Butterflies of Suriname by LM Publishers

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Butterflies of Suriname A natural history

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Butterflies of Suriname A natural history

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Butterflies of Suriname A natural history Hajo B.P.E. Gernaat, Borgesius G. Beckles, Tinde van Andel

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Colophon Butterflies of Suriname. A natural history

KIT Publishers

Hajo B.P.E. Gernaat, Borgesius G. Beckles, Tinde van Andel

Mauritskade 63 Postbus 95001

This publication has been produced with the support of:

1090 HA Amsterdam E-mail: publishers@kit.nl www.kitpublishers.nl

NCB Naturalis

NCB Naturalis Darwinweg 2 Hugo de Vries Fonds

2333 CR Leiden E-mail: contact@ncbnaturalis.nl

Uitvoeringsorganisatie Twinningfaciliteit

Website: www.naturalis.nl

Suriname -Nederland

The Netherlands Centre for Biodiversity (NCB Naturalis) contributes to the preservation of biodiversity by using its extensive collection and Alberta Mennega Stichting

the results of its research to foster knowledge and awareness of life on earth through its educational and public information programmes.

SLI

ÂŠ2012 KIT Publishers â&#x20AC;&#x201C; Amsterdam Uyttenboogaart-Eliasen Stichting (UES)

Lay out: Ontwerpbureau Agaatsz BNO, Meppel

Production: High Trade BV, Zwolle

ISBN 978 94 6022 171 2

Every effort has been made to contact and obtain permission from owners of copyrighted material quoted in this book. In case of oversight, please contact the publisher.

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Contents

Foreword Acknowledgements How to use this book Part I

Part II

Part III

Biological names and classification, geography, geology and soils 1. Latin names and biological classification 2. Geography, geology and soils Plants 3. Some basic plant anatomy and physiology 4. Introduction to the rain forest Complexity Bromeliaceae Ecological succession Plant defenses Pollination and pollination syndromes Fruits and frugal syndromes 5. Plant diversity in Suriname Plant diversity in the Guianas Plant diversity in Suriname 6. Habitat overview The coastal plain The savanna belt The inland mountainous region The butterflies of Suriname â&#x20AC;&#x201C; introduction and history 7. The butterflies of Suriname â&#x20AC;&#x201C; an introduction Butterflies and moths Butterfly families Form and function of butterflies Male versus female butterflies The life cycle of butterflies Migration Butterfly enemies Parasitoids Predators Butterfly defenses Plates 1-7 Mimicry........................... The butterflies of Suriname in the context of the world and neotropical butterfly fauna

9 11 14 17 19 23 35 37 47 47 52 52 54 56 59 67 67 68 73 73 76 76 83 85 85 87 90 92 96 102 103 103 105 110 122 136

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Part IV

8. History of the study of the butterflies of Suriname Introduction Maria Sibylla Merian and the Metamorphosis Insectorum Surinamensium Carl Linnaeus and the Systema Naturae Daniel Rolander and the Diarium Surinamicum Pieter Cramer, Caspar Stoll, the Uitlandsche Kapellen and the Aanhangsel Plates 8-11 The Cramer butterfly types for Suriname in NCB Naturalis Johann Christian Fabricius J.C. Sepp and the Surinaamsche Vlinders August Kappler (1815-1887) Heinrich Benno MĂśschler and the BeitrĂ¤ge zur Schmetterlingsfauna von Surinam The Penard brothers and others The great expeditions (1901- 1938) Dirk C. Geijskes (1907-1985) Piet, Van Dinther, Belle and Mees Heinrich B. Heyde (1921-1993) Hermann J.T. Stammeshaus (1918-1991) E.H. Jonkers, D. Schilder and R. de Jong The National Zoological Collection of Suriname The Neotropical Insects and Butterfly Park in Lelydorp Summary The butterflies of Suriname - species accounts and Plates 9. Checklist and Plates of 150 butterfly species of Suriname 10. Hesperiidae (the skippers) Subfamily Eudaminae Subfamily Pyrginae Myrtaceae Combretaceae Subfamily Hesperiinae 11. Papilionidae (the swallowtails) Annonaceae Aristolochiaceae Rutaceae Piperaceae 12. Pieridae (the whites and sulphurs) Subfamily Dismorphiinae Subfamily Coliadinae Fabaceae Subfamily Pierinae

143 143 143 146 149 151 156 164 164 166 168 168 168 172 174 175 177 178 179 180 181 183 185 273 274 279 280 282 288 293 297 303 310 319 321 321 323 330 338

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13. Lycaenidae (the blues and coppers) Subfamily Theclinae Subfamily Polyommatinae 14. Riodinidae (the metalmarks) Araceae 15. Nymphalidae and Danainae Nymphalidae (the brushfooted butterflies) Danainae (the monarch butterflies) Caricaceae Apocynaceae 16 Ithomiinae (the glasswing or clearwing butterflies) Solanaceae 17 Morphinae (the morpho and owl butterflies) Arecaceae Musaceae Heliconeaceae 18 Satyrinae (the brown and ringlet butterflies) Poaceae 19 Charaxinae (the charaxes butterflies) 20 Biblidinae (the small pages) Euphorbiaceae Moraceae 21 Nymphalibae (the peacock and buckeyes) Cecropiaceae Acanthaceae The Phyciodina subtribe 22 Limenitidinae (the admirals) Rubiaceae 23 Heliconiinae (the passion flower or longwing butterflies) Passifloraceae Appendix I Appendix II Appendix III Appendix IV Appendix V Appendix VI Appendix VII References Index

How to enjoy the forest â&#x20AC;&#x201C; what to do and what not to do The butterflies of Suriname and their natural history in the Metamorphosis Insectorum Surinamensium Maria Sybilla Merian The Cramer butterfly types for Suriname in the Netherlands Centre for Biodiversity (NCB) Naturalis Data on figured specimens (Plates 1-52) Glossary of Surinamese, scientific and English names for plants and animals used in this book Glossary of technical terms Authors, photography and the SLI

347 347 352 355 365 375 375 375 378 385 393 397 407 428 435 441 449 454 465 473 479 495 499 500 516 521 527 529 535 553 577 579 591 599 619 627 636 643 661 7

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Foreword

This is the first book of an intended series on the butterfly fauna of Suriname. This part is meant to provide background data and to introduce the reader to the butterflies of Suriname. There are three groups of people for which this book is meant: the people of Suriname, tourists and other interested nonprofessionals, and students of biology. During the process of writing, it has become a bit of an unconventional book. We originally intended to write an identification guide to the common butterflies of Suriname, i.e. species likely to be seen by an average tourist on a three-week trip, supplemented by some species of special interest. Suriname is a wonderful country with great people, beautiful butterflies, many other animals and plants as well. All of these make for great tourist appeal. For many people on such a trip, however, it is their first encounter with the rainforest. For some, this turns out to be a disappointment. They are, on the one hand, primed with adventurous or fantastic stories of extraordinary animals and plants; and, on the other hand, by dismal articles on deforestation, global warming and similar themes, suggesting a sense of urgency to visit the forest ‘before it is too late’. People pay a lot of money for a visit, only to find that they see ‘green, green and green everywhere’, occasionally interrupted by an insect or a bird call, and end up wet, muddy and tired. ‘Is this all,’ they wonder ‘and worth all this trouble and money?’ There is one phenomenon that is usually not mentioned in advertisements for tourist trips: most species in the rainforest are rare and seldom seen. Some of them, like howler monkeys and birds, you may hear, you may see tracks or droppings of others like the jaguar, but most remain hidden. There are at least 1,300 species of butterflies in Suriname and most of them are rare, which means that there are not many individuals of a given species at a certain point in time. Chance, therefore, is a major factor in spotting a species or not. A visit to the rain forest, however, can be extremely rewarding and fill you with a pervading sense of awe and admiration. For this, two things are essential: you have to be patient and pay attention to detail. If you do that, there are treasures to be found around every corner. For life in the forest is hidden. It is a place where every species has to go to great lengths to survive. Over millions of years, life forms have adapted and become specialized to an extent that is hardly seen in temperate climates. The diversity of life is gigantic and, even today, not yet fully comprehended. Imagine a 5 km stretch of track through the forest, for example the Brownsberg 9

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Nature Reserve. You can walk this track every day for three years and see new butterfly species each day. Butterflies are an excellent way to be introduced to the intricacies of life in the forest. During a visit, you will see many butterflies that with, the help of this book, are fairly easy to identify. You will notice, that if you pay attention to butterflies, you will see many other wonderful things in the forest and, instead of â&#x20AC;&#x2DC;green everywhereâ&#x20AC;&#x2122;, the forest may be transformed to an unprecedented treasure house of life forms. Butterflies are linked to plants: they lay their eggs on them and caterpillars eat them. Plants, caterpillars and butterflies all try to avoid being eaten and have developed specialized ways to survive. Other species are also connected to plants. So, in this book there are stories on butterflies, but also on plants and other species that have some link to them. By paying particular attention to photography, we have tried to convey our sense of respect for nature and our enjoyment of its beauty. We did not intend to be complete or systematic, but only to tell stories that are interesting. Hopefully, you will become fascinated too. Two final things about butterflies. First, some people in Suriname believe that they may be dangerous, for example venomous or able to bite or sting. Be reassured, this is definitely not the case. Butterflies are entirely harmless, ready to be enjoyed. Beware, however, of hairy caterpillars. These may indeed cause irritation or they may even be poisonous. Second, we have heard some people in Suriname believe that a butterfly resting on an individual would be a sign something bad is going to happen to that person. We believe that there is actually no evidence for this. We have had butterflies on us scores of times over the last ten years and remain in good health. So, donâ&#x20AC;&#x2122;t be distraught by such ponderings and enjoy the book! The authors

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Acknowledgements

Although writing a book is in large part a solitary endeavor, a large number of people helped most generously and enthusiastically. This book had its awakenings in 2001, when the first author, rather accidentally, discovered in the collection of NCB Naturalis in Leiden, The Netherlands, a great many cabinets containing unidentified butterflies from Suriname and hundreds of specimens in papillots, several of which collected more than a hundred years ago. A subsequent literature search showed that very little had been published on Surinamese butterflies in the last 150 years. After Paul Ouboter and Helene Hiwat, then curator of the invertebrate section of the Anton de Kom University in Paramaribo, and Marie Djosetro, then head of the research department, later director, of Stinasu, had expressed the need and enthusiasm for an inventory of the Surinamese butterfly fauna, the decision to write a book was made and a first collection trip took place. Then, the first author received an email from Helene Hiwat, reporting that a certain lawyer, called Borgesius Beckles, had contacted the university requesting details on butterfly species he had photographed. He was quite unknown at the university, but had the most magnificent photographs and said he planned to write a book on butterflies. Quickly, contacts were made, forces were joined and cooperation developed, which gradually turned into a friendship. As the need for botanical expertise grew, Tinde van Andel joined in. During the production process of the book, Eelco Kruidenier, Frans Barten and Liz Steele-Gernaat were so essential, that they are mentioned in Appendix VII. Over the years, the staff and trustees of NCB Naturalis have been most supportive, always creating a warm and welcoming atmosphere. Rienk de Jong was very helpful, always willing to share some of his vast knowledge on Lepidoptera, provided literature, had suggestions and reviewed an earlier draft of this book. Eric van Nieukerken was always available to make arrangements for good working conditions and publication details. Cor Lepelaar and, later, Behnaz van Bekkum-Ansari made for warm welcomes at NCB Naturalis. We are very grateful for Behnaz’s search for Surinamese specimens in the vast Lepidoptera collection and for Rory’s perseverence in databasing many Surinamese butterfly specimens. Cees van Achterberg had the patience to answer questions and provide additional information to an absolute layperson with regard to parasitoids. Klaas Douwe (‘KD’) Dijkstra had an important part in the early stages of the book. During his stay in Suriname, he became enthusiastic about the concept of the book and later managed to secure funds from the Twinning project to 11

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ensure publication for a reasonable price. Rene Dekker and Sancia van der Meij provided the necessary managerial support for the project. The staff of the Zoological Museum in Amsterdam, before merging with Leiden, has been most supportive. We enjoyed the cooperation very much with Sandrine Ulenborg on Maria Sybilla Merian for the exhibition in the Rembrandt House Museum in Amsterdam and the J. Paul Getty Museum in Los Angeles as well as for Ella Reitsmaâ&#x20AC;&#x2122;s book on Merian. Willem Hogenes has, over the years, been most supportive in allowing access to the ZMA collection, providing information and has shown an unflagging interest in the project. The library of the Netherlands Entomological Society (NEV), then housed in the ZMA building, is one of the most comprehensive of Europe with about 2.2 km of bookshelves with 21,000 books, 4400 journals and 100,000 reprints. Godard Tweehuysen and Danny Boomsma somehow managed to extract from this vast depot of information, in an impossibly short time, books and publications in Dutch, English, French, German, Spanish, Portugese, Japanese and Latin. Nothing was ever too much to ask and their support has been greatly appreciated. There have been many people who were willing to share expertise from their own field of interest without whom this book would be quite impoverished. We thank them all. Early on, Bert Jongerling showed his own magnificent collection of Surinamese butterflies, provided lots of information and ensured his support for the book. His emigration to the USA, unfortunately, prevented his continued support. Andrew Neild was willing to share the results of his Junonia quest even before his second book on the butterflies of Venezuela was published and we had lively email discussions. Christian BrĂŠvignon sent his beautiful posters on French Guinean butterflies and many times we exchanged information on butterflies. Rene Marcelis shared data from his own collection of Surinamese butterflies. Mike Gilman gave information on butterflies from Guyana. Matthew Cock sent his publications on the Hesperiidae of Trinidad with many life history details. Theo Wong reviewed the geology and soil section and thus prevented quite a few omissions. Christian Feuillet at the Smithsonian Institution in Washington generously provided additional information on the Passifloraceae of Suriname. Martin Heyde and his wife generously provided information on his father, Heinrich Heyde. Dre Teunissen gave data on Engbert Jonkers. Jan-Hein Ribot and Otte Ottema were always willing to discuss and add data on Surinamese birds. Paul Maas identified some of the plants figured, especially Heliconiaceae and Passifloraceae. Jan Mol shared information on fishes. Piotr Naskrecki at Harvard University and Elsa Youngsteadt looked for photographs of Camponotus femoratus (which, unfortunately for this book, they did not have), but their efforts are greatly appreciated. Many people made their photographs available, for which we are very grateful: Bobby Angell, Frans Barten, David and Maurits Gernaat, Carol Gracie, Dan 12

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Janzen, Andrew Neild, Riad J. Nurmohamed, S. Naipal, Marcel Pollack, Ro Sewdien, L. Westra and Mat Wijnen. Plant identification and botanical drawings by Hendrik Rypkema and W.H.A. Hekking were provided by the National Herbarium, now part of NCB Naturalis. And last, but certainly not least, we were encouraged and helped over the years by several institutions and individuals in Suriname. Stinasu, both its management and the research department (currently Serano Ramcharan) have been most supportive, granting research permits for the Raleigh falls and, especially, the Brownsberg Nature Reserve. We were particularly grateful for the license to collect, apart from the adult butterflies, also their early stages and foodplants, many data of which are presented in this book. As butterflies also have relationships with other organisms (various predators, parasitoids, ants, mites, fungi, bacteria etc.) and so much is unknown of the ecological relationships of butterflies in Suriname, a great deal of work remains to be done. People from the Anton de Kom University in Paramaribo, notably Paul Ouboter, Helene Hiwat and Jan Mol, have consistently encouraged the project and allowed ready access to the Lepidoptera collection. Plans to find a common publisher for books on amphibians, fishes and butterflies unfortunately foundered, but we are glad that our good relationship did not suffer from this. We thank Aniel Gangadin and Sheryl Starke for their willingness to help in collection matters and various other areas. METS Travel & Tours specializes in the development of sustainable tourism in Suriname and has shown a keen interest in nature as well as the development of local communities in the inland of Suriname. We are highly appreciative that the director of the METS, Armand Bhagwandas, has repeatedly supported us by favorable ticket conditions and allowed us to transport our research equipment. Without this, the data collection and photography would have suffered considerably. His help has greatly benefited the contents of this book. Bart DeDijn has a great deal of entomological and botanical expertise in Suriname. His emails and information on pollination syndromes have been of great help. Ewout Eriks has decades of experience on the natural history of butterflies in Suriname, as well as their breeding. He was always willing to share information, allow access to his facilities for photography and readily informed us of new findings. Over the years, Gerda Kajuffa has been as accommodating and hospitable as anyone could wish for, enduring many an intrusion in the daily household routine, including, among other things, the breeding of butterflies in the living room, plant samples stored under the bed and boxes of dried butterflies in the fridge. Lastly, we profoundly thank our partners, Heidi and Gerda, and children, David, Sofie, Maurits, Benajo, Alvin and Borger, for sacrificing so much of our time and activities together. 13

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How to use this book

As noted in the foreword, this book is meant as an introduction to the butterflies of Suriname in an ecological context. We have tried to make a book for a general readership with lots of photographs, but also with sound biological information. For the general reader, all technical terms are explained in the book starting with biological classification and Latin names. Then, in order to better appreciate the environment butterfly species live in, the geography, climate, geology and soil types of Suriname are briefly discussed in part I. This is followed in part II by an introduction to plants and the rain forest, which covers about two thirds of the country, plant diversity in Suriname, as well as Suriname’s main habitats. In part III, butterflies enter the picture. After an introduction to their biology, the butterflies of Suriname are dealt with in the context of the world and neotropical butterfly fauna. This part is concluded with an account of the rich history of the study of the Surinamese butterflies. In part IV, 150 butterfly species are discussed in detail. In Appendix I - VII some additional information is provided, among other things, on how to enjoy the forest and avoid trouble, details on Maria Sybilla Merian and the Cramer types, and glossaries of Surinamese names and technical terms. Most books are probably meant to be read from the beginning till the end and, therefore, have a more or less systematic line of reasoning, as hopefully this book has as well. However, we realize that only a minority of the readers will use it in this way. Most users will probably want to look at photographs and read ‘around’ them or they may look for a figure that most closely resembles the butterfly they have just seen and want to read up on it. For this reason, the book is full of cross-references, enabling the casual reader to go ‘from story to story’. This may possibly disturb the more systematic reader but it has been a deliberate choice. Also, because of this approach, some information has been included more than once. The choice of which butterfly species to include has been only vaguely guided by reason. We had a prototypical tourist in mind who books a three-week summer holiday in Suriname with some trips into the interior and looks for butterflies but not exclusively. We estimated this tourist would see about 100 different species, most of which are discussed in this book. The other fifty we chose rather at random. Some are rare or interesting, others are representative of certain groups. We had to decide to use Latin names, as popular names for butterflies hardly exist in Suriname and popular names usually vary between countries. The 14

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local, Surinamese names are usually mentioned, if known to us, and summarized in Table VIII for birds and Appendix V for other animals and plants. The best way to deal with the Latin names is not to skip them when reading, but to read them slowly and pronounce them softly to yourself. In this way you quickly get used to them. Scientific names of butterflies were based on Lamas (2004) or more recent literature, the names of plants on Funk (2007), Feuillet (2009) and The Plant List of the Royal Botanic Gardens Kew (www.theplantlist.org). In the butterfly species accounts, we decided not to be too schematic to improve readability. Still, there is a standard structure: - plate and figure numbers of the species and subspecies occurring in Suriname - identification characteristics - type description - distribution of the species - number of subspecies - distribution of the subspecies occurring in Suriname - distribution within Suriname - habitat and behavior - foodplants - (possible) foodplants in Suriname - data on life history - data on parasitoids The introductory accounts on plant families similarly have a standard structure: - family name, English name followed by a well-known family member in Suriname - butterfly species in the book that have a family member as their foodplant - number of species worldwide, in the Neotropics, in the Guianas and in Suriname - origin of the family name - identification characteristics of the family - some data on its biology (pollination, pollinators, means of seed dispersal) - human use of some plant family members Other deliberate choices to improve readability were to omit author names of species (for butterflies, they are mentioned only once, in the species checklist; for other animals and plants, they are not mentioned) and, in the text, to refer to literature only by the first author and year of publication. We hope other authors do not feel offended by this. In the references section, full data for publications are given.

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Part I Biological names and classification, geography, geology and soils

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Figure 1-01 Three butterfly species of Suriname; a. Phoebis sennae marcellina, male, upp.; b. Phoebis philea philea, male, upp.; c. Morpho menelaus menelaus, male, upp.

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Latin names and biological classification

In this book the â&#x20AC;&#x2DC;officialâ&#x20AC;&#x2122; biological, Latin names are used for animals and plants. This is necessary because popular names are not available for most species and, moreover, the same species may have different popular names in different countries. The only truely international names, therefore, are the Latin ones. When available and known to us, popular names used in Suriname will be mentioned. Latin names may pose difficulties for some people who may be tempted to skip over them in the text. However, if you pay a little attention to them, you will soon get used to them. When reading this book, try to pronounce the Latin names carefully once or twice and then proceed. After a while, to your own surprise, you may discover that you actually start to remember them. In biology, naming is done according to a classification system that aims to indicate the relationships between living organisms as they are thought to have occurred in evolutionary history. In this system, the species is the basic unit. There have been several definitions of a species, but a useful one is that of a population of individuals that can freely interbreed and produce fertile offspring. For example, consider a horse and a donkey, two separate species. They can interbreed and produce a mule. However, the mule is sterile and cannot produce young and this is the reason that horses are considered a different species from donkeys. If mules were able to produce young, and these could reproduce, a horse and a donkey would have to be, by definition, the same species. To illustrate the principle of naming and classification, compare a big blue butterfly with two yellow ones (Figure 1-01). It is clear that the two yellow ones are not identical to each other and probably belong to different species. A species is given a single, Latin name. The smaller yellow butterfly was first described in 1758 by Linnaeus and he named it sennae, the larger yellow one was named philea in 1763 by Linnaeus and the large blue one was called menelaus by Linnaeus in 1758 (species names are always written in italics by convention). On examination of the three butterfly species, it is clear that the two yellow butterflies are more similar and probably more closely related to each other than either of them is to the blue one. Biologists acknowledge a close relationship between species by grouping them into a category called a genus. So, the two yellow butterfly species are grouped into the same genus, in this case Phoebis. The blue species menelaus is grouped, together with other similar species, 19

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into the genus Morpho (by convention, genus names are also written in italics). When indicating a particular species, it is internationally agreed that both the genus and species names are mentioned (binomial or binary nomenclature). So, when referring to the above-mentioned species we speak of Phoebis sennae, Phoebis philea and Morpho menelaus. Note that the genus name is a noun and the species name an adjective or a noun. Often, the species name is followed by the author that described the species and the year in which he did so. As such, the above-mentioned species should be noted as Phoebis sennae (Linnaeus, 1758), Phoebis philea (Linnaeus, 1763) and Morpho menelaus (Linnaeus, 1758). For brevity’s sake, in this book, species names will be indicated without authors and year except in the species checklist. Of course, relationships are more encompassing than genera (plural of genus). For example, there are also yellow-and-white and white-only butterfly species, belonging to different genera than Phoebis and these are related more to each other than each of them is related to Morpho menelaus. Different, closely related genera are grouped into tribes. By a similar line of reasoning, different tribes are grouped into subfamilies, subfamilies to families, families to orders and orders to classes, classes into phyla (plural of phylum), phyla into kingdoms and kingdoms into domains. In the classification of plants the term division is used instead of phylum. As an example, the formal classification of Morpho menelaus is: menelaus Morpho Morphini (Morpho-like butterflies) Morphinae (Morpho and owl butterflies) Nymphalidae (brush-footed butterflies) Lepidoptera (butterflies and moths) Insecta (insects) Arthropoda (‘jointed foot’ animals: insects, spiders, crabs and others) kingdom : Animalia (animals) domain : Eukarya (all organisms with a eukaryotic cell (e.g. with a cell nucleus) as opposed to e.g. bacteria) Note that only genus and species names are written in italics, other Latin names are written normally. species genus tribe subfamily family order class phylum

: : : : : : : :

In biology the rules for name-giving are very strict. For animals, they are stipulated by the International Commision on Zoological Nomenclature and laid down in the International Code of Zoological Nomenclature, currently the fourth edition, 1999 (ICZN 1999; http://iczn.org). For plants, they are stipulated by the International Commision on Botanical Nomenclature and laid down in the International Code of Botanical Nomenclature, current edition, 20

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2006 (ICBN 2006; http://ibot.sav.sk/icbn/main/htm). There are some slight differences in zoological and botanical rules for nomenclature that do not concern us here. As noted above, the biological classification system aims to reflect the relationships as they have developed in the course of evolution. This means that each category is thought to have contained the ancestral species that has given rise to all species in the category below it and that each category arose from a single ancestral species. For example, millions of years ago, in the class of Insects one species is assumed to have developed from which all butterflies and moths (order Lepidoptera) would later develop. Similarly, the Lepidoptera is supposed to have contained an ancestral species from which later all brush-footed butterflies (family Nymphalidae) originated. The Nymphalidae is supposed to have contained an ancestral species from which later all Morpho and owl butterflies arose, and so on. This holds for each category (Judd 2002). Biologists study the pattern of evolutionary relationships between living organisms, for example by closely comparing organisms and by studying their DNA.There is a massive amount of data supporting the overall correctness of evolutionary history and the classificatory system, although refinements are constantly being suggested and tested (e.g. Simpson 2006, p. 17-48). This means that, as new data become available and insight into evolutionary relationships alters, the classification system may change accordingly. For example, in 2009, based on an analysis of DNA and anatomical data, the classification of the genera within the butterfly family Hesperiidae was altered, presumably to better reflect the course of evolution (Warren 2009). Generally, however, the classification of butterflies has been rather thoroughly studied, more than of most other insect groups (Lamas 2004). A final point on classification: variation within species. If one compares Morpho menelaus from Suriname with e.g. specimens from Venezuela, they do not look completely identical. Yet, they can interbreed and produce fertile offspring (or so it is assumed, to our knowledge this has not been tested). For this type of variation, the concept of subspecies is often used although its overall usefullness is debated among biologists (see e.g. Winston 1999, pp. 323-336). A subspecies is a population within a species with a special characteristic in a different geographical area. By definition, subspecies can freely interbreed and produce fertile offspring (if not, they would be separate species, not subspecies). The name of the subspecies comes after the species name. So, the proper name of a butterfly for which several subspecies have been described, consists of three names: genus, species, subspecies. The subspecies of Morpho menelaus in Suriname is menelaus (it was the first form of menelaus described and, as other forms are being reported, the first described name automatically gets the subspecies name identical to the species name), so the correct name is Morpho menelaus menelaus (species 73 in this book). The yellow butterfly mentioned above, Phoebis sennae, was originally described, probably, from Jamaica in 1758. 21

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In 1777, Cramer described a similar butterfly from Suriname, which he named marcellina. Later, it was found that these are two regional forms of the same species. So, the proper name for the Jamaican butterfly is Phoebis sennae sennae (Linnaeus, 1758) and for the Surinamese one Phoebis sennae marcellina (Cramer, 1777) (species 28). As subspecies can freely interbreed, mixed forms can be found where two subspecies meet (more properly said, where the geographical ranges of two subspecies meet or overlap). This is for instance the case for Heliconius erato (species 147) at the Brownsberg Nature Reserve, where different kinds of beautiful intermediate forms between subspecies erato, amalfreda and hydara can be enjoyed (Plate 52).

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Geography, geology and soils

Suriname is situated on the north coast of South America, between 2o and 6o northern latitude and 54 o and 58 o west longitude. It is bordered by Guyana on the west, French Guiana on the east and Brazil on the south. Its total surface area is 165,942 sq. kms, of which 75-80% is covered with forest. Suriname and the Guiana Shield Suriname is part of the Guiana Shield (Figure 2-01). This is an area of about 2.2 million square kilometers in northeastern South America, roughly bordered by the Amazon, Rio Negro and Orinoco rivers. It comprises parts of Brazil, Colombia and Venezuela, and the countries Guyana, Suriname and French Guiana (Table I).

Figure 2-01 Location and areal extent of the Guiana Shield. Amazonas (small letters), Bolivar: states of Venezuela; Amapรก, Parรก, Roraima, Amazonas (larger letters): states of Brazil; (adapted from Hammond (2005, p. 10).

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Land area (km2)

Table I Estimated area of the Guiana Shield and its breakdown by constituent country

Country

entire country

in Guiana shield

Brazil

8,456,510

Colombia

1,038,710 88,150

French Guiana

% in Guiana shield

% of Guiana shield

1,204,279

14.2

52.6

170,500

16.4

7.5

88,150

100.0

3.9

Guyana

207,710

214,980

100.0

9.4

Suriname

163,270

156,000

100.0

6.8

Venezuela

882,060

453,950

51.5

Guiana shield

2,287,859

19.8 100.0

Note: adapted from Hammond 2005a, p. 11.

The Guiana Shield has been described as a land of ancient rock, poor soils, much water, extensive forests and few people. It contains as much as 34% of the worldâ&#x20AC;&#x2122;s tropical moist forests. Its population is about 1.5 - 2 million, or 0.6 - 0.8 persons per square km (Conservation International 2002, Hammond 2005a). There are many reasons to consider the Guiana Shield a distinct subregion of the Neotropics and there are important differences with the Amazon basin: it is geologically much older with poorer, more acidic soils (see below) and plants and animals that have adapted to these conditions. This is illustrated by the relatively large percentages of endemic species (i.e. species that occur only in the area of the Guiana Shield and nowhere else) (Table II). Table II Estimated diversity and endemicity of

Group

Number of species

Number of endemic

Percentage endemicity

(estimated)

species (estimated)

(%)

20,000

7,000

Birds

975

150

Mammals

282

Reptiles

280

selected groups of flora and fauna of the Guiana Shield

Vascular plants

Amphibians

272

127

2,200

700

Hymenoptera, Meliponinae

Hymenoptera, Vespidae

200

1,500

300

Freshwater fishes

Hymenoptera, Formicidae Isoptera (termites)

225

Lepidoptera, Sphingidae

125

Notes: Hymenoptera: sawflies, bees, wasps and ants; Meliponinae: stingless bees; Vespidae: family of wasps, mainly social wasps; Isoptera: termites; Formicidae: ants; Sphingidae: hawk moths. Data from Conservation International (2003) and Lacomme (2007).

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Figure 2-02 Contour map of the total annual rain-

Butterflies are not shown in Table II, because they have not been sufficiently researched (see, however, p. 139 for a discussion of some groups of Surinamese species in the context of the Guiana Shield).

fall (mm) in Suriname (averaged data from 1961 to 1985; due to the civil war (1986-1992) large parts of the (hydro)meteorological network has been closed down to date); solid dots indicate rainfall

Climate Surinameâ&#x20AC;&#x2122;s climate is tropical with an average temperature of 27.3 o C, an average minimum temperature (mostly at sunrise) of 23 o C and an average maximum temperature around 31 o C. The highest temperatures occur in September and October (with peaks of 38 o C), the lowest in January and February. The average total annual rainfall varies between about 1,650 mm in the low-lying areas in northwest Suriname to about 2,800 mm in hills and mountains in the inland (Figure 2-02) (Hammond 2005b, Nurmohamed 2006). There is substantial monthly variation in rainfall. Figure 2-03 shows

stations (reproduced from Nurmohamed (2006) with permission from the authors).

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500

Figure 2-03 Average monthly rainfall (mm) at four rainfall stations, Suriname (1960-1999); data from

375

Naipal (2004).

250 Nw Nickerie (1960-1999) 125

Cultuurtuin (1900-1999) Tafelberg (1959-1986)

Sipaliwini (1961-1986) J

months

the average monthly rainfall at four rainfall stations in Suriname (Nieuw Nickerie (northwest), the Cultuurtuin in Paramaribo, the Tafelberg in the country’s center and Sipaliwini near the border with Brazil (data from Naipal 2004). Generally, four seasons are discerned: • long rainy season: late April – mid-August • long dry season: mid-August – November-December • short rainy season: December – early-February • short dry season: early-February – end-April. There is, however, great variation from year to year and a season might well start a month earlier or later. Generally, the wettest month is May with 250400 mm rainfall, and the driest month is October, with about 50-100 mm. The relative humidity for Paramaribo is 76-80%. In the forest, relative humidity rises to about 95% at night and during daytime it is at least 75% (e.g. at the Brownsberg 88%) (De Dijn 2007). Geology Generally, geologists distinguish three basic types of rocks: 1. Magmatic or igneous rocks formed because magma (very hot, liquid rock) from the mantle of the earth solidifies at relatively shallow depth or at the surface. Examples are basalt (especially on the sea floor), granite, gabbro and diorite. Igneous rocks occur mainly in the mountainous interior of Suriname. 2. Sedimentary rocks. Through erosion (e.g by abrasion of rocks through wind and/or water), sand, gravel, etc., are transported, often by the sea or rivers, and deposited elsewhere. Sediments are formed by accumulation of this loose material and, through hardening, consolidation may occur. Examples are limestone and sandstone. Sedimentary rocks are found in the Tafelberg, the savanna belt and coastal plain of Suriname. 3. Metamorphic rocks. These are rocks of the above two types, which have subsequently been subjected to very high pressures and temperatures. During the metamorphosis new minerals may have been created or the 26

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Figure 2-04 Map of the coastal area of Suriname

existing ones may have been reordered. Examples include slate, schist, gneiss and marble (resulting from limestone). Metamorphic rocks in Suriname are commonly found in the interior, e.g. the Bakhuis mountains and the Greenstone Belt (Marowijne Group). The latter unit hosts important goldbearing rocks in Suriname.

showing, from north to south, young coastal plain, old coastal plain, the savanna belt and the mountainous interior (reproduced from Wong (1998) with permission from the Royal Netherlands Academy of Arts and Sciences).

The geology of Suriname is covered in detail in Hammond (2005b, p. 15-48) in the context of the Guiana Shield and in Wong (1998, p. 15-127). Geomor phologically, Suriname can be divided into three regions (Figure 2-04): 1. the inland mountainous region (more than 80% of Suriname). 2. the savanna belt (Zanderij Formation). 3. the coastal plain, subdivided into a young coastal plain in the North and an old coastal plain to the south of the young coastal plain. The coastal plain and savanna belt vary in width from 180 km in the west to 20 km in the east. Ad 1: the inland mountainous region This part, covering 80% of Suriname, consists of igneous and metamorphic rocks, which were formed in the Precambrian era. They are generally 2000 2200 million years old. Through erosion, the top layer has largely worn away. Where this has not happened or happened to a lesser extent, mountains are found, with the highest peak the Juliana Top (1280 m) in the Wilhelmina mountains. Igneous rocks are also exposed as smooth hump-shaped rocks in the rapids (sulas) (Figure 2-05). However, most of the surface of the Precambrian rock is strongly weathered to a depth of up to 40 m. This Precambrian terrain was covered in the past (1900 million years ago) with sedimentary rocks of at least 1000-2000 m thick, which have been subsequently eroded away. Remnants of this are the Roraima Formation (tepuis 27

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Figure 2-05 Igneous rocks at the Raleigh falls in the early morning (BGB, 06-09-2009).

or table mountains) in Venezuela and Guyana. The most eastern occurrence is the Tafelberg in Suriname (1080 m), which mainly consists of sandstones and conglomerates. Approximately 60 million years ago the southern edge of the Guiana Shield was lifted (this is probably still going on), forming a watershed between the Amazon Basin and the basins of the Guianese rivers. Later, this formed the basis for the establishment of the Brazilian-Surinamese border. After millions of years of erosion the watershed is now at 1000-2000 m altitude. Ad 2: the savanna belt (Zanderij Formation) This unit was formed in the Pliocene (5.3 â&#x20AC;&#x201C; 2.6 million years ago), when quartz particles were released from the granitic surface of the interior. The eroded material formed large alluvial fans or was transported by rivers and creeks and a large sheet of sandy sediment was deposited on the northern edge of the Guiana Shield. The sands underwent severe leaching resulting in predominantly white quartz grains forming the Zanderij Formation, on which the savanna belt of Suriname developed.

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Ad 3: the coastal plain The Amazon river carries a constant flow of fine sediment, mainly formed by erosion of the Andes mountains, into the Atlantic Ocean. This material, consisting of a mixture of clay and very fine sand, is transported westward by the Guiana Current towards the Orinoco river in Venezuela through a system of extended migrating mudflats, alternating with sandy shores with cheniers (beach ridges). This mudflat migration along the shore of the Guianas takes place with a spacing of 45 km between the fronts of adjacent mudflats and a migration rate of about 1.5 km/year. The coast experiences a cyclicity of about 30 years. The retained sands, and often also shells, accumulate and form near shore sand bars which are pushed towards the high-water line by incoming waves, thereby forming cheniers that migrate over the clay deposits of the mudflats. The number of ridges decreases towards the West. This process started in the Pleistocene, approximately 1 million years ago when the Old Coastal Plain was formed, continuing over the last 10,000 years (Holocene era, during the formation of the Young Coastal Plain). The sand from the beaches (Galibi, Matapica) is mainly derived from the Marowijne river and French Guiana. Therefore, the sandy beaches are mostly situated in the east of Suriname. Soils Generally speaking, soils of the wet tropics are acidic (pH 3.5 - 6), poor in available nutrients (especially phosphorous) and calcium, with high levels of aluminium and iron. It is generally accepted that the main factors in soil formation are the mineral composition of the parent material (e.g. granites), the dynamics of water, acid chemistry, biological action, human activity and the duration of weathering and erosion. Each soil type is the product of its own unique life history (Hammond 2005b, p. 48-64). During the last 65 million years, strong climate changes took place and numerous periods of wet tropical forests with little or no erosion were alternated with dry periods of savannas with periodic heavy rains, which caused severe erosion. During wet periods, degradation products from vegetation (e.g humic acids) penetrated deep into the rock and reacted with it, turning it into soft, so-called rotten rock. In a subsequent dry period, when there was little vegetation due to savanna formation, the rock was further eroded by the heavy rains. Through these processes over millions of years, the soil in the mountainous interior has become highly leached, poor and infertile. During the leaching of the rock, the poorly soluble oxidized iron accumulated, forming red soils. In places with stagnant groundwater, hard laterite caps developed, especially consisting of hematite (Fe2O3), which protected the underlying soil and rotten rock against further erosion. An example of this is the Brownsberg (Figure 2-06). Under similar climatological conditions, aluminum silicates were altered into Al2O3. High concentrations of this 29

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Figure 2-06 Reddish brown colored soil at the Brownsberg (HBPEG, 08-08-2008).

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Figure 2-07 View of the Voltzberg. The height gives

mineral are called bauxite. These bauxite concentrations also formed hard caps, topping several plateaus, e.g. Mungo area (see also plinthosols below). The two main types of soils of the mountainous interior of Suriname are ferralsols and acrisols. Ferralsols are deep soils typified by a highly weathered subsurface dominated by low-activity clays, such as kaolinite with a relatively high iron content. Acrisols are also deep soils characterized by a dense clay layer just below the surface. They are generally associated with younger, less highly weathered parent material, and a clay subhorizon developed largely through downward clay migration of deposited materials, rather than those formed from rock decomposition. They contain higher concentrations of weathered minerals and active clay particles than the ferralsols, though there is overlap and the two soil types are often tightly interwoven. In places where no forests could develop, e.g. by severe drought sensitivity,

an indication of the original minimum height of the igneous rock in the mountainous interior (Frans Barten, 17-07-2006).

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Figure 2-08 The Kasikasima mountains and surrounding rainforest (Mat Wijnen, 22-10-2010).

there was less formation of rotten rock and erosion, and â&#x20AC;&#x2DC;kopjesâ&#x20AC;&#x2122; or inselbergs formed. Examples are the Voltzberg (Figure 2-07) and Kasikasima mountains (Figure 2-08). These inselbergs, therefore, give an indication of the original minimum height of the igneous rock in the mountainous interior. The Zanderij Formation, constituting the savanna belt, has as main soil types arenosols (about 17%) and clay and silt-containing ferralsols (the remainder). Arenosols consist largely of coarse sand with relatively small amounts of clay, silt or loam, and no rock fragments. In extreme cases (albic arenosols: the white, bleached sands) this extends to a depth of several metres and the sand has little, if any, water-holding capacity, so that dry seasons result in virtually no moisture remaining in the top layers. Other arenosols in slopes and valleys show an increase in clay content in depth, but remain highly infertile. The coastal plain is fertile. The sediments from the Amazon river are relatively fresh and have not yet been exposed to severe weathering and leaching. Nowadays, roads and villages cover the ridges and crops are grown there. Along the river banks, where there were once sugar cane, coffee and cocoa

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plantations, there are now polders with citrus, banana and rice. The Old Coastal Plain is less fertile, and there is more extensive agriculture and livestock. The main soil types of the coastal plain are gleysols, histosols and plinthosols. Gleysols and histosols belong to the alluvial soils (soils that form where flooding is permanent or seasonal and fresh material is regularly deposited and removed). Gleysols are formed where flooding occurs for most of the year. It is present in a narrow band in the Young Coastal Plain. Histosols are gleyosols that accumulate organic, vegetative matter to form a humus-rich upper layer, 2-3 m in Suriname. They are present in the coastal swamp areas. Plinthosols (‘groundwater laterites’, ‘scholsoils’) are characterized by a hardpan at or near to the surface (< 100 cm depth), formed by the cementation of iron or aluminium-rich kaolinite (plinthite). This hardpan, better known as laterite, is linked to clay and iron rich soils that have been exposed to fluctuating water levels on a low-lying sedimentary plain. Soils and soil formation are dealt with in Hammond (2005b, pp. 48-66) and in Wong (1998).

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Part III The butterflies of Suriname â&#x20AC;&#x201C; introduction and history

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The butterflies of Suriname â&#x20AC;&#x201C; an introduction

Now that the stage has been set by briefly describing biological classification and naming, the geography, geology, soils, plants and main habitats of Suriname, let us focus on the subject proper of this book: butterflies. After an introduction on butterflies and moths, butterfly families and the functional anatomy of butterflies is dealt with, followed by sexual differences, the life cycle, butterfly migration, the enemies of butterflies and their means of defense. Finally, the butterflies of Suriname are discussed in the context of the world and neotropical fauna. This is followed by an account of the history of the study of the Surinamese butterflies. In part IV, a checklist of the 150 butterfly species dealt with in this book is followed by the species accounts. Butterflies and moths Butterflies, together with moths, are insects (insects have six legs, in contrast to e.g. spiders which have eight) of the order Lepidoptera. In ancient Greek lepidos means scale and pteron means wing. The wings of butterflies and moths are covered with scale-like hairs, like the tiles of a roof (Figure 7-01).

Figure 7-01 Detail of the ventral side of female Caligo teucer teucer (Morphinae), showing covering of the wing with scale-like hairs (HBPEG, 02-06-2011).

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It is difficult to give accurate numbers of species of moths and butterflies, since new species are described regularly. Worldwide there are about 18,000 to 20,000 different species of butterflies. Larsen (2005) estimated the world total at 18,460 species and Heppner (1991) at 19,238. The number of moths is even more difficult to estimate as they comprise a large number of families, many of which are insufficiently known. Heppner (1991) calculated the number of known species in 1991 at 127,039 and estimated the world total number of Lepidoptera to be as high as 255,000. In 1998 he suggested it might well be even higher (Heppner 1998). The number of undescribed moth species is far and far greater than the number of undescribed butterfly species. If one assumes that 50% of moth species and 90% of butterfly species are known to science at present, it can be deduced that the likely number of moth species is 7.6 â&#x20AC;&#x201C; 11.6 times the number of butterfly species. The total number of butterfly and moth species of Suriname is not known. The present authors conservatively estimate that Suriname has 1,325 species of butterflies (see Table X, p. 137) and, consequently, should have 10,070 â&#x20AC;&#x201C; 15,370 moth species. How to distinguish between a butterfly and a moth? Most people associate butterflies with bright colors and daytime activity, and moths with dark brown-grey colors and nighttime activity; however, there are many exceptions to this. Biologists distinguish about 121 different families of Lepidoptera, of which only six comprise butterflies and the remainder all moths (Heppner 1991, Scoble 1995). For everyday purposes, some charcteristics are listed in the table below. Table VI Some characteristics to distinguish

Butterflies

Moths

butterflies from moths.

usually diurnal

usually nocturnal

usually not attracted by light at night

attracted by light at night

usually brightly colored

usually drab colored

antennae clubbed or hooked

other antennae, e.g. pointed, feathered

A reliable, easy and quick way to distinguish between butterflies and moths is by examination of the antennae (Figure 7-02). Butterflies have clubbed or hooked antennae, whereas any other antennal form indicates a moth. Only one family of moths with a handful of species in Suriname, the Castniidae, also have clubbed antennae. A perhaps surprising result is that some species, popularly called butterflies in Suriname, are actually moths. Examples are: Urania leilus (witbroek, family Uraniidae) (Figure 7-03), members of the Castniidae (Figure 7-04), of which the caterpillars bore into palms or sugar cane and are known as pests, and Melanchroia subnotata (family Geometridae) (Figure 7-05), which feeds on the wild birambi (Phyllanthus species).

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Figure 7-02 Some antennal forms among Surinamese Lepidoptera; a: hooked (Pyrrhopyge amyclas amyclas, Hesperiidae); b: clubbed (Battus polydamas polydamas, Papilionidae); c: pointed (Urania leilus, Uraniidae); d: pointed (Melanchroia subnotata, Geometridae); e: feathered (Automeris liberia, Saturniidae).

Figure 7-03 Witbroek (Urania leilus, Uraniidae).

Butterfly families In this book the butterfly species are presented according to family (Lamas 2004, see Vane-Wright (2003) for an overview of butterfly systematics). The major characteristics of the families for Suriname are as follows (see also Table X, p. 137): 87

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Figure 7-04 A day-flying moth, Eupalamides guyanensis (Castniidae) (BGB, Paramaribo, 06-01-2005).

Figure 7-05 Melanchroia subnotata (Geometridae).

1. Hesperiidae (skipper butterflies; Plates 12 and 13): usually rather small, dull, moth-like butterflies with a stout body, the head wider than the thorax, and hooked or pointed antennae; three subfamilies (Eudaminae, Pyrginae, Hesperiinae); 426 species recorded in Suriname. 2. Papilionidae (swallowtail butterflies; Plates 14-18): large butterflies, often black or yellow with red; keep their wings fluttering when drinking nectar or minerals; use six legs for walking; one subfamily (Papilioninae); 33 species in Suriname. 3. Pieridae (whites and sulphurs; Plates 19-22): medium-sized or small butterflies, mostly white or yellow; use six legs for walking; three subfamilies (Dismorphiinae, Coliadinae, Pierinae); 31 species in Suriname. 88

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4. Lycaenidae (blues and coppers; Plates 23 and 24): small butterflies, usually with a blue upperside and small tails on their hindwings; the females use six legs for walking and the males four; two subfamilies (Theclinae, Polyommatinae); about 191 species in Suriname. 5. Riodinidae (metalmarks; Plates 25 and 26): small delicate butterflies of various appearance; the females use six legs for walking and the males four; two subfamilies (Euselasiinae, Riodininae); about 360 species in Suriname. 6. Nymphalidae (brush-footed butterflies): large to small butterflies; use four legs for walking; 284 species in Suriname; represented in Suriname by 11 subfamilies: a. Libythaeinae (snout butterflies): one species in Suriname, not treated in this book. b. Danainae (monarch butterflies;Plate 27): orange-black butterflies with eversible hair pencils in abdomen and/or black androconial patches on the hindwings; four species in Suriname. c. Ithomiinae (glasswing butterflies;Plates 28 and 29): medium-sized, partly transparant or orange-black-yellow butterflies; the males have long hair-like androconial scales on the hindwings; about 35 species in Suriname. d. Morphinae (morpho and owl butterflies; Plates 30-36): morpho butterflies are large, often shiny metallic blue and spectacular (ten species in Suriname); owl butterflies usually fly at dusk, are often large, somberly colored with large eyespots on the underside (about 26 species in Suriname). e. Satyrinae (the browns; Plates 37-39): medium-sized, usually brown butterflies with eyespots (ocelli) on the underside; about 68 species in Suriname. f. Charaxinae (Charaxes butterflies; Plates 40 and 41): medium-sized to large, strong, swift-flying butterflies with a stout body, large palpi and a short proboscis, often blue and black colors; about 30 species in Suriname; g. Biblidinae (small pages; Plates 42-44): medium-sized, often beautifully colored butterflies; about 50 species in Suriname. h. Apaturinae (the emperors): medium-sized butterflies, two species in Suriname, not treated in this book. i. Nymphalinae (the peacocks and buckeyes; Plates 45-47): medium-sized butterflies, not easily classifiable into another nymphalid group; about 26 species in Suriname. j. Limenitidinae (the admirals; Plate 47): medium-sized orange-brown butterflies, about 12 species in Suriname. k. Heliconiinae (passionflower or longwing butterflies; Plates 48-52): butterflies with elongated forewings, large eyes and long antennae; 30 species in Suriname. 89

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Figure 7-06 Major anatomical features of Protographium agesilaus autosilaus (Papilionidae): a: antenna; b: compound eye; c: extended proboscis; d: first leg; e: last leg; f: abdomen; g: hindwing tail; h: hindwing; i: forewing; note that six legs are used for walking (Frans Barten, 26-07-2006).

Form and function of butterflies A butterfly has three main parts: head (with antennae, eyes and mouthparts), thorax (with six legs and four wings) and abdomen with the digestive and reproductive tracts. (Figure 7-06). The head The antennae are sensory organs that are used for finding food, for locating a sexual partner and during mating. In addition, female butterflies, e.g. Ceratinia neso nisea (species 70), can be seen, just before laying an egg on a leaf, to carefully probe the surface of the leaf with the antennae. Most probably, they are checking the chemical make-up of the leaf and whether the plant is the correct one to lay their eggs on. The eyes of butterflies are very different from human eyes. They are of the compound type, made up of numerous so-called ommatidia. Each ommatidium functions as a kind of miniature eye, in which light is transmitted that generates an electrical impulse to the brain. The number of ommatidia per eye vary greatly between species of Lepidoptera, from about 200 to 27,000 (Scoble 1995). The compound eye is very sensitive to movement, colors and light, but is unsuitable for sharp vision because it is incapable of focusing. Butterflies have relatively large eyes and their field of vision is about 300 degrees, so they can almost literally â&#x20AC;&#x2DC;watch their backâ&#x20AC;&#x2122; (Rutowski 2003, Briscoe 2003). The mouthparts mainly consist of a proboscis, a hollow tube (like a drinking straw), which is coiled beneath the head. When feeding it is uncoiled and can then reach surprising lengths. For example, the uncoiled proboscis of the 90

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Figure 7-07 Male (a, c, e) and female (b, d, f) first legs of Nymphalidae species; a, b: Adelpha iphiclus (Limenitidinae); c, d: Hamadryas februa (Biblidinae); e, f: Memphis acidalia (Charaxinae) (drawing by AFE. Neild. Reproduced from Neild (2008) with permission from the author).

skipper Pyrrhopyge amyclas amyclas (species 5) is about as long as its body. Because a proboscis functions like a straw, a butterfly is restricted to a liquid diet: nectar from flowers, sap from rotting fruit or trees, etc. The thorax The thorax has three segments, to which six legs, two forewings and two hindwings are attached. In the Papilionidae, Pieridae and Hesperiidae all six legs are used for walking, but in the Nymphalidae walking is done on four legs. The two first legs are reduced in size and are used by the female butterfly as a sensory organ. With special spurs on the first legs, she can prick into a substance (e.g. leaf tissue) and ‘taste’ it. Female butterflies do this to identify the proper plants to lay their eggs on (Figure 7-07). Butterflies have four wings: two forewings and two hindwings, which are attached to the thorax by strong muscles. The wings consist of a thin membrane covered with scale-like hairs. In the vast majority of species, pigments in these hairs give wings their colors (Figure 7-01). The brilliantly colored Morpho (species 71-76, Plates 30-33) get their color not by pigments, but by a modified anatomical structure of the wing scales, reflecting light in such ways that humans perceive blue, purple and greenish hues depending on the angle of view. There are veins in the wings, through which haemolymph flows, providing nourishment to the wing tissues. The pattern of veins is widely used in the classification of Lepidoptera (e.g. Heppner 1998). Biologists have given symbols to each of the veins and names for different wing areas. There are basically two systems for this: Herrich-Schäffer’s, in which the veins are numbered from 1 to 12 (Herrich-Schäffer 1843-1856) and Comstock’s, in which the veins have been given various names and symbols (Comstock 1918). We will use the Comstock system with some generally accepted modifications (Figure 7-08). The technical terms are avoided as much as possible throughout this book, but in the identification sections of the species they are often necessary. The abdomen The digestive and reproductive organs are located in the abdomen. The abdomen consists of ten segments; the first seven or eight contain the digestive system 91

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Figure 7-08 Wing venation and wing areas. a. wing venation of a butterfly, following the modified Comstock system of numbering; Sc = subcostal vein; R = radial vein; M = median vein; Cu = cubital vein; A = anal vein; the costal vein forms the costal wing margin and is not shown separately; the small forked vein that branches of the hindwing Sc+R1 is the

and the last two or three the genitals. Usually, the abdomen is quite slender, but in butterflies that have eaten from a banana bait the first seven or eight segments can be enormously expanded, even to the extent they can hardly fly. Female butterflies more often have a distended abdomen due to a large number of eggs (see below). The reproductive system in butterflies is quite complicated. The anatomy of the genitalia is widely used in classification, but is beyond the scope of this book.

humeral vein; b: names of the major wing areas referred to in this book (drawings by PJ de Vries; reproduced from DeVries 1987 with permission from Princeton University Press).

Male versus female butterflies There are a number of ways to distinguish male from female butterflies, some of which occur in all species; others are species, genus, subfamily or family specific. 1. In many species, males and females have a quite different appearance (sexual dimorphism). For example, male Heraclides androgeus androgeus have bright yellow-black wings, whereas the females are black with iridescent green or greenish-blue (Plate 17, Figure 5-8). In Itaballia demophile de-

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Figure 7-09 External appearance of genitalia in male and female butterflies; a: male Lycorea halia halia (Danainae); note protruding penis (aedeagus); next to it on both sides, are the clasping organs (valvae); b: female Danaus plexippus megalippe (Danainae); there are three openings: the anus, the copulatory pore and the egg pore.

mophile, the males are white on the upperside and the females predominantly black (Plate 22, Figure 1-4). In species where the sexes are more alike, female butterflies are usually a little larger with duller colors and more rounded wings, e.g. Archaeoprepona demophon demophon (Plate 41, Figure 1-4) or Dryas iulia alcionea (Plate 49, Figure 1-3). Appearance of the abdomen. Generally, males have a slender abdomen with the genitalia at the end consisting of two valves (the claspers, to hold the female during mating) and an aedeagus or penis in between. Females are often filled with eggs which give the abdomen a stuffed, rounded appearance. The abdomen ends in three openings: the anus, the copulatory pore used for mating, and an egg pore used for laying eggs (Figure 7-09). The anatomy of the front legs (Figure 7-07). In the Lycaenidae (Plates 23 and 24) and Riodinidae (Plates 25 and 26) the front legs of the males are strongly reduced in size, whereas the females use six legs for walking. In the Nymphalidae (Plates 27-52) the two front legs are not used for walking in both sexes. In the males the two front legs have no function and are rudimentary. In the females the front legs have evolved into sensory organs (see above). The anatomy of the front legs is a reliable way to distinguish males from females in species where sexes are similar (e.g. Chloreuptychia chlorimene (Plate 38, Figure 5 and 6), Junonia evarete (Plates 45 and 46) or Telenassa fontus fontus (Plate 47, Figure 9-12), or when doubt remains as to the sex after inspection of the abdomen. In most cases a microscope is needed. Secondary sexual characteristics. Pheromones are chemical substances that are produced in an organism and induce certain behavior in other individuals of the same species. In butterflies, pheromones are mainly related to reproduction. Females produce pheromones to lure males and male pheromones make the female respond to the maleâ&#x20AC;&#x2122;s â&#x20AC;&#x2DC;courtingâ&#x20AC;&#x2122; and to induce her to mate (Scoble 1995). Male butterflies often have specialized scales 93

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Figure 7-10 Some examples of secondary sexual characteristics in male butterflies; a: white androconial hairs on fold of dorsal hindwing of Parides sesostris sostris (Papilionidae); b: pink androconial patch on ventral forewing of Enantia aloikea (Pieridae); c: light yellow androconial patch on dorsal forewing and hindwing of Aphrissa statira statira (Pieridae); d: black androconial patch on dorsal forewing of Arawacus aetolus (Lycaenidae); e: eversible abdominal hair pencils of Lycorea halia halia (Danainae); f: black androconial patch on dorsal hindwing of Danaus plexippus megalippe (Danainae); g: grey androconial hair-like scales on dorsal hindwing of Hyposcada dujardini dujardini (Ithomiinae); h: androconial hair tuft on inner margin of dorsal hindwing of Caligo illioneus illioneus (Morphinae); i: yellow androconial hair tuft on inner margin of dorsal hindwing of Pierella lena lena (Satyrinae); j: yellow androconial hair tuft on inner margin of dorsal hindwing of Agrias narcissus narcissus (Charaxinae); k: light androconial band near costal margin of hindwing of Dryas iulia alcionea (Heliconiinae); l: grey androconial band near costal margin of hindwing of Neruda metharme metharme (Heliconiinae); m: light androconial band near costal margin of hindwing of Heliconius melpomene meriana (Heliconiinae); n: grey androconial band near costal margin of hindwing of Heliconius erato hydara (Heliconiinae).

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or hairs on their wings or sometimes other body parts (androconia) that store and emit pheromones (Figure 7-10). Many male Parides butterflies (Plates 15 and 17) have androconial folds with white hairs on their hindwings. Male monarch butterflies (Danaus plexippus megalippe and Danaus eresimus eresimus, Plate 27) have black and Pierella species (Satyrinae; Plates 37 and 38) may have yellow androconial patches on their hindwings. Many Satyrinae (Plates 37-39), Hesperiidae (Plates 12 and 13) and Lycaenidae (Plates 23 and 24) have androconial streaks or patches on their forewings. Male Ithomiinae (Plates 28 and 29) have androconial hairs and Heliconiinae (Plates 48-52) an androconial band on their hindwings at the junction with the forewings. Many Charaxinae (Plates 40 and 41) have androconial tufts (bunches of hairs) on their hindwings. All Surinamese Danainae (Plate 27) have eversible androconial hairs at the end of their abdomen. 6. Perhaps most importantly, male and female butterflies behave differently and the way they behave differs between species (Wiklund 2003). a. Males have one ultimate goal in life: to mate with as many females as possible. To do this, they must (besides feed and survive) compete with other males, find the females and entice them to mate. Males do this in different ways. Heraclides thoas thoas (species 23, Plate 18), as well as many whites (Plates 19-22), fly about and check anything that looks vaguely female. Male Morpho menelaus menelaus (species 73, Plate 31) flies in a fairly straight line along tracks in the forest in search of females. Many male butterflies (e.g. many Nymphalidae (Plates 27-52), Lycaenidae (Plates 23 and 24) or Riodinidae (Plates 25 and 26)) are overtly territorial. They claim a certain territory for themselves, sit and watch at an advantageous spot and may attack anything in their vicinity. Hamadryas butterflies for instance (Plates 43 and 44) sit head down with their wings spread on tree trunks and attack any other approaching butterflies or unsuspecting tourists (Figure 7-11). Territorial contests between male Hamadryas are accompanied by a clear crackling sound made by the fighting butterflies. Butterflies are, to an important extent, visual animals and, in addition to pheromones, visual signs are most probably important in the determination of which male is finally allowed to mate with a female. So, males look flashy, conspicuous and show themselves readily. The dark side for them is that predators (e.g. lizards, birds) also readily spot them and, consequently, males generally do not live very long. Being flashy has its price to pay. b. Female butterflies have another ultimate goal in life: (after having mated) to lay as many eggs as possible on the right plant to ensure the survival of the species. So, generally, they are better camouflaged than males, are less flashy with subdued wing coloration, show themselves less readily and lead a more hidden life, flying from plant to plant, examining the plants with their antennae and front legs, and laying eggs. Generally, they live longer than males.

Figure 7-11 Male Hamadryas arinome arinome (Biblidinae) perching head down on a tree trunk (HBPEG, Brownsberg Nature Reserve, 23-10-2009).

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Figure 7-12 Butterflies mating; a: Parides lysander lysander (Papilionidae) (BGB, Lelydorp butterfly farm, 01-032006);

b: Danaus plexippus megalippe (Danainae) (BGB,

Paramaribo, 23-03-2005).

The life cycle of butterflies A female butterfly lays an egg, from which a tiny caterpillar (larva) emerges. After having grown sufficiently, the caterpillar pupates and from the pupa a fresh butterfly emerges. Each butterfly species completes this life cycle in its own characteristic way: the plant on which the egg is laid, the appearance of the eggs, the foodplant, the way the larva deals with the plantâ&#x20AC;&#x2122;s defenses, the appearance and behavior of the caterpillars, the way they pupate, the appearance of the pupa and, of course, the adult butterfly and the way it behaves; all of this is characteristic of a peculiar species. For the vast majority of the Surinamese butterflies the life cycle in Suriname is unknown. Throughout this book we illustrate aspects of the life cycle, if known to us, and report on basic characteristics of each species. Some general information on the life cycle follows here. a. Courting and mating. As noted above, both male and female butterflies produce pheromones to ensure mating. Typically, female pheromones work at long range: the male detects the pheromones with his antennae and follows the chemical trail towards the female. When close, he sprays her with his own pheromones, after which they mate (Figure 7-12). Inside

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Figure 7-13 Mechanitis polymnia polymnia (Ithomiinae), laying a cluster of eggs (BGB, Catharina Sophia, Saramacca, 17-09-2005).

the female body, the sperm is temporarily stored and subsequently the eggs inside the female’s abdomen are fertilized. Overviews can be found in Deinert (2003) and Wiklund (2003). b. Eggs. The female lays her eggs on the foodplant or, in some species, in the vicinity of the foodplant. The decision where to lay the eggs is an extremely important one. If the female picks the wrong place, the chances of survival of the larva are very small indeed. There are great differences in egg-laying (Chew 1984). Some species, e.g. Hesperiidae (species 1-10), do so with great speed, only momentarily interrupting their flight, whereas others appear to be very careful. Females may flutter around a potential plant for considerable time, examining leaf after leaf, sometimes walking up and down the leaf. Some distinguish between old and fresh leaves and microclimatic factors such as shade, sunlight, wind, temperature probably also play a role. The female probably identifies the foodplant visually, but also chemically. She probes the leaf with her antennae, ‘tastes’ the leaf with the sensory organs on her front legs and some species probe the leaf surface with their proboscis (Singer 1984, 2003). Most butterfly species lay one egg per leaf or per plant, others lay clusters of eggs (e.g. Mechanitis polymnia polymnia (species 64, Figure 7-13), Heraclides anchisiades anchisiades (species 19; 97

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Figure 7-14 Eggs of Caligo illioneus illioneus (Morphinae) on banana leaf (BGB, Paramaribo, 17-09-2005).

Figure 11-14a) or Brassolis sophorae sophorae (species 78). The eggs hatch, sometimes after a few days, sometimes after several months in species that only occur as adults in a certain period of the year. The eggs have an opening on top, the micropyle, through which the sperm enters the egg in the female body for fertilization and through which later, possibly, the larva breathes. Eggs often have a delicate, sculpture-like structure (Figure 7-14). Generally, the form and appearance of eggs differs in different butterfly families. E.g. the eggs of Hesperiidae (species 1-10) and Papilionidae (species 11-23) are round whereas eggs of Pieridae (species 24-37) are spindle-shaped and eggs of Riodinidae (species 45-55) are often hairy. c. Caterpillars. When the tiny caterpillar hatches from the egg, it first eats the egg shell and then starts to feed on the foodplant. As it grows, it sheds its skin (molts) several times, enabling it to grow bigger. The body of a caterpillar between two molts is called an instar (the first instar is the caterpillar that emerges from the egg). Until the pupa is formed, the caterpillar usually molts four times, so most species have five instars. Molting is necessary mainly for developing a larger head. The body is soft and flexible, enabling it to expand as it grows, but the head is enclosed in a head capsule which has a fixed size. Just before molting, the caterpillar stops eating and a new, soft skin is made underneath the old one. Then the head capsule splits, the caterpillar (the new instar) wriggles out and usually eats its old skin except for the head capsule and other hard parts (Figure 11-18d). Before the new head capsule hardens, the head expands considerably and the caterpillar is ready for further growth. Like the butterfly, the caterpillar has three main parts, the head, thorax and abdomen (Figure 7-15). On each side of the head there are a number simple eyes (mostly three), called ommatidia, stemmata or ocelli (Figure 7-16b). 98

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Figure 7-15 Caterpillar of Agraulis vanillae vanillae (Heliconiinae); a: head with horned head capsule; b: thorax; c: abdomen; d: anal end; e: true leg; f: proleg (BGB, Paramaribo, 21-10-2007).

The front of the head is made of hard material (chitin) and is called the head capsule (Figure 7-16a). At the base there are strong mouthparts and short antennae, probably used for food discrimination. The silk-spinning organs (spinnerets) are situated near these (Figure 7-16c). Caterpillars almost continuously make sure they are connected to the plant by means of silk. This prevents them from accidentally falling off the plant (which would mean death if they are unable to find a suitable plant again), enables them to follow other larvae and find their way to a communal nest in certain 99

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Figure 7-16 Head of mature caterpillar of Caligo illioneus illioneus (Morphinae); a: head capsule with horns; b: simple eyes (ommatidia, stemmata or ocelli); c: silk-spinning organ (spinneret) (BGB, Paramaribo, 06-01-2008).

species (e.g. Brassolis sophorae sophorae, species 78), and provides a means to escape from predators (see below). Prior to pupation the larva spins a silken pad to which the later pupa will be attached. The thorax consists of three segments and has three true legs. The abdomen consists of ten segments. The third through the sixth and the tenth segments bear a pair of false legs or prolegs. These are extensions of the body wall with little hooks or crotchets at the underside that can hook into the plant part to prevent detachment. The number of false legs, the number and structure of the crochets and the body segments that bear them can be of help in distinguishing the caterpillars of butterflies from larvae of other families. E.g. the vast majority of larvae of Geometridae, a large family of moths with more than 20,000 species worldwide and more than 7,900 species in Central and South America (Heppner 1991), have only two pairs of prolegs, situated on the sixth and tenth abdominal segments, causing them to walk in a â&#x20AC;&#x2DC;curlyâ&#x20AC;&#x2122; fashion (Wagner 2005) (Figure 7-17). Functionally, a caterpillar consists of a big gut with a mouth at the front and an anus at the end. Caterpillars have three tasks in life: eat and grow, avoid being eaten and develop into a pupa. Therefore, many feed at night and hide during the day. Various 100

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Figure 7-17 Caterpillars of Melanchroia subnotata (Geometridae) on gooseberry tree (Phyllanthus elsiae) (BGB, Paramaribo, 11-01-2009).

Figure 7-18 Pupa of Danaus plexippus megalippe (Danainae) (BGB, Paramaribo, 16-11-2007).

adaptations for the survival of caterpillars will be discussed in the next section. d. Pupae. Just before pupating the caterpillar stops eating, voids its guts, sometimes changes its color and starts walking in search of a place to pupate. This stage of the life cycle is called the prepupal. So, if you see a caterpillar on the move, often it will pupate shortly. The prepupa attaches itself to a substrate (e.g. a twig or stem) by a silken pad at its rear end and transforms into a pupa. Pupae, as well as caterpillars, come in many forms and colors and are usually cryptic (see below). The development of the actual butterfly is very complex indeed. Inside, caterpillar tissues are broken down and new tissues forming the adult butterfly are built up (Figure 7-18). The pupal stage lasts 1-2 weeks or sometimes months. Then, the pupal skin splits, the butterfly wriggles out and hangs on it for a few hours, so that its wings can expand and harden. After this, the butterfly is ready for action. 101