Natural History and Ecology of suriname by LM Publishers

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Natural History and Ecology of Suriname

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Natural History and Ecology of Suriname

Editor Bart De Dijn, Environmental Services & Support NV www.ess-environment.com Graphic design Brigitte Küchler, René Bosshard, Aisthesis.ch aisthesis.ch

Production Hightrade bv, Zwolle Artwork on cover by / © Isabella Kirkland

ISBN 978-94-6022-438-6

This publication has been made possible with the support of: WWF Guianas The Embassy of the Kingdom of the Netherlands in Suriname

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Contents Preface

Acknowledgements

Introductory chapters General Introduction / History Bart De Dijn

Geology, Landforms and Soils Salomon Kroonenberg, Dirk Noordam

Climate and Hydrology

Riad Nurmohamed, Jacobus Groen, Sieuwnath Naipal

Biogeography

Brice Noonan

Ecosystems

Bart De Dijn, Pieter Teunissen, Jan Mol

Species account chapters Plants

Bruce Hoffman, Sofie Ruysschaert, Sabitrie Jairam-Doerga

100

Insects and their Relatives

Bart De Dijn, Hélène Hiwat

164

Freshwater Fishes

Jan Mol, Kenneth Wan Tong You

206

Amphibians and Reptiles

Paul Ouboter

256

Birds

Brian O’Shea

304

Mammals

Burton Lim, Marilyn Norconk

346

Final chapter Use, Threats and Conservation Chantal Landburg, Jan Mol

350 388

Glossary

414

Information sources / references

416

Index

467

Authors

479

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Preface

‘Natural History and Ecology of Suriname’ takes you on a journey through the natural world of what still is one of the greenest countries on earth. Indeed, Suriname maintains one of the highest forest covers of any nation worldwide, a treasure trove of natural richness. In addition, its savannah landscapes, swamps, rivers and mountains, all contribute to the colorful diversity of plant and animal species to be found. To a wide range of nature lovers, naturalists and professionals, this book aims at unraveling some of the secrets of Suriname’s natural world. As it reflects on the history and status of our nature to date, this book offers a reference for those who wish to know more about and explore Suriname’s natural world. Suriname’s unique and little explored natural world deserves more of international attention and recognition. This book offers readers a wide knowledge base to help motivate and implement nature conservation for the years to come and to increase the awareness and appreciation of the values of our natural world. WWFs’ mission is to create a future in which humans live in harmony with nature. Knowing our natural history through the information in this book, we at WWF Guianas as a part of WWF, can actively work towards that goal. We are grateful to Bart De Dijn and all contributors for all their hard work in producing this book. We want to thank the Embassy of the Kingdom of the Netherlands for providing the financial support to realize this unique publication. We know that you will enjoy it.

Laurens Gomes WWF Guianas Regional Representative

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Acknowledgements

he editor extends thanks to WWF, in the first place the former director of the Guianas office, Dominique Plouvier, who initated the project to make this book. Also thanked are all others at WWF Guianas involved in the project, especially the current director Laurens Gomes for continued support, and Mark Wright, Sofie Ruysschaert and Jerrel Pinas who supervised, helped and gave valuable inputs. Barney Jeffries is thanked for language-checking semi-final chapters (his advice we mostly took, but remaining errors in the final product are not Barney’s). The Government of The Netherlands is thanked for funding the entire project via the Royal Dutch Embassy in Paramaribo. It was implemented by a grant to WWF, which assigned project execution to Environmental Services & Support N.V. (ESS).

For their dedication and hard work, the authors are of course recognisized, as well as all who supplied images. Special thanks to Jan Mol, who not only wrote a chapter on his beloved fishes, but at the editor’s request also co-wrote two more chapters. Junior author Chantal Landburg is recognized for taking charge of writing the book’s closing chapter. Pieter Teunissen should probably get a statue for collaborating on this project while at some point facing life-threatening illnesses (and overcoming them). Brigitte Küchler and René Bosshard of Aisthesis spent many crazy hours on the layout of the book, and implementing final text corrections. Their hard work and flexibility was greatly appreciated. Peter Sanches and Ron Smit of LM Publishers are thanked for taking this project to the next level: final layout, printing and distribution. For help with tracking down artwork by Gerrit Schouten, John Henri Lance, Maria Sibylla Merian and Louise van Panhuys for the introductory chapter, Bart De Dijn wishes to thank Clazien Medendorp. She and Pieter Teunissen are also thanked for a quick review of the chapter. For access to artwork, thanks go to Charlotte Brooks of the Lindley Library of the Royal Horticultural Society in London, also Jip Binsbergen of the Artis Library in Amsterdam, and Cornelia Gilb of the Zentralbibliothek of the Johann Christian Senckenberg Museum in Frankfurt am Main. Susanne Kridlo and Johannes Lerp of the Museum Wiesbaden are thanked for allowing access to insect specimens likely collected by Maria Sibylla Merian. Many thanks to Guido and Rosine De Dijn for hospitality abroad, and Afi for patience and a lot of breaks. Salomon Kroonenberg and Dirk Noordam thank professor Theo Wong of the Anton de Kom University of Suriname for critically reading the geology part of this second chapter. Ton Markus of Kartomedia is thanked for professionally drawing two figures (Figures 6 and 15) for this chapter. Riad Nurmohamed, Koos Groen and Sieuwnath Naipal gratefully acknowledge the Meteorological Service Suriname, the Hydraulic Research Division, the Maritime Authority Suriname and the Suriname Water Supply Company for data provided. For support in his fieldwork in the region and study of the Guiana Shield, Brice Noonan would like to thank CNRS, NSF, the Centre for the Study of Biodiversity (Guyana), and the Nature Conservation Division (Suriname). Nathan Lujan, Brian O’Shea and Andrew Snyder kindly shared their thoughts and insights during the writing of the chapter on biogeography. Brice’s wife Danielle and two

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children (Alivia and Caden) are thanked for having kindly tolerated his absence during many extended trips to the Guianas. Bart De Dijn, Pieter Teunissen and Jan Mol wish to thank Salomon Kroonenberg, Dirk Noordam and Sofie Ruysschaert for helpful review of a draft version of the chapter on ecosystems. Dirk is also thanked for feedback on the deep history of Suriname’s landscapes and the landscape-soilvegetation relationship. Verginia Wortel and her employer CELOS are thanked for images of estuarine lakes. Bruce Hoffman, Sofie Ruysschaert and Sabitrie Jairam-Doerga thank the following collaborators who contributed significantly to the completion of the chapter on plants: Pieter and Siela Teunissen (photography), Chantal van den Bergh-Lodeweyckx (background research, photography), Eric Gouda (Vriesea splendens species account), Scott Mori (Lecythidaceae species accounts), Reinaldo Aguilar (photography; bejucososa.blogspot.com/), Olivier Gaubert (photography; floredeguyane. piwigo.com/), Omar Kasijo (contributing artist; Figures 55, 56, 58, 59), Sara Fuste and Brigitte Küchler (image editors). Credits for many additional contributers, no less appreciated, are provided in image captions. Bart De Dijn and Hélène Hiwat wish to thank those who contributed species accounts and photographs to the chapter on insects & relatives: Menno Reemer, Alies van Sauers-Muller, Hajo Gernaat, Simon Clavier, Atilano Contreras-Ramos, Nicolas Dedieu, Jocelia Grazia, Cristiano Schwertner, David Sillam-Dussès, Virginie Roy, Dominic Evangelista, Julio Rivera, Antonio Agudelo, Marcel Wasscher, Johan van’t Bosch, František Kovařík and Volker Mahnert. Also thanked are Anil Gangadin, Yanouk Epelboin, Auke Hielkema and Vanessa Kadosoe for providing additional photographs. Maurice Leponce is thanked for a quick review of part of the chapter. Paul Ouboter wishes to thank all scientists and technicians that assisted him in the field, especially Rawien Jairam, Vanessa Kadosoe, Gaitrie Satnarain and Gwen Landburg. Brian O’Shea wishes to thank Otte Ottema, Serano Ramcharan, and Arie Spaans for their contributions to the species accounts, and Sean Dilrosun for sharing his extensive knowledge of indigenous names and stories of birds in the Interior of Suriname. Johan Ingels and Thomas Valqui assisted in contacting some of the photographers who kindly shared images to illustrate the species accounts: Carlos Alcantara, Nick Athanas, Hervé Breton, Marc Chrétien, Foek Chin-Joe, Kester Clarke, Ian Davies, Mathias Fernandez, Harold Greeney, Audrey Hogenboom, Sean McCann, Leon Moore, Meshach Pierre, Jean-Louis Rousselle, Frédéric Royer and Lloyd Spitalnik. Burton Lim wishes to thank the Nature Conservation Division of the Suriname Forest Service for research and export permits. His fieldwork in Suriname has been funded and supported by Conservation International, SRK Consulting, ESS, University of Suriname, and Royal Ontario Museum Governors. Marilyn Norconk is grateful to the many students and colleagues in Suriname and from the US, UK and the Netherlands with whom she has worked at Raleighvallen-Voltzberg and at Brownsberg Nature Park. Chantal Landburg and Jan Mol wish to thank the Foundation for Forest Management and Production Control (SBB) for providing land-use and deforestation maps through their online portal; Sarah Crabbe of SBB customized these maps for the final chapter of the book. They also recognize Aad Versteeg of Stichting Surinaams Museum who made images available of prehistoric man-made raised fields near Galibi and mounds near Wageningen.

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General Introduction / History Bart De Dijn

his book is about natural history and ecology, a subject currently within the domain of the biological sciences. It is also about Suriname, located in the American tropics (Neotropics), and part of the larger Amazon region (Amazonia). So it may be fitting that an introductory chapter that offers a historical perspective is told by a biologist who has been living and working in Suriname for the last 27 years. This chapter reflects upon the study of the country’s natural history and ecology within the context of the ‘Western’ scientific tradition. This tradition is rooted in European culture, and is distinct from the Indigenous (‘Amerindian’), African, Hindustani (‘Indian’), Indonesian and other non-European traditions that are represented in Suriname. Yet it is informed by these traditions, often building on them: like Suriname itself, Western science is a bit of a melting pot, and no island. It is all about observing, collecting specimens, hypothesizing, testing, record keeping, and ultimately publishing findings based upon all this. The biological specimens and the scientific publications represent physical evidence that helps to reconstruct the history of the study of Suriname’s nature in the Western tradition. The evidence is presented below, in its larger historical context. Europeans came to the Americas as explorers from the late 16th till well into the 19th century. But exploring was more of a means than an end. The European goal at the time seems to have been to get richer and more powerful by any available means, be it trade and plunder, the appropriation of land and other resources, or the production of valuable commodities by a workforce of subjugated or even enslaved people (see for example Hemming 2009). Many practitioners of Western science at the very least piggybacked on what amounted to a sinister project, and the early stages of Western scientific study of Suriname are somewhat soiled by the link to this chapter in history.

Small steps, late 16th – early 18th century RENAISSANCE UPHEAVAL. What happened in Suriname from the late 16th till early 18th century should be understood in the context of the Renaissance. During the Renaissance there was upheaval in many aspects of society, such as on how knowledge was created and transmitted. Recent historical accounts of the Renaissance, such as by Cameron (2001) and Wiesner-Hanks (2006), explain this in some detail; below, a summary is provided based on these accounts. As the Renaissance proceeded from the 15th century onward, the full scope of the classical Greek and Roman arts and sciences was rediscovered in the Western European, Christian world. Europeans followed in the footsteps of Near-Eastern and North-African Muslim scholars. The latter had translated and drawn from the scientific works of, especially, the ancient Greek ‘natural philosophers’, and added new discoveries of their own to this body of work. The Western Europeans started to shift from scavenging the actual and proverbial ruins of classical civilization to wanting to fully understand, recreate and surpass the achievements of this civilization. Instead of uncritically accepting the dogmatic ‘truths’ found in the Bible and decreed by the Roman Catholic church, Western academics started to turn more to the empirical methods developed by ancient Greeks: observing the natural world, developing theories based on these observations, testing the theories in the real world, and thus making new discoveries.

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The Renaissance was also a time of great financial and technological development in Western Europe. The first European banks date back to this time, as well as the first personal firearms, ships better suited to open ocean travel, and advanced astrolabes to aid in navigation. This enabled voyages into the unknown and the conquest of new territories, such as Suriname. In addition, this was a time of religious upheaval in Western Europe. Various reformed and protestant churches and nations were established, including in the Low Countries and parts of what we now call Germany. The colonies of the Guianas, such as Suriname, were established by exponents of protestant European nations; the rest of South America was the dominion of Iberian Roman Catholics. When Roman Catholic rulers expelled the ‘heretic’ Jews from the Iberian Peninsula and the Portuguese colony of Brazil, they were accommodated in protestant enclaves, including in the New World. Another significant Renaissance development, given the importance of publishing in science, was the shift from manual copying of books to the technology to print books with text and blackand-white illustrations. This meant that ideas and knowledge could be shared faster, wider, and cheaper. Like translations of the Bible and protestant religious ideas, scientific ideas and accounts could thus reach a broader public, more widely sowing the seeds of change. THE NEW WORLD DISCOVERD AND MAPPED. The year 1492 AD marks the ‘discovery’ of the New World by Columbus during his first expedition to find a western passage to the spice islands of

Figure 1. Hondius’ 1599 ‘Guiana’ map showing the coastline and main rivers of the Guiana Shield, with the Suriname coast located above the leopard image (source: Bijzondere Collecties van de Universiteit van Amsterdam, UBM: Kaartenzl : 104.05.04).

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Southeast Asia (‘the Indies’). Maps produced during the Renaissance confirm that shortly after 1492 the coast of the Guianas was explored by Iberian mariners, subjects of the Spanish and Portuguese kings (see Helman 1982, Bakker et al. 1998). One of the captains in Columbus’ fleet during his second expedition to the New World in 1493 was Alonso de Ojeda. He sailed straight to the Guianas during a first expedition of his own in 1499, to explore and map the coast. Juan de la Cosa participated as a master navigator in that expedition, and made a crude map of the world (‘Mappa Mundi’) in 1500, on which the coastline of the Guianas can be recognized. These Iberian expeditions and the maps resulting from them were treated as state secrets. An example is the secret 1529 world map (the ‘Padrón Real’) of Diogo Ribeiro which shows 13 rivers between the ‘Rio Dulce’ (clearly the Essequibo), east of the island of Trinidad, and the ‘Marañon’ (the lower Amazon). No local names of these rivers are mentioned on this early map. A century later, after the Englishman Sir Walter Raleigh’s fleet explored the Orinoco and the coast of ‘Guiana’ in 1596, the secret was out. Based on this expedition, detailed maps were published in Dutch and other languages by Hondius and De Bry in 1599. Between the very recognizable Orinoco and Amazon, the maps show rivers with names quite similar to those on present-day maps of Suriname: ‘Coreteny’, ‘Mickhery’, ‘Copanamo’, ‘Saramo’, ‘Saranano’, ‘Comojowini’ and ‘Cimarawini’) (Figure 1). These maps were illustrated with an armadillo, jaguar, river turtle and peccary, alongside out-of-place illustrations of animals from the Old World. Apart from maps, narratives about the Amazon, Orinoco and the Guianas by for example Amerigo Vespucci (Markham 2010) and Walter Raleigh (Whitehead 1997) were published too. The narratives tell of mariners going upstream into the rivers and ashore, contacting the natives to extract information from them, as well as to acquire goods, including animals and plants. During the early 17th century, maritime merchants from protestant nations found the Guianas poorly defended by the Spanish and Portuguese, and thus available for first establishing trading posts, later colonies. The Dutch from Zeeland established trading posts at Cayenne and on the Corantijn (Helman 1982). They successfully established colonies along the Essequibo in 1616, and the Demerara in 1627; these were the ‘Dutch Guiana’ colonies, in present-day Guyana. The first successful colonization of Suriname wasn’t initiated until around 1650, by Englishmen hailing from Barbados, under the leadership of Francis Willoughby. They established sugar plantations, using the advanced ‘factory’ technology and practices – such as sugar mills and slave labor – that were introduced by displaced Jewish plantation owners from Brazil. This Jewish-protestant connection was a consequence of the religious persecution in Portuguese Brazil. In protestant territories like Suriname the authorities opportunistically welcomed the know-how and capital of the religious refugees. Initially, the colony in Suriname had just one small town, Torarica, some 40 km up the Suriname River. By 1665, a Jewish town, Jodensavanna, had developed some 10 km further upstream. In 1667, the thriving English colony was taken over by an expeditionary force from Dutch Zeeland. At the time, it already had 175 plantations along the Commewijne River, the lower Suriname River, and their tributaries (Figure 2). By 1686 a town called Paramaribo had become Suriname’s main settlement, located close to the mouth of the Suriname River, in the shadow of pre-existing fort Zeelandia (locally called foto, meaning fort; this word in present usage also refers to Suriname’s capital Paramaribo). Aside from the English interregnum of 1804-1816, the colony essentially remained under control of the Dutch until Suriname’s independence in 1975.

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Figure 2. Anonymous 1667 map of ‘Surranam’, present-day Suriname (source: https://coursewikis.fas.harvard.edu/aiu18/File:Surinam.jpg).

NATURALIA. The Renaissance’s focus on discovering and observing the natural world led to new developments in the arts and sciences – which, at that time, were not really separated. The realistic paintings of still life scenes became fashionable: vases with flowers, insects and other animals added as embellishments, as well as baskets with fruits, fishes and game catches. There was a definite taste for depicting the exotic, such as the New World. Drawings and paintings were commissioned by noblemen, as well as merchants, whose wealth and power had been increasing. The members of this elite acquired a variety of ‘naturalia’: rocks, minerals, fossils, dried plants, stuffed animals, and their depictions (van Gelder 1993, Swan 2002, 2004; Reitsma 2008, Davenne & Fleurent 2011). Besides these naturalia, exotic ethnographic objects were collected. They were combined into a ‘cabinet of curiosities’ or ‘cabinet of wonders’: typically a single room, or even a single piece of furniture, with all objects on display (Figure 3). The cabinets were mostly private spaces in which the rich and powerful could retreat and enhance their knowledge by actively studying the collection of objects or simply passively contemplating their esthetics and meaning. Some cabinets comprised considerable collections of exotic objects, and were famous and more public, such as the Rumphius cabinet. The fascination with naturalia and exotic places at the time led to the publication of catalogs (for example Rumphius 1705) and natural history books. Brazil’s environment, flora, fauna and people featured extensively for the first time in an illustrated book by Piso et al. (1648). Suriname and its flora and fauna were introduced to the world in a beautifully illustrated book by Maria Sibylla Merian (first edition 1705; second, posthumous expanded edition 1719) (see Box 1). Bart De Dijn

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Figure 3. ‘The Cabinet of a Collector’ painting of part of a cabinet of wonder, by Frans Francken the Younger, 1617 (source: Royal Collection Trust / © HM Queen Elizabeth II 2017, RCIN 405781).

Living exotic plants and animals were shipped to Europe and kept in-house or in private ‘gardens’ (precursors of public botanical gardens and zoos). Plants were of interest for use as ornamentals, food and medicines. Records indicate that seeds from Suriname were shipped to the Netherlands from the mid 17th century onward (van Andel et al. 2012c). At that time, several botanical gardens (‘hortus botanicus’) already existed, such as the one of the University of Leiden (established 1590) and the city of Amsterdam (established 1638). Early, published accounts of useful plants and animals from Suriname can be found in Merian’s work (see above and Box 1), and in early descriptions of the colony (van Berkel 1695, Harcourt 1707, Herlein 1718).

Adolescent leaps, mid 18th – late 19th century LINNAEUS’APOSTLES. From the 1730s onward, the Swedish naturalist Linnaeus named and classified naturalia (see Box 2). He described what we now consider biological species based on specimens obtained from curiosity cabinet owners, as well as traders and correspondents in distant territories. He inspired others to collect, name and classify species, including two botanists that went to Suriname but alas died not long after their arrival in the country (Lanjouw & Uittien 1935, van Andel 2010). He sent out 17 students – his ‘apostles’ – to the planet’s far corners to collect biological specimens (see www.ikfoundation.org/downloads/LACompleteCatalogueCopyrightIK2.pdf). Several of his apostles died on the road, or not long after returning to Sweden. Daniel Rolander was an apostle that survived. He was sent to Suriname in 1754 and left in 1763. Rolander collected mainly plants and insects in Suriname, where he was a guest of plantation owner Carl Gustav Dahlberg (van Andel et al. 2012a, de Moraes et al. 2014). Dahlberg was a friend of Linnaeus and sent him plant specimens for description, including the kwasibita plant (Quassia amara) with feversuppressing properties (Linnaeus 1763, 1775; see also Oudschants Dentz 1941, Dragtenstein 2004).

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BOX 1

Maria Sibylla Merian: naturalist-artist in a strange land

Maria Sibylla was the daughter of Matthäus Merian, a Swiss-German copper plate engraver and publisher. He was famous for his books with copper plate prints of landscapes (Wettengl 1998, Reitsma 2011). She followed in her father’s footsteps, making watercolors and engravings of flowers and insects. She developed an interest in the metamorphosis of butterflies, a novel subject at the time. Exploring nature, looking for caterpillars and their food plants, made Maria Sibylla an early ecologist. Maria Sibylla Merian married, but in the 1680s separated and left with her two daughters to join the Labadists, a reformed religious sect with a secluded congregation in the north of the Netherlands. This was located on lands of the van Sommelsdijck family. At the time, Cornelis van Aerssen van Sommelsdijck was co-owner and governor of Suriname. The Labadists also had a small beachhead in Suriname, on a plantation owned by governor van Sommelsdijk. Maria Sibylla and her daughters left the Labadist community to start an engraving and publishing business in Amsterdam in 1691. Yet the Labadist connection remained strong, at least via the oldest daughter’s husband Jacob Hendrik Herolt, a devout Labadist with business interests in Suriname. In 1699, Maria Sibylla traveled to Suriname with her youngest daughter. She stayed there at the Labadist outpost for less than two years. After returning to Europe, she published her observations and illustrations in a book on the insects and plants of Suriname. It was originally published in a limited edition in 1705 (Figure 4), but became a bestseller, well after Maria Sibylla’s death in 1717. Her hand-colored prints remain popular to this day, and are still of considerable scientific interest. Both daughters continued to make watercolors under their mother’s name after her death. Maria Sibylla’s elder daughter moved to Suriname in the 1710s with her husband; she was buried in Paramaribo in 1728. The younger daughter married and moved to St. Petersburg (Russia) with her husband, where she died in 1743. Figure 4. Watercolor of the butterfly Morpho deidamia (adult and caterpillar) on Suriname cherry, Eugenia surinamensis, by Maria Sibylla Merian, 1902-3 (source: Royal Collection Trust / © HM Queen Elizabeth II 2017, RCIN 921160).

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BOX 2

The Linnaean system for naming and classifying species

Carl Linnaeus (1707-1778) – the ‘Father of Taxonomy’ – formalized the system for naming and classifying species that we still use today. Linnaeus referred to what we now call species with universal two-part (‘binomial’) names. These were unique, short names that expressed key characteristics of the species. Latin, the universal language of science in Linnaeus’ time, was used to form the names. Before Linnaeus’ innovation, different names – in Latin or local languages – were often assigned to the same species by different authors, which caused much confusion. Some authors used descriptive sentences to name species; these were hard to memorize. The usefulness of Linnaeus’ innovation is evident from the fact that it has remained the basis for current scientific species names. The scientific name for the jaguar, for instance, is Panthera onca. It is written in italics in the modern literature. The first part of the name is the generic part, a single word with the first letter always capitalized in modern usage. This part is typically shared by several rather narrowly related species, which are then referred to as belonging to the same genus. It is also known as the genus name, and for example Panthera spp. is used as shorthand for all species of the genus Panthera. The second part of the name is the specific one, a single word that is never capitalized in modern usage. It makes a unique combination with the generic name, distinguishing, for example, Panthera onca from Panthera tigris (tiger) and Panthera leo (lion). In 1753 Linnaeus published the first edition of ‘Species Plantarum’, in which all plants known to him were named and classified. The classification was hierarchical, and based on structural characteristics, such as the position and shape of leaves, flowers and fruits. Earlier, plants were often classified based on their usage. Linnaeus assigned structurally very similar plants to the same genus. Genera (plural of genus) with broad similarities were grouped into families. Families were grouped into higher categories, such as orders. The same principles were applied to other naturalia. By 1758, the tenth, extended edition of his publication ‘Systema Naturae’ was printed, in which all naturalia known to him were classified, including animals. Linnaeus’ publications include descriptions of plants and insects that were based on the drawings and notes of Maria Sibylla Merian (1705). By the end of the 18th century, Linnaeus’ works had become authoritative, and the basis of universal scientific names. Many scientific species names that are still in use today can be traced back to Linnaeus’ 1753 and 1758 publications.

In the end, Linnaeus did not describe any of the plants Rolander brought back, because Rolander refused to hand over his collections, and tried to independently embark on an academic career. Rolander failed miserably at this, and ended up selling his collection to make ends meet; in this way at least some of the material he collected led to published descriptions of species (mainly by Rottbøll 1778). Rolander did process his notes and species descriptions into a manuscript that remained unpublished until it was translated 250 years later (Rolander 2008; also known as ‘Diarium Surinamicum’).

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NATURAL HISTORY BOOKS AND ILLUSTRATIONS GALORE. In the mid 18th century there seems to have been an increasing appetite in Europe for natural history accounts of exotic territories. Several books on the subject as it related to Suriname were published in a variety of languages: by Fermin in French (1765), by Bancroft (1769) in English, and by Hartsinck (1770) in Dutch. A late 18th century bestseller that included much information on the country`s natural history was John Gabriel Stedman’s 1790 ‘Narrative’ (reprinted repeatedly, such as in 1971). Stedman was a professional soldier, part of a contingent sent to Suriname to protect the colony against runaway slaves who had escaped from the plantations and taken refuge in the forest to establish independent Maroon communities there. Stedman’s narrative contains a lot of information on natural history, but its prime purpose was to expose the cruel treatment of slaves in Suriname. From the mid 18th till late 19th century many collecting activities took place in Suriname by amateur naturalists, often expatriates who were part of the country’s colonial elite. Although these amateurs did not publish their findings, they showed their scientific interest by sending specimens to museums in Europe for further study. Examples include Dahlberg (already mentioned above), Dieperink, Hering, Hostmann, Lammens, Kegel, Wullschlägel and Voltz (Teunissen 1979; see also Medendorp 1999). Definitely not part of the elite was August Kappler, a low-level public servant stationed at the periphery of the colony. He published a narrative (Kappler 1854), as well as an extensive description of Suriname and its natural history (Kappler 1887). The English interregnum (1804-1816) seems to have been a period of great synergy for the production of natural history illustrations in Suriname. At least four people were involved at the time: Adriaan François Lammens, a Dutch judge stationed in Paramaribo, John Lance, his English counterpart in the mixed Dutch-English court at Paramaribo, Louise van Panhuys, the wife of a plantation owner that was also briefly governor of Suriname, and Gerrit Schouten, a local artist (Burkhardt et al. 1991, Medendorp 1999, 2008) (see Box 3). Subsequently, some naturalists with more professional ambitions were active in Suriname. One was Hendrik Charles Focke; born in Suriname, he had studied law in The Netherlands, and was undoubtedly part of Paramaribo’s elite. He published several works on the orchids of Suriname (such as Focke 1849, 1855). Another publishing amateur botanist in Suriname was the Dutchman Splitgerber (for example 1940, 1942).

Mature strides, early 20th century onward RECENT HISTORY. It is challenging to present the highlights of natural history studies in Suriname from the early 20th century onward. So much biological exploration and research activities took place in this period, and relatively little time has passed to digest what has transpired. Below an overview is provided of the kind of efforts that were undertaken, the key institutions and people involved, and the historical context. As in earlier times, scientific developments reflected the social and political ones. The historical backgrounds are outlined in what follows largely based on van Traa (1946), van Lier (1971), Helman (1982), Bakker et al. (1998), and the author’s personal observations in the period 1990 till present. Biological research in Suriname till the late 1970s has been reviewed by Teunissen (1979), botanical expeditions by van ’t Klooster et al. (2003). Below a limited number of references are mentioned; the other chapters in the present book contain more extensive references to recent research in the country by researchers mentioned ones.

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BOX 3

Gerrit Schouten, home-grown artistic genius

Gerrit Schouten was a multi-talented professional artist: skilled in taxidermy, drawing and the making of dioramas (Medendorp 1999, 2008). He was a mixed-race Surinamer, and is assumed to have been largely self-taught. He likely absorbed the skills of people like Louise van Panhuys, who drew and painted plants and landscapes (Görner & Dobat 1991), and John Henri Lance, an amateur naturalist and draughtsman. Schouten sold his beautiful watercolors of plants and animals (Figure 5) to Lance and to the judge Adriaan François Lammens. These watercolors rival those of Maria Sibylla Merian.

Figure 5. Watercolor of the lizard Iguana iguana, with line drawing added to show details of the plates and scales on the head, by Gerrit Schouten, 1800-1839 (source: Artis Bibliotheek, Bijzondere Collecties van de Universiteit van Amsterdam, IZAA100171).

Schouten’s drawings are currently in the Surinaams Museum in Paramaribo, in the Artis Library in Amsterdam (recently integrated in the ‘Bijzondere Collecties’ of the University of Amsterdam), in the Naturalis Museum in Leiden, in the Lindley Library of the Royal Horticultural Society in London, and in the Fine Arts Museum in San Francisco. The Lindley Library has the largest known collection of Schouten’s watercolors, mixed with those of Lance in two large bound volumes. Many of Lance’s and Schouten’s drawings are near-identical in subject and composition, but not in quality: Schouten’s drawings were executed with superior artistic skill. They were intended for use by naturalists, as is obvious from the fact that no backgrounds were drawn, and that the subjects are positioned to show all essential anatomical features to identify the species depicted. Schouten’s most amazing artistic product can be seen in the Boerhaave Museum in Leiden: a box filled with Surinamese butterflies. They look like regular pinned butterflies, but they are entirely artificial. The likeness to real butterflies is uncanny. This appears to be the only surviving box of three that were made by Schouten (Clazien Medendorp, personal communication) (see ‘vlinderkast’ video at www.youtube.com/watch?v=WgqVia55CVg).

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A PERIOD OF CHANGE. In the early 20th century and the period leading up it, Suriname went through dramatic changes. While the Dutch banned the transatlantic slave trade in the early 19th century, slavery itself had not been abolished in Suriname. As the supply of slaves to the country gradually diminished, the management of plantations became difficult to impossible. Suriname’s economy was entirely based on these plantations. As July 1st 1863, the date of the abolition of slavery by the Dutch, approached, the pace of abandonment of plantations increased. On that date, only 163 of the 500 previously active plantations remained. This number decreased further to 45 at the start of the 20th century. Some freed slaves stayed on the plantations, but many went to the capital city to settle in a part of Paramaribo aptly called frimangron (literally: free man’s land). The colonial government managed to somewhat delay the demise of the plantations by recruiting an alternative workforce in the form of impoverished Dutch farmers and ‘East Indian’ laborers recruited in what is currently China, India and Indonesia. For a few decades, the economic decline in the colony due to failing plantations was partly compensated by development of the gold mining and balata plant gum extraction industries (see chapter on Use, Threats and Conservation). Gold production peaked around the turn of the 19th20th century, balata production some 20 years later. In the period 1901-1911, a railroad was built from Paramaribo deep into the Interior (Wicherts & Veltkamp 2012). The railroad track opened up previously inaccessible parts of the country, which led to social and economic development, as well as scientific exploration. The colonial government was eager to discover more potential sources of wealth in the Interior. It started a program to demarcate the southern borders of Suriname and map the geological riches in Suriname’s entire territory. From 1901 till 1911, the Royal Dutch Geographical Society (KNAG) organized a total of seven expeditions to the far reaches of all the main rivers of Suriname. Typically one of the participants in these expeditions was an amateur naturalist, often the medical doctor that was part of the team to keep it healthy (Pieter Teunissen, personal communication). Between 1920 and 1944 at least ten expeditions took place; professional biologists and foresters led these expeditions or participated in them, such as Stahel, Gonggrijp, Geijskes and Maguire. Zoologists and botanists, such as Pulle and Lanjouw, also undertook several collecting trips in Interior during this period. Pulle, a botanist at the herbarium of Utrecht University, initiated an important, high quality series of publications in 1928: the ‘Flora of Suriname’. Pulle’s initiative was continued by other Utrecht University botanists: Lanjouw, Stoffers and Lindeman. At present, the Flora of Suriname series encompasses over 2000 pages of species descriptions, including identification keys. During 19081910, two Surinamese brothers, Frederik Paul and Arthur Philip Penard, compiled ‘De Vogels van Guyana’, the first monograph on the birds of the three Guianas (present-day Guyana, Suriname and French Guiana). During this period, applied biological research was also undertaken in Suriname. Forestry research took place under the auspices of Gonggrijp. Research on agricultural pests was undertaken at the newly (1904) created Agricultural Experiment Station (Landbouwproefstation) in Paramaribo by such researchers as entomologist (insect specialist) Geijskes and botanist and mycologist (fungi specialist) Stahel. Research on human and livestock parasites was undertaken by researchers such as Flu and Frickers. Medical interest in the transmission of malaria led to a monograph on the mosquitos of Suriname by Bonne and Bonne-Wepster (1925).

Bart De Dijn

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SECOND WORLD WAR UNTIL INDEPENDENCE. The Second World War (WWII) created major economic opportunities for Suriname. Bauxite was used to make aluminum for the war industry in the USA, and Suriname was a major source of it. Airstrips were built as part of ‘Operation Grasshopper’ in remote corners of the Interior to facilitate mineral exploration. The bauxite industry in Suriname boomed and led to the construction of the Brokopondo hydropower reservoir in the 1960s in the Suriname river basin (see also chapters Climate and Hydrology, and Use, Threats and Conservation). A road was built all the way around the reservoir to Pokigron, on the middle Suriname River. Road development increased in the decades after WWII: an east-west road in the swampy Coastal Plain, one in the Zanderij Belt with its savannas and forests on white sand, and a number of connections between the two, as well as extensions to the south, into the densely forested Interior. The coastal road served to connect population centers and new agricultural projects, such as wet rice farming in the northwest of Suriname. The other roads provided access to timber and mining concessions, and allowed for the creation of palm oil plantations in parts of the Interior. This airstrip and road development opened up parts of Suriname previously inaccessible for researchers. It made biological exploration much easier than before, and offered opportunities for Dutch researchers, who were no longer welcome in the former colony ‘Nederlands Oost Indië’, which became independent shortly after WWII as the Republic of Indonesia. The work of Dutch researchers was mainly funded by the newly (1954) created research funding agency in The Netherlands, Wosuna, which became Wotro in 1961. Their publications were included in the Flora of Suriname series, in the newly created serial ‘Studies on the Fauna of Suriname and other Guyanas’ (started 1945), in pre-existing zoological serials of the Museum of Natural History in Leiden, and in a number of separately published studies. At that time, more than 30 extensive collecting trips were undertaken to many corners of Suriname by researchers such as the botanists and vegetation specialists Lindeman, Jonker, Kramer, Hekking, Wessels-Boer, Florschütz, Maas, Moolenaar, Schulz, Heyligers, Van Donselaar and Oldenburger, entomologist Geijskes, fish specialists Boeseman and Nijssen, amphibian and reptile specialist Hoogmoed, bird specialist Mees, and mammal specialist Husson. Important ecological research was also done in relation to the Brokopondo reservoir area, such as by Leentvaar on the aquatic environment (see Ouboter 1993), and by Genoways on small mammals. Spaans undertook ecological studies on shorebirds in the context of nature conservation. Publications of these researchers are referred to in other chapters of this book. Inventories of mollusks, such as snails and mussels, in the Estuarine Zone and elsewhere in Suriname were undertaken by van Regteren Altena (for example 1960, 1975). A number of publications deserve specific attention here, because they are still commonly used. Lindeman and Mennega (1963) published a Suriname tree guide; a fully revised, English-language edition has recently been published (Bhikhi et al. 2016). Mennega (1976) edited ‘Fa joe kan tak mi no moi’, a pocket flora of mostly common plants in the Coastal Zone. Husson (1978) published a monograph on the mammals of Suriname, Haverschmidt (1968) on the birds, Schulz (1975; Dutch edition in 1980) on the marine turtles, and Hoogmoed (1973) on the lizards. In the domain of applied science, there were important publications on agricultural pest insects by van Dinther (1960), and on useful plants by Ostendorf (1962). IMMEDIATELY AFTER INDEPENDENCE. During the first few years after Suriname’s independence in 1975, biological research flourished as never before. New research and development initiatives

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financed by The Netherlands created new opportunities. There was a strong focus on ecology, a reflection of the global trend at the time. An agreement was made between the University of Wageningen in The Netherlands and the University of Suriname (AdeKUS, with its research institute CELOS), to strengthen forestry and forest ecology research. This led to research on the so-called CELOS management system (CMS), a timber harvest and sylvicultural system described in a multi-author volume edited by Werger (2011). During the late 1970s to mid 1980s, a number of important studies were published that were based on fieldwork done in the preceding years: on primates (monkeys) by Mittermeier (1977) and van Roosmalen (1985b), on river otters by Duplaix (for example 1980), on wild fruits by van Roosmalen (1985a), on vegetation and ecosystems by Teunissen (see references in Ecosystems chapter), and on the traditional use of plants (ethnobotany) by Plotkin (1986). What drew a lot of attention internationally was a popular publication by Trail (1983) on the behavior of one of Suriname’s most charismatic birds, the Guianan Cock-of-the-rock. Much of the fieldwork of these authors took place in protected areas or focused on protected species. This was facilitated by several governmental institutions: the Nature Conservation Division of the State Forest Service (LBB), and the Foundation for Nature Preservation in Suriname (Stinasu). The role of the so-called ‘Teacher’s College’ (IOL) merits mentioning here, because IOL offered the only advanced courses in biological sciences in Suriname at the time. IOL students and graduates were involved in some of the research mentioned above. TRYING TIMES. The period from 1982 till the mid 1990s was politically tumultuous. An eight-year civil war led to the destruction, or at least sharp decline, of many development projects initiated in the previous decades. It also ruined Suriname’s economy, and negatively affected governmental institutions, such as the Central Bureau for Aerial Survey (CBL), the State Forest Service (LBB), the Geological Mining Service (GMD), and the AdeKUS university. From the mid 1990s onward, the political and economic situation improved. Biological research activities in Suriname followed this trend of decline and subsequent improvement. Just ahead of this period, in 1979, two new institutions had been created at AdeKUS: the National Herbarium of Suriname (BBS), and the National Zoological Collection of Suriname (NZCS). BBS was headed till the early 2000s by botanist Marga Werkhoven, who published a popular book on the orchids of Suriname (Werkhoven 1986). NZCS was and still is headed by herpetologist (amphibia and reptiles specialist) and ecologist Paul Ouboter, who in 1993 edited a multi-author volume on the ‘Freshwater Ecosystems of Suriname’ and in 1996 published his thesis on the ecology of caimans. At CELOS fish specialist Jan Mol did research on fish ecology, and published parts of his thesis work on armored catfish (for example Mol 1995). The NZCS and BBS undertook collecting expeditions to the remote corners of Suriname (see van ’t Klooster et al. 2003, Ouboter & Jairam 2012). This has led to publications on Suriname’s freshwater fishes (Mol 2012b) and amphibians (Ouboter & Jairam 2012), part of a series on the fauna of Suriname, which also includes a field guide to the country’s birds (Spaans et al. 2016). Several international herbaria have collaborated with BBS since its establishment, resulting in reports and publications on biodiversity and ethnobotany by Raghoenandan, van Andel, Ruysschaert, Hoffman, ter Steege, Banki and collaborators (see references in other chapters). Much of the fieldwork of these researchers, as well as researchers of the previous generation, has been facilitated by Frits van Troon, a Surinamer with a remarkable, hybrid scientific-local knowledge. He remains the most knowledgeable person on rainforest plants in Suriname.

Bart De Dijn

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BOX 4

Taxa and molecular taxonomy

A taxon (plural: taxa) is a coherent group of species or populations that are part of a single species, ideally representing a complete lineage, meaning all descendants of a closest common ancestor (de Quieroz & Gauthier 1990). Taxa are organized hierarchically; species are very basal in the hierarchy, yet may be further subdivided in subspecies (distinct, geographically isolated populations). The species Homo sapiens (modern humans), for instance, belongs to the human taxon (genus Homo), which is a group of species most closely related to and including Homo sapiens (the only surviving species of the group); the genus Homo in turn belongs to the taxon of Hominidae (‘great apes’), which also includes closely related chimps, gorillas and orangutans (McNulty 2016). Taxonomy is the naming, description and classification of taxa, such as species, based on shared and distinguishing characteristics (de Quieroz & Gauthier 1990). It is traditionally based on morphology: the presence, size, shape and structure of body parts, taking into account what is understood about their evolution. Contemporary taxonomy often makes use of a wide range of data: morphological, ecological and behavioral, as well as ‘molecular’. The latter refers to data encoded in biological molecules, particularly RNA and DNA which contains the genetic code. In this genetic material lies a detailed record of ancestry. But what is a species? Based on the ‘biological’ species concept, individual organisms are considered part of the same species if they can interbreed (mate and produce fertile progeny) (discussion of this and other species concepts: Mallet 1995, de Queiroz 2007). Most extant (currently living) species were never proven by breeding experiments, but assumed not to be able to interbreed with other species based on obvious morphological differences between the species, and the absence of extant intermediary forms in nature. Currently, species can be delimited based on molecular data (see for example Rannala 2015). No interbreeding between lineages, means that they do not physically exchange genetic material (no ‘gene flow’), and that specific changes in the genetic code due to random mutation that occur over time in one lineage are very unlikely to be shared with other lineages. The accumulation of changes in the genetic code ultimately results in lineages whose representatives lose the ability to interbreed with those of other lineages, producing what may be considered different species (de Queiroz 2007). Over the last three centuries, taxonomists (practitioners of taxonomy) have named and described taxa based on data that was available to them at the time and considered relevant, and based on different concepts of what should be considered a taxon, especially a species (see de Quieroz & Gauthier 1990, Mallet 1995, de Queiroz 2007). As a result, species and other taxon names, as well as taxonomic hierarchies, have often changed dramatically over time. In recent decades, better delimitations of species have been achieved based on molecular evidence of the interruption of gene flow; the classification of taxa was also improved based on genetic evidence of ancestral relations (see for example Heled & Drummond 2010).

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In the late 1990s and the 2000s, Stinasu facilitated research by a number of visiting specialists and their associates, such as marine turtle specialists Hilterman and Goverse (supported by WWF), primatologists Boinski and Norconk, small mammals specialist Lim, ornithologist Oâ&#x20AC;&#x2122;Shea, herpetologist Noonan, and seed dispersal expert Forget; Conservation International, in collaboration with local and international scientists, undertook several so-called rapid biological assessments (RAP; reports edited by Alonso and others) (see references in other chapters). Their work has produced important insights into biological diversity, ecology and nature conservation. Comprehensive biological studies were also undertaken in Suriname in the context of environmental and social impact assessment (ESIA): at bauxite mines and oil prospection and drilling sites in the Coastal Zone, and at bauxite and gold concessions in the northern part of the Interior. These studies were implemented by researchers such as Noordam, Teunissen, Bordenave, ter Steege, Ouboter, Mol, Lim, Oâ&#x20AC;&#x2122;Shea, and De Dijn (see reports referenced in other chapters, but not all are in the public domain). As part of some of these recent studies, tissue samples have been taken of birds, small mammals and amphibians. These samples have been used to generate so-called molecular data that are of critical importance in contemporary taxonomic research (see Box 4) (for example Velazco & Lim 2014), as well as biogeographical research (for example Noonan & Wray 2005) (see Biogeography chapter). This gives Suriname a taste of the new direction biological research is taking in the 21st century.

Bart De Dijn

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Geology, Landforms and Soils Salomon Kroonenberg, Dirk Noordam

hoever flies to Suriname for the first time cannot but wonder at the seemingly pristine landscapes of the country seen from the air. Upon approaching the coast from the north, you first see a half-submerged mangrove-clad coastal plain with narrow ridges. Then comes a somewhat dissected, but also partly drowned landscape, followed by white sand savannas just before you land. Further away to the south you see endless expanses of tropical forest disappearing into the hills. Before you touch down you have already seen the four main landscapes of Suriname: the Holocene Young Coastal Plain, the Pleistocene Old Coastal Plain, the Pliocene Zanderij Belt, and, in the distance, the Precambrian basement of the Interior. On the geological map of the country (Figure 6) they all come out clearly. The landscapes, rock units and their ages, and the sequence of events are given in Table 1. The Precambrian basement occupies about 80 percent of the land surface of Suriname, and is a part of the Guiana Shield. The Guiana Shield is the northern half of the Amazonian Craton, which is divided into two parts by the river basin of the Amazon and its tributary the SolimĂľes. Shields are the oldest cores of the continents, and the Precambrian spans the first 80 percent of the history of the earth, from its origin 4.6 billion years ago to 541 million years ago. The Guiana Shield has a long geological history, spanning the time between 3.5 billion and 600 million years ago (Gibbs & Barron 1993, De Vletter et al. 1998, Delor et al. 2003a,b; Fraga et al. 2009, Kroonenberg & De Roever 2010). However, almost all rocks in Suriname are about 2 billion years old, and were formed mainly as a result of a pervasive period of mountain building that affected large parts of eastern South America: the Trans-Amazonian Orogeny (Priem et al. 1971, GMD 1977, Bosma et al. 1983, De Vletter 1984, Kroonenberg et al. 2016) (see Table 1). The major rock types that were formed during that period are distinguished on the simplified geological map in Figure 6.

Geology of the Precambrian basement MAROWIJNE GREENSTONE BELT. The northeastern part of the country is occupied by a greenstone belt, so called because of the predominance of green rocks. It consists of a series of volcanic and sedimentary rocks that have been folded and metamorphosed (recrystallized under high pressure and temperature) deep in the earthâ&#x20AC;&#x2122;s crust. This greenstone belt is also the main area with gold mineralization (Daoust et al. 2011). Along its southern border large ellipsoidal bodies of granitelike rocks have intruded, also called TTG (tonalite-trondhjemite-granodiorite) bodies. OLDER GRANITES AND GNEISSES. The area south of the greenstone-TTG belt, more or less coinciding with the Marowijne and Tapanahony drainage basins, is occupied by rather inhomogeneous granites, often showing slight banding or parallel orientation. Close to the greenstone belt, older gneisses (strongly banded and oriented metamorphic rocks) predominate.

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Figure 6. Geological sketch map of Suriname, simplified after Kroonenberg et al. (2016) (image ÂŠ Kartomedia).

BAKHUIS GRANULITE BELT. A striking unit on the map of Figure 6 is the Bakhuis horst in the NW part of the Precambrian basement. This area consists mainly of granulites, dark grey metamorphic rocks that have been subjected to much higher pressures and temperatures during the TransAmazonian Orogeny than the greenstone belt. They show features of early stage melting, and may once have been as deep in the earthâ&#x20AC;&#x2122;s crust as 30 km (De Roever et al. 2003a). Here too there are intrusions, mainly of charnockites and anorthosites, equally dark gray intrusive rocks (Klaver et al. 2015). Salomon Kroonenberg, Dirk Noordam

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Natural History and Ecology Suriname | Geology, Landforms and Soils

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COEROENI GNEISS BELT. In southwestern Suriname a third unit of metamorphic rocks occurs, mainly gneisses with large amounts of micas. Some of them represent older sediments, others possibly older granitoid rocks. These rocks also have been subjected to intense pressures, high temperatures and incipient melting deep in the earth during the Trans-Amazonian Orogeny, though not as deep as the Bakhuis rocks (Kroonenberg 1976). In contrast to the Bakhuis Mountains, the Coeroeni Gneiss Belt occupies a lowland area, in which different types of gneisses give different landforms (Kroonenberg & Melitz 1983). YOUNGER GRANITES, GABBROS AND VOLCANIC ROCKS. The western half of central Suriname, especially in the Wilhelmina Mountains (Verhofstad 1971) and in the Corantijn and Sipaliwini drainage basins, consists of very fine grained volcanic rocks (Dalbana Formation) and granite bodies intruded at a shallow level in the earth’s crust. Both rock types stem from the same magma sources. The volcanic rocks have been deposited mainly by pyroclastic flows (glowing ash clouds), and have been slightly recrystallized in contact with hot intruding granite magma. Small, round gabbroic intrusions of similar age (Lucie Gabbro) are widespread throughout the basement. Gabbros are dark-colored iron-rich rocks. TAFELBERG FORMATION. The Tafelberg is an eastern outlier of the Roraima Supergroup that forms impressive sandstone plateaus in Guyana, Venezuela and Brazil (Bisschops 1969, Bosma et al. 1983, Santos et al. 2003b).The Tafelberg Formation consists of at least 700 meters of indurated pinkish sandstones and quartz conglomerates. Sedimentary structures as trough bedding and ripple marks suggest deposition by rivers. The contact with the underlying granites is marked by a Precambrian fossil soil (Kroonenberg 1983). The age of the formation was debated in the past, until Priem et al. (1973) established its Proterozoic age from red volcanic ash layers between the sandstones. Since then, more accurate data have been obtained in Brazil by Santos et al. (2003), giving an age of 1873 million years. DOLERITE DYKES. There are three generations of dolerite dykes, kilometers-long fractures filled by basaltic magma during periods of extension in the Guiana Shield. The oldest one, the Avanavero Dolerite suite (Reis et al. 2013), forms conspicuous mountain ridges such as the Van Asch van Wijckgebergte. The Käyser Dolerite suite in SW Suriname with predominant N-W orientation is slightly younger (De Roever et al. 2003b). The youngest one, the Jurassic Apatoe suite, with predominantly N-S orientation, is the only rock unit in the basement younger than the Precambrian; it marks the extension where Wegener’s supercontinent Pangea started to break up and South America and Africa started to drift apart (Deckart et al. 2005).

Landforms and landform formation in the Precambrian basement PHYSIOGRAPHY. The Guiana Shield area is dominated by an endless mosaic of low to moderately high convex hills. The northern part of the shield is characterized by undulating to moderately steep-sided, low hills with elevations of up to 100 m. Further south, large areas show a rolling relief with very steep-sided, low to high hills with elevations between 100 m and 250 m. Above this hilly terrain, mountains, plateaus and residual hills form prominent features up to 750 m. The higher summits are often covered with laterite or bauxite duricrusts (see below and Figure 10). Conspicuous mountainous terrain and high plateaus occur locally within almost all of the above zones, in some

Salomon Kroonenberg, Dirk Noordam

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Figure 9. Tafelberg, view from subtop to the west (image © Bart De Dijn, 2008). Figure 7. Landforms in the Marowijne Greenstone Belt strongly influenced by rock type (image © INPE (Brazil); Landsat image #2763050-130119-7 (band 7, infrared), October 31st, 1976). BXT-MB: Bauxite plateau on mafic (iron-rich) rocks (Paramaka Formation, see Table 1); MTB: metamorphosed turbidites (Armina Formation); MST: metasandstone (Rosebel Formation); AMF: Amphibolite; MQA: Meta-quartzites; TTG: Tonalite-TrondhemiteGranodiorite plutons. Note also cataracts (sula) in Marowijne and Tapanahony rivers.

Figure 8. Devil’s egg, granite inselberg (bare mountain top) and tor (the loose core stone) in the Wilhelmina Mountains, Central Suriname (source www.pinterest.com/pin/336573772124194603/; author unknown)

cases extending to elevations in excess of 750 m. The Juliana Peak at 1230 m is the highest point in Suriname. Each rock type of the Precambrian basement produces its own landforms and soils (O’Herne 1969, Melitz 1975, Kroonenberg & Melitz 1983). The basaltic rocks in the greenstone belt (Paramaka Formation) form the typical bauxite plateaus of Brownsberg, Nassau and Lely mountains and smaller occurrences. The TTG rocks, on the other hand, form low areas: the Brokopondo storage lake (Brokopondo Reservoir) occupies such a low area (Figure 7). Even narrow ridges of more resistant rock types are directly visible in the LANDSAT satellite images. In granitic terrains, domed inselbergs (bornhardts) crop out (Figure 8); rock pavements (ruwares) and boulder inselbergs (tors) are also present here, in general covered by forest. In the Sipaliwini Savanna and elsewhere, outcrops of fine-grained volcanic rocks of the Dalbana Formation (see above) may form long ridges in the landscape. Gabbro intrusions form

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Figure 10. Plinthite horizon at Mosqueiro, near BelĂŠm, Brazil (image ÂŠ Salomon Kroonenberg, 2013). A plinthite horizon in a soil profile is the result of changing ground water levels in the soil. When plinthite is exposed to the air, for instance if the overlying soil is eroded away, or, as is the case here, the profile is exposed in a cliff, plinthite hardens into a duricrust, i.e. a laterite cap on mafic (iron-rich) basement rocks, bauxite on aluminous basement rocks as in the Bakhuis Mountains, or in aluminous sediments in the Suriname Coastal Plain.

Figure 11. Stages in the development of an etchplain (Kroonenberg & Melitz 1983). Stages 1) Humid climate: deep weathering, formation of plinthite over mafic rocks, shallow weathering in slightly jointed granite, deep weathering in gneisses; 2) Arid climate: stripping of the weathered rock, plinthite hardens to laterite or bauxite, inselbergs and tors are exposed; 3) Humid climate and 4) Arid climate: the processes are repeated, height differences increase.

conspicuous hills due to laterite duricrusts. Some of the rocks from the Bakhuis Granulite Belt contain so much aluminum that bauxite plateaus have been formed upon them by chemical weathering. The prominent horizontal sandstone plateau of Tafelberg in central Suriname reaches an elevation of 1026 m (Figure 9). A plateau at a lower level around the Tafelberg itself, including the Kappel Savanna, consists of the lowermost part of the same formation. Dolerite dyke suites form lateritecapped ridges protruding above the surrounding basement landscapes. DIFFERENTIAL ETCHING AND STRIPPING. Rocks in humid tropical environments experience intense chemical weathering. Rainwater mixed with acids from rotting leaves and trees penetrates through cracks in the rock, and changes the most susceptible minerals such as feldspars and micas into clay and iron oxide, so that only a red loamy soil, often with residual quartz, remains. Open pit mining operations and drill holes show that sometimes you have to go down 100 m through soil and weathered rock before you touch the fresh rock. But not all rock types are equally susceptible to chemical weathering. Mica-rich rocks like gneisses are often deeply weathered, whereas granitic

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rocks usually weather less deeply. Iron-rich and aluminum-rich rocks may develop plinthite horizons (layers within the soil) rich in iron oxides which may harden upon exposure, and then develop into laterite or bauxite duricrusts (see explanation in Figure 10). This differential chemical weathering of rocks is called etching (Thomas 1975, Rabassa 2014). Now imagine that a drier or at least more seasonal climatic episode sets in, in which the tropical forest is replaced by savanna vegetation. Savanna protects the soil much less efficiently than the tropical forest, and soon soil erosion leads to stripping of the weathering mantle and planation of the land. The deepest weathering mantles get most affected, and become deeply eroded terrains. Granites with widely spaced vertical joints (fissures), in the order of tens to hundreds of meters, have shallow weathering mantles and become exposed as fresh rock outcrops such as inselbergs and tors (Figure 8). Granites with more closely spaced joints may weather more deeply. Plinthite horizons developed from iron-rich rocks become exposed, harden, and form laterite or bauxite caps that prevent further stripping. When in a subsequent climate cycle moister conditions return, the tropical forest comes back again, but less luxuriously on laterite plateaus and not at all on exposed granite outcrops, and deep weathering is resumed. After a few of such cycles of etching and stripping, the height differences gradually increase between the laterite caps and the granite outcrops on the one hand and the gneiss landscapes on the other. Thus the well-known basement topography of plateaus and inselbergs and tors appears. The sequence of events is depicted in Figure 11. These climate cycles are usually correlated with supposedly drier climates during the Pleistocene ice ages (Van der Hammen & Absy 1994), although recent research shows little support for dry climate during the ice ages in Amazonia (Hoorn 1997, Hoorn et al. 2010). TECTONICS. Erosion is only capable of removing the soil mantle if there is potential energy in the landscape, i.e. a slope from the headwaters of the rivers towards the sea. What causes that elevation? The only way to achieve it is tectonic uplift. Without uplift it is impossible to imagine that rocks which once were 30 km deep in the interior of the earth are now exposed at the surface. It is also impossible to imagine the origin of the Tafelberg Mountain otherwise: in the Precambrian this must have been the lowest basin in the landscape to be able to receive river sediments. Now it is a high top: a clear case of relief inversion as a result of tectonic uplift. The rate at which the surface of the land is being lowered by the combination of uplift on the one hand and etching and stripping on the other is called denudation rate. The denudation rate in the Guiana Shield was calculated from the water chemistry of the Orinoco River at 15-20 m per million years in highlands and 5 m per million years in the dissected lowlands (Edmond et al. 1995). The clearest example of uplift is in the Bakhuis Mountains. This entire area is a horst, meaning that it is uplifted with respect to the surrounding land. The resulting mountainous character is further enhanced by the presence of prominent bauxite caps. Although the SW-NE direction of the border faults is inherited from Precambrian events, the Bakhuis horst is still active under the sediments of the coastal plain, and has had an impact on the location of the oil fields in the Saramacca area (Bosma et al. 1983, Wong 2014, Nelson, 2016). Other areas have subsided, such as the 6 km deep Jurassic Takutu graben in Guyana. The fact that the coastal area in western Suriname is much broader than in the east is also related to the subsidence of that graben. Suriname is tilting northwestwards. These up and down movements of

Natural History and Ecology Suriname | Geology, Landforms and Soils

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the Guiana Shield are probably an echo of the separation of South America and Africa since the Jurassic. PLANATION LEVELS. In many areas underlain by iron-rich rocks such as the Paramaka Formation, several laterite and bauxite plateaus at different heights above each other have been found, separated by steeper steps in the terrain, for instance at Brownsberg Mountain. As denudation results in lowering of the surface, higher plateaus are considered older than lower ones; the elevation of a laterite plateau is thus regarded as a measure of its age. The Tertiary age (Eocene-Oligocene, see Table 1) of the oldest laterite plateau in the basement is confirmed by the occurrence of bauxites of the same age in the coastal plain (Wong 1989), and by absolute dating of the laterites themselves (ThĂŠvĂŠniaut et al. 2002). The level of the highest laterite plateaus in Suriname slopes towards the north from Lely Mountain (700 m), Nassau Mountain (600 m) to Brownsberg Mountain (500 m). It is therefore tempting to connect them as evidence of a countrywide ancient surface sloping to the sea: a planation surface. The concept has been extended to surfaces without laterite caps as well: the whole continental surface was thought to be subdivided in planation surfaces separated by steps in the topography, formed by parallel slope retreat during periods with dry climate independent of rock type. The highest level was called the Early Tertiary Level, followed by the Late Tertiary I and II Level (Zonneveld 1993, Zonneveld & Krook 1998). But the steps between levels outside the lateritized areas appear to coincide with boundaries between rock types, suggesting they are formed by etching and stripping, not by parallel slope retreat (Kroonenberg & Melitz 1983). Planation by backwearing of slopes as envisaged for examples by Kips and Snel (1979) is not a viable process in crystalline rocks. The lithological control of unlateritized surfaces means that their altitude should not be taken as a measure of age. The landforms of the basement outside the laterite and bauxite plateaus are essentially an etchplain, not a succession of planation levels (Kroonenberg & Melitz 1983). RIVERS AND CREEKS. A striking feature of the rivers in the basement is the occurrence of cataracts or rapids (known as sula in Suriname): sudden changes from a single channel pattern to a multichannel pattern, not unlike that of braided rivers (Figure 12). These cataracts develop at sites where Figure 12. Bonnidoro Falls, Marowijne district, typical sula complex in Armina Formation meta-greywackes (image ÂŠ Google Earth, 2015).

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Figure 13. Creeks following fault patterns in dissected granite landscape, W of Oelemari river, at about 2°N55’30” N , 54°W 40’30” W (image © Centraal Bureau Luchtkartering, Suriname; aerial photograph run 13ZO, 6739, Z=4500 m f= 152.9 mm, November 1977).

rivers, while incising into the weathering mantle, suddenly encounter hard rock. Chemical weathering is so intense in the drainage basin that no gravel is formed, and therefore the rivers do not carry bedload coarser than sand. As a result they do not have enough erosive power to incise into the bedrock, and instead they try to negotiate any hard rock obstacle by passing it sideways. In this way the river splits in different channels around hard rock outcrops in sometimes very complex ways (as in Figure 12). River courses are also strongly influenced by linear features in the bedrock. They tend to follow faults, fractures and shear zones, and to avoid or be deflected by dolerite dykes. Along the major rivers, staircases of river terraces are found: these are abandoned floodplains at a certain elevation, often at about 6 m and 15 m above the present floodplain (De Boer 1972). Some terraces contain rounded gravel, and some are gold bearing, whereas at present no gravel is being formed. This indicates that weathering conditions in the basement might have been less severe during the deposition of the terrace sediments than at present. Terraces might have been deposited by braided rivers in Pleistocene periods with more arid climate and less vegetation cover, and become dissected at the return of the tropical forest. During the next more arid climatic event a new terrace would have been deposited at a lower level, because tectonic uplift would have left the previous terrace at too high an elevation above the river to be flooded again. River terraces are especially well developed along the Marowijne, as that is where uplift is more vigorous (De Boer 1972). Absolute ages from these deposits are not available. The smaller creeks that dissect the relief in numerous small hills are deeply incised into the weathered rock, and rock outcrops are often found at the creek bottoms; elsewhere, the valley bottoms are swampy and filled with young sediments. The creeks often follow rectilinear lineaments in the basement (Figure 13). Tectonic uplift is probably the main reason for this dissection. Sea-level change, as in the Old Coastal Plain (see below), cannot play a role, because dissection occurs everywhere in the Interior, not only close to the sea. Moreover, the cataracts form local base levels that prevent gradient changes in the coastal area from moving upstream.

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Figure 14. Sipaliwini Savanna, with boundary savanna-forest maintained by burning (image ÂŠ Salomon Kroonenberg, October 2013).

SAVANNAS IN THE INTERIOR. There are two larger savannas within the Interior, the Sipaliwini and the Gros savannas. These savannas cover approximately 0.5 percent of the Interior, but their combined area exceeds the combined area of the savannas in the Zanderij Belt (see below). The Sipaliwini Savanna (Figure 14) is underlain by exactly the same rocks as the surrounding forested area. Climatic conditions are also the same, as is the general morphology of low dissected hills. However, due to more active soil erosion, gully formation and slope movements, valley fills in the savanna are more prominent, leading to adverse soil factors (soil-water relations; Riezebos 1979). The Gros Savanna is underlain by sandy soils derived from quartz arenites of the Rosebel Formation. This might represent a model for river and slope processes under more arid conditions, though the origin or at least the persistence of the savanna is due to human activity, namely burning, rather than due to the prevailing climate (see Ecosystems chapter). SOILS OF THE PRECAMBRIAN BASEMENT. The soils of the Precambrian basement are predominantly developed in the loose regolith that overlies the fresh, unweathered rock. Regolith is in-situ weathering material, colluvium (slope deposits) local alluvium (river sediments) or a combination thereof. The often deeply weathered parent material and its colluvium or alluvium are already strongly leached and poor in nutrients, and thus the soils that develop from it are mostly chemically poor as well. Kaolinite is the main clay mineral, which has only a limited capacity for retaining plant nutrients. There is a fair proportion of iron oxides. In spite of the high amount of litter production above the ground, organic matter is low in the soil, because of the high mineralization rate under the humid and warm conditions in the tropical forest. Also, nutrient content is low, because most nutrients from the litter are already scavenged by the standing vegetation before they can be incorporated into the soil. Texturally soils in the basement differ widely, mainly depending on the parent materials. Soils derived from coarse-grained granitoid rocks are usually rather sandy because of the presence of residual quartz, while rocks from gneisses are more clayey due to the high amount of micas weathered to clay and far less residual quartz. Soils on iron-rich rocks such as the meta-basalts in the greenstone belt, the gabbros and dolerite intrusions are deep red due to the

Salomon Kroonenberg, Dirk Noordam