jio aromuticu ( Lam.) Eichl.

X,fmresccu.\· A ubi.

freq uent herbs: frequent trees:

Pmticummaximum Jacq.

Byrsauinw cra.uifolia (L.) H.B. K. Psidium guajm•a L. [Myers (1969), Santiago Plain, Veraguas: as "Cu/'({fella Savannas"]:

Cumtel/a americana L.

Didymopauax morolotoni (Aubl.) Dec. and [Bennett (1968): as " Grassy Plains"]:

Planeh.

Diphysa robiuioides Benth. and Ocrst.

Enterolobitmt cyclocarpum (Jacq.) Griseb.

Luehea carrdida (DC.) Mart. and Zucc. (?)

Pitltecellobium samau (Jacq.) Benth.

Platymiscium polystacltyum Seem. {As P. pimratum (Jacq.) D ugand) Prosopisjulif/ora (Sw.) DC. Sciadodendron excelsum Griseb.

Simarouba glauco D C. Sterculia opera/a (Jacq.) Karst.

S weetia panamensis Benth.

Tabebuia chrysamha (Jacq.) Nicholson Xylopia frutesceus A ubi.

most frequent shrubs:·

Anacardittm occidentale L.

Byr.wuima crassifolia (L.) H.B.K. Cumtella americana L.

[Bennctt(l 968}: as " Low T rees and/or Oush"] : common shrub of drier phase:

Cumtella americana L. o ther c haracle ristic taxa :

Anacardirmt occiderrtale L. Byrsonima crassifo/ia (L.) H.B.K.

Nectandra s pp.

Xylopia spp. abundant and frequent associate of shrubs:

Hyparrhenia rufa (Nees) Stapf

[Holdridge a nd Budowski (1956): as " Tropical Dry F orest", association on poorer soils]: [Johnston (1949), San Jose Island, Panama: as " Deciduous Forest" ]: fire-resistant species : common canopy trees: Cedrela augustifolia DC. (as C.fissilis Veil.)

185

occasional canopy trees : Bombacopsis quina/a (Jacq.) Dugand (as Bombax quinatum Jacq.)

Curatella americana L.

Xylopia.frutescens Aubl.

186

D. M.I'ORTER

T HE VEGETATIO N O F PANAMA : A REVIEW

[Jo hnston (1 949): S<1n rose Isla nd, Pan:una: as " Herbaceous Flora at East Harbor"]: trees :

Curatel/a tmu:riCll/111 L. herbs:

Acisalltilera quadmta Pcrs.

Aeschynome11e amcricmw L. A. brasifit11w (Poir.) DC. Anemia pasli11acaria Prantl Audropogo11 llllgustatlls (Prest) Stcud.

A. brevifolius Sw. Arislidujomllensis Kunth A. lemipesCav. Bletia purpurea (Lam.) DC. Bouneria dc11sif/ora DC. B.latifolill (A ubi.) Schum.

B. ocymoidcs (Burm.) DC. Brtm avola 11odosa ( L.)'Lindl.

Bt~clmera sp.

Calopogouitmtnwctmoides Dcsv.

Cassia diphylla L. Cli!Oria rubigi11osa J uss.

Couto11bea spicata Aubl.

Crotoutriuitatis Millsp.

Cypcms diffusus Yah !

C.ligularis L. C. tenerrimm· Pres!

Desmodium barbttlll/11 (L.) Bent h. and Ocrst.

Dichrome11a ciliata Yah! Dioscorea sapi11doides Pres!

Eleoclwris illlerstichla (Vahl) Roem. and Schult.

Evoll'ldus jilipes M art.

Fimbrislylis amwa (All.) Roem. and Schult.

Habcllariapaaci/lora (Lind!.) Reich b. f.

Hyptis pulegioides Bent h.

H. recurvata Poi!.

Jacquemoutia tamnifolia (L.) Griseb.

Saltu·ater S wamp Forest

Kollleria lubiflum (Cav.) Hans!.

Lep!Ochloaji/iformia (L'Im.) Ocauv.

Mamilla arwulinacea L. Mar>)'lliantlws dwmacdrys (Vahl) 0. Ktze.

Mimosa pu.>·i/la Bent h.

Mitracm]ms brevif/orlls Gray Op/ismemts bamumnii (Rctz.) Beauv.

PauiCilllllllOIIe Sw. Pttspa/am clavalifemm Wright Pectis elougara H .B.K. Pcntarrlwphis scabra 1-I.B.K. Phaseo/us /ougipedancu/alus Mart.

P. ste11olobas Standi. l'hyllautlms caro/iuensis Wa lt.

P. stipu/atus (Raf.) Webster (as P. dif}imt;· Kl.) Po/yga/a bryzoides St.-H i!.

P.laugicaulis H.O.K. Pterolepis pumila (Bonpl.) Cogn.

Rhynclwspora armcrioides Pres!

R. mexicaua Steud.

R. micrautha Vahl Rolula mexicana C hmn. and Schlecht.

Sauvagesia pulchella Seem.

S. tc1tella Lam. Sc/wllesia gaianensis (A ubi.) Mal me Schwackea cupheiodes (Benth.) Durand Scleria illlerrupla L . Rich.

S. micrococca (Liebm.) Stcud.

S . setacea Poir.

Setaria /enax (L. Rich.) Dcsv.

Sida liuifolia Cav. Spigelia anthelmia L. Spiractmllw comifo/ia H .B.K. Sporobolas ci/iatas Pres!

Wallheria indica L. (as W. americana L .)

[Duke and Po rter (1970), Darien]: locally dominant trees :

Lagu11cularia racemosa (L.) Gacrtn . f.

Pel/iceira rhizoplwrae Tr. and Planch. frequent trees:

Posoqaeria lalifo /ia (Rudge) Roem . and Schult.

Curate/Ia americana L. Psidiam salutare (H.B.K.) Berg (as P. oerste· deanum Berg) herbaceous plants:

Buclmera spp.

Cassia tagera L.

Tabebuia palnm·is Hems!. locally dominant ba nana-like or small palm-li ke plants: Baclris nwjor Jacq.

Rltizop/wra brc>·islyla Salvoza

[Standley (1928), Pacific side of Canal Zone : as "Mangrove Swamps"]:

Pelliceira rltizoplwraeTr. and Planch.

[Standley (1 928), Pacific side of Canal Z one : as "Saline Flats"]:

Sesul'ium portulacastmm (L.) L.

Sporobolus >•ilgiuicus(L.) K un th

Saltwater R iparian Forest [D uke and Porter (1970), Darien]: locally do minant canopy·trecs :

locally don1i nant s hrubs :

Rltizoplwra breo•istyla Salvoza R. mangle L.

Rltizop/wra samocnsis ( Hochr.) Salvoza

locally dominant trees: Lagaucularia racemosa (L.) Gacrtn. f.

Conocarpas erecta L. Tabebaia pa/ustris Hems!.

frequent trees :

frequent shrubs:

freq uen t lianas:

Cilharexylam caadalum L. Plumeria rubra L.

Rhabdadeuia bi/lora (Jacq.) M uell.-Arg.

[Mayo Melendez ( 1965), D arien : as "Rltizophora brevistyla Salv. (Mangle Colorado) Asociaci6n"]:

Rltizopllora brevistyla Salvoza

Eupatorium amygdalinum Lam. Psidium sa/utare (H.B.K.) Berg (as P. oerstedeanum Berg)

Brackish S wamp Forest [Duke and Porter (1970), Darien]:

[Standley ( 1928) : hill at eastern end ofTaboga Island, Panama): stunted bushes on steep s lopes : Byrsonima crassifo/ia (L.) H.B.K.

frequent shrubs:

Conocarpus erecta L.

[Johnston (1949), San Jose fsla nd, Panama: as "Coastal Swamp Forests"]: flooded by sea wa ter at high tide: Pel/iceim rltizopltorae Tr. and Planch.

[Standley (1928), Ancon Hill, Canal Zone]: herbs and shrubs on steep grassy slopes : Curtia tenella (!Vfart.) Cham.

Dec/ieuxia mexicona DC.

187

Clitoria guyauensis (Aubi.) Benth.

Crota/aria pilosa Mill. (as C. ptewcaula Desv.)

Cyperaccae

Eupatorium amygda/inw11 Lam. £volvulus s pp.

Indigo/era /espedezioides H .B.K. Sida linifo/ia Cav. Tephrasia nitens Benth Zornia diphylla (L.) Pers.

locall y dominant canopy trees :

I

l

Mora oleifera (fr.) Ducke Rhizophora brel'istyla Salvoza R. mangle L. locally dom inant trees:

A>•iccnnia bicolor Sta ndi.

A. gcrminans (L.) L. Laguucalaria racemosa (L.) G aertn. f.

\

locally dominant shrubs :

Molllrichardia m·borescens (L.) Schott

Rhizophora samoensis ( Hochr.) Salvoza frequent shrubs :

Conocarpus erecta L. locally dominant palm-like plants Bactris major Jacq.

banana-like or

locally dominant herbs:

Philoxerus vemlicularis (L .) R . Br.

small

D. M. I'ORTER

THE VEGETATION OF I'ANAMA: A REVIEW

[Mayo Melendez ( 1965), Darien: as •· A Pict'lmia gl'l'miurm.,· L. (Mangle Negro) Asociacion ''I :

-Anosticlwm aarcul/t L. A ••iceunitt germinal/.\' (L.) L.

Graminac Hdicouia spp.

lm:nlly do minant canopy trees:

Momriclull'lfia urhoresccn.v (l.) Schott

(Mayo Melendez (1965), Darien: as " Mora olcifera (Tr.) Duckc (Aicornoquc) Asociaci6n"]:

Aslrocaryum slaudlqmunl/ Bailey Campa guiaueusis A ubi.

Diejfeubacltia sp.

Graminac

Mora ofeiferu (Tr.) Duckc

Brackish Riparian Forest [Duke and Por ter ( 1970), Darien) :

Rltizopfwra bre•·istyla Salvoza

(Mayo Melendez ( 1965), Darien: as "Momriclwrdiaurbore.>cenv Asociacion"]:

Pacltim rtlftWiim A ubi.

Posoqueria fati/olia (Rudge) Rocm. and Schu ll.

PriOt·ia copai/em G riscb.

Pterocarpu.> o}/iciualis Jacq.

[Holdridge and Budowski {1956) : as "Mangrove Swam1>s", Bocas del Toro): swamp fringe:

Rftizopftora mangle L. area toward dry land:

A t•iceuuia germiuaus (L.) L.

frequent sh rubs: loca lly dom inant trees: Laguucularia racenu1.1a (L.) Gacrtn. f.

Rapftia /aedigera Mart.

Rftizop!wra ma11gle L.

[Johnston (1949), San Jose Island, Pananui: as "Coasta l Swamp Forests"]: d omi nants where nooding with sea-w<11er is shallow or infrequent: A••icemtia bicofor Standi.

A. germiltam·(L.) L. (as A. nitida Jacq.)

Laguncufaria racemosa (L.) Gaertn. f. areas Hooded by brackish water: 1'Vlora oleifcra (Tr.) D ucke

Montriclwrdia arborcsccus (L.) Schott Pterocarpu.1· officina/is Jacq. frequent in shallowly flooded brackish swamp soi Is: Acrosticlumt aureum L. Euallagma fat!f'olill (L.) Small Rlwbdadenio bi}lora (Jacq.) Mueii.-Arg. Tabebuia palustris Hems!.

[Standley ( 1928), Canal Zone : as "Mangrove Thickets", Caribbean side]: trees :

Avice1mia germinotts (L .) L. (as A. nilida Jacq.)

Conocarpus erecta L. Laguncufaria mcemosa (L.) Gaertn. f.

Rftizophora sp.

shrubs and trees of lesser abundance: Cassipourea elliplica (Sw.) Poir.

Pat•onia spicata Cav. (as P. scabra (Vogel) Cifferri) Rhabdadenia biflora (Jacq.) Mueii.-Arg. (as R. paludosa (Yah!) Miers)

[Standley (1928), Cana l Zone: as "Mangrove Swamps", Pacific side]: marshes back of mangroves:

Acrostichum aureum L.

Cyperaceae Graminae

locally dominant banana-like or s111all palmlike l>lants: Gyuerium sagitlalwn (A ubi.) Bcauv.

Cil!ttm:x)'lwn caudatwu L. locally dominant shrubs:

frequent lianas:

Moulric!umlia arbore!>Cens (L.) Schott

Rfwbdttdenia bif/ora (Jacq.) Mucii.-Arg.

Freshwater .Marslt [Kenoyer (1929), GatLI!l Lake, Cana l Zone: as "Emerged Aquatic Association"] :

Acrostic!twn daucacfoliwn Langsd. and Fisch.

Crinum erubesceus A it.

Hibiscus somrius L. f.

Sagiluwia flmcifolia L. Typ!ta angusli/olia L.

(Kenoyer ( 1929), Gatun La ke, Canal Zone: as " Eroding Shores'']:

Begouia}ilipes Bent h.

Lycopodium cerm~11111 L.

Bryophyta

Musci

Gleic!teuia f/exuosa (Schrad.) Met!. (as Dicanop/erisflexuosa Schrad.)

Ko!tleria tubif/om (Cav.) Hanst.

P!tyffantlws uimri L. Pit)'rogr·muma ca/omelauos (L.) Link Tiboucftina lougifolia (Vahl) Bail I.

level swamp periodically flooded by fresh water:

Conocnrpus creela L. Coutarea ftexmulra (J acq.) Schum.

frequent trees :

Laguucnlariaracemosa (L.) Gacrtn. f. pure stands in periodically inundated tidal swamps, DariCn:

R11.\tia uccidemafis(Bcnth.) Hems!.

Tabefmia penlap!t)·lla (L.) Hems!.

[Holdridge and lludowski (1956): as "Aicornoq ue Swamp Forests'', Darien] :

Mort/ o/eifera (Tr.) Ducke

IS9

[Kenoyer ( 1929), Gatun Lake, Canal Zone: as "Floating Aquat ic Association"]:

Cham keuo)'eri Howe Eicft!tomia azurea (Sw.) KLmth (as Piaropus t1zureus (Sw.) Raf.) Ludwigia ltefmiut!torrftiza (Mart.) Hara (as Jus-

siaca untaus Hum b. and Bon pl.)

Nvtia stmtiotes L. Sal vinia auricula/a A ubi.

Utricularia 1ltixla Barn h.

[Kenoyer ( 1929), Gatun Lake, Canal Zone: as " Marsh Shrub Association"]: Dalbet~t:ia

ecastap!ty/fa (L.) Tau b.

Momriclwrdia arboresceus (L.) Schott

[Kenoyer {1929), Gatun Lake, C a nal Zone: as "Sedge Association"]:

Cypemsgigameus Vahl C. luzu/ae (L.) Retz.

Fuireua umbel/a/a Rottb.

Gynerium sagillalwtt (Aubl.) Beauv.

Hymeuac/me amp/exicaulis (Rudge) Nees Ludwigia oclova!vis ssp. sessilij/ora (Mich.) Raven (as Jussiaea stt}frulicosa L.) Nephrofepis biserrala (Sw.) Schott

Phragmites commwds Trin.

Rftyncftospora corymbosa (L.) B ritt.

Scirpus cubensis Kunth Scleria eggersiaua Boeck!. (as S. pa/udosa K uoth) Tfte/yp/eris gougy/odes (Schk.) Small (as Dryopleris IJOitgy/odes Schk.)

T. serra/a (Cav.) Alston (as Dryoptef'is serrata (Cav.) C. Chr.)

[Kenoyer (1929), Gatun Lake, Canal Zone: as "Stump Islands"):

Atrdropogon bicomis L. Boehmeria cy/indrica (L .) Sw.

Cfusiaspp.

EcfitJta alba (L.) Haask.

190

D. M. I'O RTER

FiCIIJSpp. Filicinac O rchiduceac

Typha ang11.1tijiJ/ia L. Vigna vexillllla (L.) A. Rich.

T I-l E VEGETA TIO N O f' PA N AMA: A REVi EW

PoiOquerialattfolia (Rudge) Roem . and Schult.

Quamribea astem/epi.1 l'illicr Q. bmcteolosa (Duckc) Cuatr.

Q.plerocalyx Hems!.

[Kenoyer (1929), Gatu n Lnkc, Canal Zone: as ··water-lily Association" ]: frequent shrubs:

Nymplwra amp/a (Salish.) DC. (as Castalia amp/a Salisb.)

Cassia reticulattt Willd.

[Standley (1928): Gatun Lake, Canal Zone]: emergent vegetation:

C/adi11m jamaicense Cra ntz (as Mariscus jamaiccnsis (Crantz) Britt.)

Cypcm sgigantcus Vah l Hibiscus somrius l . f.

Hydro/ca >11inosa L. (as Nama >1Jino;·a (L.) 0 . K tzc.)

Ludll'igia spp. (as Jussiaea spp.} Miumia argentea (Sw.) DC. Pluchca pttr{mrascens (Sw.) DC. Po/ygonum punctatw11 Ell. Potttederia cordata L .

Sagittaria lancifolia L. aquatic vegetation: Ceratopteris sp. £chi11odorus sp.

Eichhomia azurca (Sw.) K unth (asPiaropusazttreus (Sw.) Raf.) Le1111tlt cyclostasa (E ll.} Chcv.

Ludwigia he/mintlwrrhiza (Mart.} I-lara (as Jus.<iaea natans Hum b. and Bon pl.} Nympftaea sp. (as Castalia sp.)

NJ•mplwides lmmboldtia11a (H.B.K.) 0. Ktze.

Pistia slratiotes L.

Safl.iniasp.

Utricu/aria sp.

[Stand ley (1928): boggy spots on savanna d uring rainy season, Panama]: Anagallis pwnila Sw. (as Centunculus pentandrus R.Br.)

Bacopa spp.

Eriocaulon seemamtii Molden kc (as E. schiedea1111111 Koern.)

Grami nae

flysantftes inaequa/is (Walt.} Pennell Lophotocat1JIIs sp.

Marsileo sp. ivlayaca aubletii Michx.

Safl.iuiasp.

Sam•agesia pulchella Seem.

[Standley (1928): open boggy land ncar Matias Hernandez, Panama]: Aesclt)•nomelle ltispida Sw. A. scnsitil'a Sw.

Habe11aria sp.

-Koste/etzkya pe111asperma (DC.) G riseb. (as K. ;·agittata Pres!)

Me/ochiome/issifo/ia Bent h.

P!tylllmtlws stipulatw· (Raf.) Webster (as P. diffusus Kl.} Sclwltesia sp.

Sesbauia emerus (A ubi.) U rb.

[Standley (1928): treeless swamp near Pacific beyond Rio Tocumen, Panama!:

locally dominant banana-like or small palmlike plants: Bactris major lacq.

191

Corozo olcifera (H.B.K .) Bailey frequent banana-like or small palm-like plants: Rcnectlmia cemult (Sw.) Macbr. .

frequent herbs :

Limuoclwris /lava (L.) Ouch.

NJ•mpltacablluulaG. F. W. Mcy. Panicttm11wxiuuun Jacq.

(Mayo Melendez (1965), Darien as " Prioria copai[eri1 G ris. (Cativo) Asociaci6n" l : trees :

Astrocaryum stamlleyamtm Bailey.

Campa guiattettsis Aubl.

Mora o/eifera (Tr.) Duckc Pcttlaclethmmacroloba (Willd .) 0. Ktze.

Prioria copaifera Grise b.

Pterocarpus o}ficirwlis Jacq.

Tabebuia rosea (Bertol.) DC.

edged with dense shrubbery of:

Acrostic/111m aurc11m L. Dieffeubacltia sp p. . Enallagmalatifo/ia (Mill.} Small Graminae Helicottiu spp .

Mora oleifera (T r.) Duckc

[Holdridge and Dudowski (1956): as "Cativo F orests", Darien]:

Campa guianeusis A ubi.

Prioria copaifem G riscb.

Quararibea sp. -

[Holdridge and Budowski (1956) : as "Mauicaria Swamps", Bocas del T oro]:

Manicaria saccifera Gaertn.

[Holdridge and Budowski (1956) : as "Orey Swamp Forests", Bocas del Toro]:

Campnosperma pmwmeusis Standi.

Symp!tottia globulifera L . f.

[Holdridge and Budowski (1956): as "Pacltim aquatica Aubl.

Canuasp.

Pac!tira aqualica A ubi.

Cyperaceae Graminae Lud,..igia spp. (as Jussiaea spp.)

Thalia sp.

Pterocarpus officina/is Jacq.

~waonp

Forest", Rio Sambu, Darien]:

[Holdridge and Budowski (1956): as "Pterocarpus o.f/ici11olis J acq. Stands") :

[Standley (1928): as "Swamps amid Pacific Savannas", Panama]:

Freshwater Swamp Forest [Duke and Po rter (1 970), Darien]: locally dominan t canopy trees:

Pachira aquatica Aubl.

Prioria copaifera Griseb.

Pterocarpus ltayesii H ems!.

P. o.ffici11alis Jacq.

Tabebuia peutaphylla (L .) H ems!. freq uent canopy trees: Copaifera aromarica Dwyer

C. pauamensis (Britt.) Standi.

locally dominant trees:

Browueopsis excelsa Pittier Erytltrina glauca Willd.

Rapltia taedigera Mart. frequent trees:

C/usiaspp.

more abundant and striking trees: Annona purpurea Dunal Byrsonima crassifolia (L.) H.B.K. Cassia moschata I-I.B.K. Cedre/a angustifolia DC. (as C.fissi/is Yell.)

Coccoloba caracasaua Meisn.

Enterolobium cyclocarpum (Jacq.) Griseb.

Erythrina glauca Willd. lfymenaea courbaril L

l ngasp.

Licania platypus ( Hems!.) F ritsch Luehea seemanniiTr. and Planch.

Nectandra globosa (A ubi.) Mez Pithecellobium saman (Jacq.) Bent h.

Pseudobombax septenatum (Jacq.) D uga nd (as Bombax barrigon (Seem.) Decne.)

Schee/ea zonensis Bailey (as Atta/ea gomp!tococca Mart.)

Sciadodendron excelsum Griscb.

Sloanea temijfora (DC.) Standi. (as S. quadriva!vis Seem.)

Sterculia apetala (Jacq .) Karst.

192

D. M. I'ORTER THE VEGETATIO N OF PANAMA: A REVIEW

Bauhiniu sp.

shrubs o r small trees: Xylopia spp.

Vi.l'miaspp.

[Standley ( 1928): as "Wooded Swamps", Caribbean side of Canal Zone]: trees:

Heli£wria sp p.

Murantaccac

Manicaria spp.

Prioria copaifem Griscb.

Rubiaceac Zingiberaceac

shrubs :

Broomea macropltylla Linden [Sta ndley (1928) : as "Wooded Swamps", cast of Rio Tapia, Pana ma]: plen tiful t rees:

Zantlroxylam sp .

Cedre/a arrgrt.\'tifuJia D C. (as C.jissi/is Yell.)

Guarea guidonia (L.) Sleumer (as G. guara (Jacq.) P. Wilson)

frcquc11tlianas:

frequent vines :

Eupatorium iresineoides H.B.K. E. macrophyllum L. E. micro.rremon Cass.

E. odoratrrm L. Garwria coci::irwa Cogn.

Mikania guaco 1-lumb. and Bon pl.

Momordica c/wrantia L.

f rcq uent herbs:

B/el'iwm pmwmensis Lindau Coix lac/uyma,jobi L. Eclrinoclrloa colotumr (L.) Link.

Eclipta alba (L.) Haask.

Justicia coma/a (L.) Lam. J."pectom/is J acq .

Ludu•igia spp.

Odontonema longifolium (Ocrsl.) 0. Ktzc.

Paniwmmaximum J acq.

Prio•a lappulacea (L.) Pcrs.

Scoparia dulcis L. Tiboachina /ongifolia (Yah!) Bail!.

[llennctt (1968): as "Galeria forest"]:

Armbidaea pac/rycalyx Sprague

Lajimrsia punicifulia DC.

Sabicea o•illosa Rocm. and Schull.

Byt/1/eria aculeata Jatq.

Mucuna bmcteara Dwyer Phryganocydia ~orymbosa (Vent .) K. Sch. Strycluros ptliWmensis Seem.

Wu/jfra baccMa (L. f.) 0. Ktzc.

vines :

193

common taxa i ncludc: A1iacardium excelsum (Bertcro and Balbis) Skeels

Cecropia spp.

Ficus spp.

Freslnrater Riparian Forest .

[Holdridge and Budowski (1956): as "Tropical Dry Forest'' ]:

[Duke and Porter (1970), D arien]: locally dominant canopy trees : Prioria copaifem Griseb .

Pterocarpa>· officina/is Jacq.

Swartzia panamem·h llcnth.

Tabebaia pelllaphylla (L.) 1-lcmsl. frequent canopy trees: Ficas insipida Willd.

Pac/rim aqaatica A ubi. locally dominant trees:

Pithecellobiaror/ongifo/iam (Humb. and Bon pl.) Standi.

Raphia taedigera Mart. freq uent trees :

Astrocarywn standleyanam Bailey Cecropia /ongipes Pittier C. obta.I'Jfolia Ber tol.

C.peltata L. Clromelia spinosa Jacq.

Clusia spp.

Cocco/oba acuminata H.B.K. Geoffi·oea inermis Sw. Gustavia superba (H.B.K.) Berg.

Jnga puncta/a W illd.

Luehea seemanniiTr. and Planch.

Myrcia sp/endens (Sw.) DC. 0/media aspera Ruiz and Pav. Pentac/ethra macro/oba (Willd.) 0. Ktze.

Pithecel/obium rufescens (Benth.) Pittier Triclranthera gigantea (Hum b. and Bon pl.) Nees

Waruewiczia coccinea (Yah!) Klotzsh. locally dominant shrubs:

Lindenia rio•alis Bcnth. frcq uent shrubs :

Cassia reticu/ata Willd.

Chiococca alba (L.) 1-l itchc.

Clibadiwrrleiocarpwn Stectz.

Gonza/agwria mdis (Sta ndi.) Standi.

I-Im·seltia/loribwrda 1-l.B.K. I-fenriettclla fascicularis (Sw.) Tr. Lturtana camara L. Leamlm dic/rotoma (D. Don) C~gn.

Mabea occidenta/is Bent h.

Mimosapigra L. Ncurolaena Iobato (L.) R . Br. Odontonema strictum (Nees) 0. Ktze.

Palicourea guianensis A ubi.

P. triphyl/a DC. P:,yc/ootria lwrizontalis Sw. R icinm· communis L. Siparuna guianensis A ubi.

Triumfetta lappa/a L. locally dominant banana-like or small palmlike plants: Gynerium sagittatum (Aubl.) Beauv.

Phytelephas seemarmii Cook frequent banana-like or small palm-like plan ts: Re~~ealmia cernua (Sw.) Macbr.

along streams or on alluvial fiats next to rivers where water table is high:

Arwcardiam excelsrmr (Bertero and

Balbis)

Skeels

[Standley (1928), tributaries of Rio.Jlaraiso near East Paraiso between Summit and Pedro Miguel, Can a l Zone]:

Tristiclria trifaria (Willd.) Tul. (as T. hypnoides

in great abundance:

(St. H i!.) Spreng .)

Monsoon Forest fOuke and Porter (1970), Darien]: locally dominant canopy trees:

Anacardium exce/swn (Bertero and Balbis) Skeels Cal'anil/esia p/atanifo/ia (Humb. and Bonpl.)

H .B.K.

Ceiba pentaru/ra (L.) Gaertri. frequent canopy trees:

• Bombacopsis quina/a (Jacq.) Dugand B. sessi/is (Ben th.) Pittier Enterolobium cyclocarpum (Jacq.) Griseb.

Licania hypoleuca Benth.

Platypodium elegans Vogel Pseudobombax septenatum (Jacq.) Dugand Ste/'cu/ia ape tala (Jacq.) Karst.

Termina/ia amazonia (J. F. Gmel.) Exell Vitex cymosa Sp reng.

V. masoniana P ittier locally dominant trees:

Mouriri parvifolia Benth.

frequen t trees:

Allophyl/as occidentalis (Sw.) Radlk.

Alseis blackiana Hems!.

Amaioua corymbosa H.B.K. Asrrocaryam stand/eyam11n Bailey Calycoplryllum candidissimum (Vahl) DC. Clramaedorea cf. pacaya Oerst.

C. wend/andiana (Oerst.) Hems!.

Chione chambersii Dwyer and Hayden Clwmelia recordii Stand!.

C. spinosa Jacq.

Clusiasp.

Cnestidium rufescens Planch.

Coccoloba darienensis Howard Cordia al/iodora (Ruiz and Pav.) Roem. and Schult.

Ficus nymplraefo/ia L. Guareaguidonia (L.) Sleumer Gustavio nana Pittier G. superba (H.B.K.) Berg.

Heisteria /ongipes Standi.

194

D. M. PORTER

flirt diu ntccmosa Lam. Luchca secm{//miiTr. and Planch.

Jvfacrocncmum g/abrcscens (Bcnth.) Wcdd.

Mico11ia argeutca (Sw.) DC. M. burea/i~· G leason Myrcia splendens (Sw.) D C. Neca muplifulia D onn. Sm. Ouratea fucens (H.Il.K.) Engl.

Pithece/lobium rufesceus (Bent h.) Pitticr Pusoqueria lattfolia (Rudge) Rocm. and Schult.

Protium pauamense (Rose) I. M. Jo hns!.

Quararibett astcro/epis Pittier Q. bracteolosa (Ducke) Cuatr.

Randia armata (Sw.) DC. Saba/ a/leuii Bailey Schee/ia zuncnsis Bailey Siparww pauciflora (Beurl.) A. D C. Sorocea affinis HemsI. Stemmadeuia gmll(/if/ora (Jacq.) Miers Swartzia simplex (Sw.) Gacrtn.

Trophis racemusa (L.) U rb.

Xylopia aromatica (Lam.) Eichl.

X. bocatorcua Schcry X. fmtesceus A u bi. locally dominant shrubs:

Faramea occidentalis (L.) R ich .

Mabea occidentalis Bent h.

Piper pinoganense T rei. frequent sh rubs:

AcalnJha diversifo/ia J acq.

Alibertia edulis (L.Rich.) A. Rich.

Aphelaudra sinclairiana Nees C/avija mezii Pittier Faramea luteovirens Standi.

Hasseltiafloribunda H.B.K. Hebracautlms silvaticus Nees

1/emmia purpurco (Pittier) R. E. Schultes Leondra dichotumo (D. Don) Cogn.

Nu:a laetivirens Sl:mdl.

Palicoarea guiauensis A itbl.

P. triphylla DC. Pipt'r dariencnsc C. DC. 1'. reticuk1tlllll L. Psyclwtria capitattl Ruiz and Pav. Quassia W1111ra L. frequent banana-lik e or small palm-like plants: H e/iconia meta/fica Pl. and Lit1den H. vagina/is Benth .

Renealmia cenwa (Sw.) Macbr.

Xiphidium caemleum A ubi. frequent lianas:

Comwmspmw11te11sis Griseb.

C. wil/iamsii Britt.

Doliocarpus de11taius (Aubl.) Standi.

D. major Gmcl.

Atfaclracri1m1 capote D ugand Mal'ipa panamemis Hems!.

Passiflora vitifolia H.B.K .

Roureag/ahra H.B.K. Tetracera ••o/ubilis L. Wu/flia baccata (L. f.) 0. Ktze. frequent vines:

Plriladendron gurtiferum Kunth

T H E VEGET ATION OF PAN AMA: A REVIEW

Cavanillesia plalanifulia (Humb. and Bon pl.)

H.ll.K.

Ceiba penttmdm (L.) Gaertn.

Gttstavia supetba (H.B. K .) Berg.

Cm•atrillesia p/atanifo/ia (Humb. and Bonpl.)

H.B.K. [ Holdridge and Budowski ( 1956): as "Tropical Moist Forest'']:

Anacardirm 1excels1m1 (Bcrtero and Dalbis) Skeels Brosimwu spp.

Carapa guiauensis A ubi.

Cecropia obtusifo/ia Bertol.

Centrolobium paraeuse var. oreuocensc Bcnth. (as C. patinense Pittier?)

Coccolobatuercklreimii Donn . Sm. Couroupita dt1rienensis P ittier (as "C. panameusis") Dipteryx ptllwmem'is (Pitticr) Record

[Myers (1 969): as "Cava11illesia p/ata11ifolia Association", eastern Panama]:

Cava11illesia p/atanifolia (Humb. and Bonpl.)

H.B.K. [Myers (1969): as "Flat Lowland F orest", cutover forest about Garach ine, Darien]:

Cava11illesia p/atanifolia (Humb. and Bonpl.)

~tze.

secondary trees: all primary and secondary species of the "North Forest" ( = Late Secondary Forest)

disti nctive secondary trees: (Jacq.) P. Wilson)

[Mayo Melendez (!965), Darien: as "Bosque Seconda rio Antiguo"] :

Bombacopsis quinata (Jaeq.) Dugand Cavm1illesia platanifolia (Humb. and Bonp l.)

H.B.K.

Cedrela odorata L. Ceiba pentandra (L.) Gaertn.

Swietenia macroplrylla G. King

[ Mayo Melendez (1965), Darien as "Cavanil/esia-Bosque Mixto Asociaci6n "]:

A11acardiam exce/sum (Bertero and Balbis)Skeels

Astrocaryum stand/eyanum Bailey

understory trees and shrubs:

ArdiJ·iacompressa H.B.K. mostly those found also in other forests on the island lianas: Bartl1inia excisa (Griseb.) Hems!. (as B. tlrompsonii r. M. Johns!.) alii ianas oft he" North F orest "(= Late Secondary Forest)

[Johnston (I 949), San Jose Island, Panama: a s " Streamside Forests"] : common characteris t ic trees : A11acardium excelsum (Bertero and Balbis) Skeels Ficus citrifolia P. Mill. (as F. duga11dii Standi.)

F. insipida Willd. (as F. crii5siuscula Standi.)

F. trigo11ata L. (as F. campbellii I. M . Joh nst.)

Lica11ia platypus (Hems!.) Fri tsch Luel1ea seemam1ii Tr. and Planch.

Guar·ea guido11ia (L.) Sleu mer (as G. guara

H .B.K.

Gustavia sp.

Herrtmia pmpurrea (Pitticr) R. E. Schultes (as Tlreobromapurpureum Pit tier) lri(ll-tea exorrl1iza Mart.

Jacara11da copaia (Aubl.) D. Don Lecytlris sp.

Lue/rea seemmmiiTr. and Pia ncb.

Pentacletlrra macroloba (Willd.) 0. K tze.

Viro/asp p.

Warszewiczia cocci11ea (Vahl) Klotzsch.

[Johnston (1949), San Jose Island, Panama: as " Marino Forest"]: principal canopy tree: Tetragastris pa11amensis (Engl.) 0.

frequent epiphytes:

frequent herbs:

Quararibeu astcrolepis Pittier S11·m·tzia sp.

Ummopsis pillieri San·.

[Hold ridge and Budowski ( 1956): as " Transition Forests"]:

Tillandsia kege/ia11a Mez Justicia pectoralis Jacq.

195

G. multiflora A. Juss. (as G. culebrana C. DC.) Gustavia superba (H.B.K.) Berg lnga spurea Willd.

Mabea occidentalis Benth.

Macrocnemum glabrescens (Ben th.) Wed d.

Marila macroplryl/a Bent h.

Piperspp.

Triclrallllrera gigalllea var. guia11eusis Gleason common palms:

Bactris ba/anoidea (Oerst.) Wend!. common vines:

Dieffenbacl1ia sp.

Philodendron er/auso11ii I. M. Johns!.

P./rarlowiii. M . J ohnst. common terrestrial bromeliad:

Aeclm1ea magdafenae (Andre) Baker epiphytes : most orchids known from the island occur here

196

D. M.I'ORT ER

TH E VEG ETATION O F PANAMA: A R EVIEW

(Kenoyer ( 1929), 13arro Colorado Island, Canal Zone ): climax forest dominants: Bombacupsis quina/a (Jacq.) Dugand (as B.fendleri (Seem.) Piu icr) · Cuumarounalwnamcnsis Piuier Ficus insipida Willd. (as F. glabrata H.B.K.) Grias fendleri Seem.

Hum crepitans L. l riartea exhorrhiza Mart.

Jacarmula copaia (A ubi.) D. Don Platypodiwn elegans Vogel (as P. llwxonimwm l'illicr)

Prioria copaifera Grise b.

Spondias mombin L. Sterculia apetala (Jacq.) Ka rst.

Symplwnia glubulifem L. f.

Tabebuiaguayacan (Seem.) Hcmsl.

Myrsin aceae

T. pentaphylla (L.) Hcmsl.

Tenniua/ia amazonia (J . F. Gmcl.) Exell (as T. lwyesii !'ill ier) Viro/a sebifem A ubi. (as V. pauameusis (Hcmsl.)

Warb.)

[Standley ( 1928): narrow belt of alluvial land bordering Rio Tapia and Rio Tocumen, Panama]: shrubs:

trees:

Auactmlium excelsum (Oertero a nd Oalbis) Skeels Dipterodendron costariceuse Rad lk.

Pem arborea M utis Pithecellobimn longifolium (Humb. and Bonpl.)

Standi. (as P. valrlianum Bent h.)

Terminalia amazonia (J. F. Gmcl.) Exell (as T. lwyesii Pi!lier)

Sipanma paucij/ora ( B~u rl.) A. DC.

Apeiba tibourbou A ubi.

Cavanil/esia platauifo/ia (Humb. a nd Bonpl.)

muddy forest a long streams: magdale~we (Andre) Standi.

and Calderon)

sticky red clay soil overflo wed at h igh water: Dict)•oxiphilllll sp.

Dieffenbachia sp.

Tectaria sp.

Ficusspp.

Gliricidia sepium (Jacq.) Standi.

Raupa/a montana Au bl. (as R. daricnensis Pittier)

H.B.K.

Cecropia spp.

Cordia alliodora (Ruiz and Pav.) Roem. and

Triplaris americana L.

Schult.

-[Standley (1 928): "as wet forests of A tlantic Canal Zone"] : large trees :

Andira inermis(Sw.) H.B.K. Castilla elastica Cav. (as C. panamensis C ook) Chrysophyllwn spp.

Ficusspp.

Oleiocarpol! panamense (Pittier) Dwyer (as Coumarouna panamensis Pittier) Pou/senia armata (Miq.) Standi. (as lnoph/oeum armatum (Miq.) Pittier) Protium spp.

Virolaspp.

Zanthoxylum spp.

Bignoniaccae ep iphytes: Araceae Bromel iaccae Filicinae Ord1idaceac

EFergreen S easonal Forest [ Duke and Porter (1970): Darien ]: locally dom inant canopy trees : Alwrardilllll excelsum (Bcrtcro and Balbis) Skeels frequent canopy trees :

tier below tallest trees: Cyatheaceae

Didymapanax morolotoni (Aubl.) Dec. a nd

Stcmmadeniagmmliflora (Jacq.) Miers Warszcwiczia coccinea (Yah!) K lotzsch. locally domina nt shrubs:

Bombacop.tis quill(lfll (Jacq .) Dugand B. sessilis (Benth.) l'ittier Brosim11111 guia11ense (Aubl.) Huber Ceiba pentandra (L.) Gaertn.

Coclr/ospemurm williamsii Macbr.

Myroxylon balsamum var. pereire (Royle) Harms Olciocarpon palwmense (Pittier) Dwyer

Acchmea magdalenae (Andre) Ba ker (as Ananas

[Standley (1928): vestiges of natura l vegetation on low hills about Corozal, Paraiso, and Summit Canal Zone]: Bombacaceae

lianas:

l'emagoni11 spp.

I':.J•c/1otria spp.

Qua.uiam11am L. Smilaxspp.

Stryclr11os spp.

Zauthox)•lmu pmmmense 1'. Wi lson stream valleys: Araccae Cyathcaccae Danaea uodo.w (L.) J. E. Sm. Dieffeubachia ocrstedii Schott Gcsncriaccac Pa lmae Polypodiaccae Trichomanes diversifi·ons (Bo ry) Sadcb.

freq uent trees :

Clusiaspp.

Cono:."legiaxalapensis (Bon pl.) D. Don. Ficus nymphaefolia L. Gustal'ia;·uperba(H.B. K.) Berg Heistcria longipes Standi.

Luchea seemannii Tr. and Pla nch.

Miconia borealis GleasonOenocarpus ponwnen1·is Ba iley Ourtrtea lucens (H. B.K.) Engl.

Pentagonia bracl1yotis (Standi.) Standi.

P. macrophylla Bent h.

Pogonopus ;peciosus (Jacq.) Schlll\1.

Posoquerialat({o!ia (Rudge) Roein. and Schult.

Po1woumu scobina R . Ben. Sapindus saponaria L .

Siparrma paucij/ora (Beurl.) A. D C.

Mabea accident a/is Dent h. frequen t shru bs:

Cephaelis glomcrrrlara Donn. Sm. Clm•ija mezii PittierFaramca luteavirens Standi.

F. occidentalis (L.) Rich.

Hasseltia/lorilumda H.B.K. Herrania p1upurea (Pittier) R. E. Schultes Letmdra dichotoma (D. Don) Cogn.

M iconia oinocflropllylla Donn. Sm. Piper daricnense C. DC. P:.-yclrotria capita/a Ruizand Pav. Quassia amara L. frequent banana-like or small palm-like p lants: Costus nutans K. Sch. Heliconiametallica P l. and Linden Renealmia cemua (Sw.) Macbr.

Stromllllthe lutea (Jacq.) Eichl.

Xipllidium caemleum A ubi. freq uent lianas:

Passif/ara viti/olia H.B.K.

[Myers (1969), Cerro Sapo, Darien : as "Lowland H ill Forest (Evergreen Seasonal or Monsoon R ain Forest)"]: conspicuous understory dominant:

Phytelcphas sp.

Planch.

Gustavia superba (H. B.K.) Berg.

Premontane Rain Forest

Pal mae

[Duke and Porter (I 970), D arien]: thickets of shrubs and vines beneath trees:

Cestrum spp.

Herrania purpurea (Pittier) R. E. Schultes (as Theobromapwpureum Pittier) Melastomataceae

197

frequent canopy trees:

Brosimum guianense {Aubl.) Huber 0/eiocai}JOII panamcnse (Pittier) Dwyer

frequent trees :

Bro!Vnea rnacrophyl/a Linden Cespedesia macrophylla Seem.

Chome/ia spinosa J acq.

Clusia spp.

198

D. M. PORTER

Cono.1·te~:ia xalt~peusis (llonpl.)

D. Don

flaistel'ialou~:ipes Standi.

M iconia borealis G Ieason Oumtea lucem (H.B.K.) Cngl.

Pentagonia brachyotis (Standi.) Standi.

P.macrophylla Bent h.

Pogonopus >peciosus (Jacq.) Schum.

Po.<Otfueria latifolia (Rudge} Roem. and Schult.

Pourowna scobina R. Den. Sipt1rt111a pauciflora (Beurl.) A. DC. Stemmadenia gmudiflora (Jacq.) Miers Warszell'iczia cocciuea (Vahl) Klotzsch.

Xylopia fmtesceus A ubi. locally dominant shrubs:

Cephaelise/ata Sw. frequent shrubs:

Til E VEGETATION OF I'ANAMA : A REVIEW

Miwnia oinuchruphylla Donn . Sm. Palicourea guiauensis Attbl.

Picramnia dwyer! Porter (as 1'. antidesma Sw.) Piper durienense C. DC. P. marginatum Jacq.

P. reticulatw11 L. Psyclwtriacapitata Ruiz and l'av.

P. im•o/ucrata Sw. Quassia t/1//(/rtt L .

[Holdridge nnd Budowski (1956): as '·subtropical Wet Forest", Cerro Azul district beyond Loma l'clado, l'anam{l ]:

A/fiu·oa co.l'llll'icemi.\' Standi.

J'odocarpus guatemalensis var. al/enii

[Lewis (197 1)', Cerro Azul, Panam{t]:

Stemmadeuia allenii Woodson

frequent banana-like or sma ll palm-like plants: Costm· mt/111/S K. Sch. Renea/mia cermw (Sw.) Macbr.

Stromauthe lulea (Jacq.) Eichl.

Xip!tidium caemleum A ubi.

[Lewis ( 1971), Cerro Jcfe, Pana ma]:

Byrsonima dres.\'leri Lewis Cephaelis rigidifo/ia Dwyer and H ayden Chione boxifo/ia Dwyer and Hayden Dukea ptmame11sis Dwyer Faromeajefensis Dwyer and Hayden Hampea micrant/w A. Robyns Lisianthus jefeusis Elias Polyga/aje[e11sis Lewis Psychotria gra11dic{IJpa Dwyer and Hayden

Drymonia spectabilis (H.B.K.) Mart.

Passiflora l•itifo/ia H.B.K. frequent herbs:

P. olgae Dwyer ancl l-layclen Quarariben dolichopoda A. Robyns Riuorea dasyadena A. Robyns Rondeletia sa/icifo!ia Dwyer and Hayden Sclu-adera blumii Dwyer and Hayden Stelis/imbriata R. K. Baker Stemmadenia allenii Woodson Voch.1•sia jefensis A. Robyns

[Lewis ( 197 1), El Valle de Anton, Cocle]:

Coccocn;,·e/um herbac~um Lam. Spigelia anthe/mia L.

Cephaelis correae Dwyer and Hayden Hasseltia rigida A. Robyns Hibiscus cocleauus A. Robyns Hoj]ii/(IIIIJia capil/acea Dwyer

[Holdridge and Budowski ( 1956) : as "SubtropiCal M oist Forest", Chiriqui] : typified primarily by:

Quercus sp.

(St<~ nd l.)

Ouchholzand Gray

frequent lianas:

Aphe/audra incamata Leonard A. sinc/airiana N ees Clm•ija mezii Pittier Faramea lute01•ireus Standi.

F. occidenta/is (L.) Rich.

Hurania pwpurea (Pittier) R. E. Schultes

199

Oumtea coc/eensis Dwyer Po/yga/a 11'1/Nittckiaua Lewis Souroubea vollicola de Roon Stemmadenia al/euii Woodson

Myrciaspp.

Eugenia spp.

[Lewis (1971), Santa Rita Ridge, Colon]:

Calderone/la sylmtica Soderstrom and D ecker Cordia dwyeri Nowicke

[Holdridge and Dudowski ( 1956): as "Subtropical Wet Forest", Chiriqui]:

Bmuel/ia cos/al'icensis Standi.

Ca/ophyllnm bl'asi/ense var. l'ekoi Standi.

Juga spp .

Lap/aceafruticosa (Schrader) Kobuski La uraceac

Melastomataceae Ow·atea sp.

Rubiaceae

C. porcata Nowicke

Elfin Forest

Temstroemia tepezapote Schlecht. and C hum. (as Taonabo seemannii (Tr. and Planch.) Standi .)

[Duke and l'orter (1970), Darien]: locally dominant trees:

frequent epiphytes:

Clusiaspp.

Cloud Forest

P>:J•clwtria pithecobia Standi.

[Duke and Porter (1 970), Darien]: locally dominant trees:

Oenocal'pus panamiinus Bailey

Miconia oinochrophylla Donn. Sm. Piper margiuatum Jacq.

Psycho/ria valeriana Standi.

AcknoiVledgemcnts

frequen t trees:

Clusia spp.

Heisteria longipes Standi. lrim·tea cometo (Karst.) Wend!.

Ow·atea lucens (H.B.K.) Engl.

frequent banana-like or sma ll palm-like plants: Xiphidium caemleum A ubi.

frequent shrubs: Besleria pauciftora Rusby

Psychotria pithecobia Standi.

frequent epiphytes:

I

I

Various drafts of the text have been read by Dr. R. K. Baker, Dr. T. B. Croat, Dr. W. G. D'Arcy, Dr. J. A. Duke, Dr. J.D. Dwyer, and Dr. W. H. Lewis. Their many helpful criticisms are gratefully acknowledged. Not only has my wife Sarah read and improved the text, she also typed the manuscript and drew the maps that illustrate it.

As alway~, her assistance has been extremely valuable to me.

[Ho ldridge and Budowski (1956): as "Lower M ontane Forest", Chiriqui]:

Cedre/a tonduzii C. DC. Persea schiedeana Nees

Quercus copeyensis Muller Weinmannia pinnata L.

'It should be noted that the species listed by Lewis are endemic to the Cloud Forest and may not be either locally dominant or even frequent.

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200

D. M. PORTER

T H E VEGETATION OF PANAMA: A REVIEW

Rct:c.renccs

Sapper, K., 1900. Miuelamcrikanische Reisen 1111d Studie11 au.1· den Jahm1 1888 his 1900. Friedrich Vicwig, Braunschweig, 426 pp.

Sauer, C . 0., 1966. 711e Early Spanish Main. Univ. California Press, '13crkclcy/Los Angeles, Calif., 306 pp.

Schery, R . W., 1942. A few facts concerning the nora of r,; nama. Chron. Bot., 7:77-79.

Shelford, V. E., 1963. The Ecology of North America. Univ. Jllinois Press, Urbana, III., 610 pp.

Smith, A. C. and Johnston, 1. M., 1945. A phytogeographic sketch of Latin America. In: F. Verdoorn (Editor), Planls and Plm11 Scien ce in L atin America. C hronicn Botanica, Walth am.

Mass., pp. ll - 18.

Standley, P. C., 1928. Flora of tire Panama Canal Zone. Con/rib. U.S. Nal. Herb., 27:1-416.

Swan, L. C., 1966: The envi ronment of the Central American coldbloodcd vertebrate fauna. Copeia, 1966:684-699.

Trcjos, E. W. and A rcher, A., 1953. Atlas Eswdistico de Costa Rica. M in. Econ. Hacienda Dir. Gen. Estadis, Censor, San J ose, 114 pp.

T rewartha, G. T., 1961. The Earth's Problem Climates. Univ. Wisconsin Press, Mad ison, Wis., 334 pp.

Viksne, A., Liston, T . and Sapp, C . D., 1970. SLR reconnaissance of Panama. Phologramm. Eng. , 36 :253-259.

Wagner, P. L., 1964. Natural vegetation of Middle America. In: R. Wauchope and R. C. West (Editors), Handbook of Middle American Indians, /. Univ. Texas Press, Austin, Texas, pp.

216-264.

Wiley, S.C., Dodd, A. V. and Cha mbers, J. V., 1955. Environmental Handbook of Fori Sherman and Fori Gulick, Panama Canal Zone. U.S. Army, Headq. Quartermaster Res. Dev. Comm., Environ.

Protection Div., Tech. Rep., EP-17, Natick, Mass., 50 pp.

Bea rd , J . S., 1944. Climax vegetation in tropical America. Ecolog)•, 25:127-158.

Beard, J. S., 1953. The savanna vegetation of northern tropical America. £col. Monogr., 23:149-215.

Beard, J. S., 1955a. The classification of tropical American vegetation types. Ecology, 36:89- 100.

Beard , J. S., 1955b. A note on g allery forests. Ecology, 36: 339- 340.

Benne ll, C. F., Jr., 1963. A phytophysiognomic recon naissance of Barra Colorado Island, Canal Zone. Smiths. Misc. Coli., 145(7): 1- 8.

Bennett, C. F., Jr., 1968. Human influences on the zoogeography of Panama. lbero-Americtma, 51: 1- 112.

Bennett, H. 1-1., 1912. The agricullllral possibilities of the Canal Zone, I. Reconnaissance soil survey.

U.S. Dept . Agr., Rep., 95:5-38.

Benne tt, H . 1-1., 1929. Soil recon naissance of the Panama Canal Zone and contiguous territory. U.S. Dept. Agr., Tech. Bull., 94 :1 - 47.

Duke, J. A. and Porter, D. M., 1970. Darien Phytosociological Diclionary. Battelle Memorial Institute, Columbus, Ohio, 70 pp.

Erlanson, G. 0 ., 1946. The vegetation of San Jose Island, Republic of Panama. Smiths. ftilisc. Coli., 106(2):1-12.

Garver, R. D., 1947. Republic of Panama Report. Office of Foreign Agr., Rei., Washington, D. C., M~ Goldman, E. A., 1920. Mammals of Panama . Smilhs. Misc . Coli., 69(5):1~309.

Goldman, E. A. and Zetck, J., 1926. Panama. In: V. E. Shclford (Editor), Naltwalist's Guide to 1/te Americas. Williams and W ilkins, Baltimore, Md., pp.612- 622.

Galley, F. B., McGinnis, J. T., Clements, R. G., Child, G. I. and Duevcr, M. J., 1969. The structure of tropical forests in Panama and Columbia. Bioscimfe, 19: 693- 696.

Holdridge, L. R., 1957. The vegetation of mainland Middle America. Proc. Pac. Sci. Congr., 81/t, 4: 148- 161.

Holdridge, L. R. and Budowski, G., 1956. Report of an ecological survey of the Republic of Panama.

Caribb. For., 17:92- 110.

Holdridge, L. R. and Budowski, G., 1959. M!!Jla eco/Ogico de Panamti: l nstituto lnteramericano Ciencias, Agricolas, Turrialba, Costa Rica.

James, P. E., 1950. Latin America (revised edition). Odyssey Press, New York, N.Y., 848 pp.

Johnston, T. M., 1949. The botany of San J ose Island (Gulf of Panama). Sargelllia, 8:1-306.

Kenoyer, L. A., 1929. General and successional ecology of the lower tropical rain forest at Barra Colorado Island, Panama. Ecology, 10:201-222.

Lamb, F. B., 1953. T he forests of Darien, Panama. Cm,ibb. For., 14:128- 135.

Lauer, W., 1960. Probleme der Vegetationsgliederuhg auf der mittclamerikanischen Landbrucke.

Tagungsber. Wiss. Ab!t., Disch. Geogr., Berlin, 1959:128.

Lewis, W. H., 1971. High floristic endemism in low cloud forests of Panama. Biolropica, 3 : 78-80.

Martini, J . A., A h Chu, R. and Nel Lezcano, P., 1960. Forest soils of Da rien Province, Panama.

Trop. Woods,l12:28-39.

Mayo· Melendez, E.,l965. Algunas caracteristicas ecol6gicas de los bosques inundables de Darien, Panama, con miras a su posible utilizaci6n. Turrialba, 15(4):336-347.

Myers, C. W., 1969. The ecological geography of cloud forest in Pa nama. Am. Mus. No vitates, 2396: 1-52.

Panama Canal Company, 1967. Climatological Data, Canal Zone aud Panama, Amwa/-1967, 59 ( 1).

P anama Canal Company, Balboa Heights, Canal Zone, 49 pp.

Park, 0., Barden, A. and Williams, E., 1940. Studies in nocturnal ecology, 9. Further analysis of activity of Panama rain forest animals. Ecology, 21:122-1 34.

Platt, R. S., 1938. Items in the regional geography of Panama: with some comments on contem porary geographic method. Ann. Assoc. Am. Geogr., 28:28.

Porter, D. M., 1970. The rape of Panama. Mo. Bol. Gard. Bull., 58(1):1 1- 17.

Riitzler, K. and Steerer, W., 1970. Oil pollution: damage observed in tropical communities alon g the Atlantic seaboard of Panama. Bioscience, 20: 222-224.

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