Spiders are of major importance in ecosystems and are recognized as effective re ee en eee cee Sa ore ee been described but the actual number of species is expected to be many times ras ‘This book provides a concise overview and descriptions of the 107 spider fami- lies that are presently recognized. Ie contains identification keys to the families and to the different kinds of spider webs, and shortcuts to remarkable types of spiders. The book fills a gap that has existed for a long time since the previous such overview dates back from more than a century Both authors of this book are professional arachnologists who have devoted ‘most oftheir careers tothe study of spiders, both froma biological/agroecolog Tere tent en ec Dr Rudy Jocqué is head of the Invertebrates (non-insects) Section in the department of Zoology at the Royal Museum for Central Africa, Tervuren, Belgium. Prof. Dr Ansie Dippenaar-Schoeman is head of the Arachnida Unit in the Biosystema- tics Division at the Plant Protection Research Institute, Agricultural Research Council ‘and extraordinary professor in the Department of Zoology and Entomology at the eee re ace SI foomenod wT a esteemed ARC - PPRI BN 90-75894.35.5, ll Peasy cee) Carrey EER RIA REC ELS f SPIDER FAMILIES | OF THE WorRLD Carat AMO SariatSPIDER FAMILIES OF THE WORLD R. Jocque A.S. DUPPENAAR-SCHOEMAN Ses] Astisa mm, wT TERVUREN SEL comae ‘ARC - PPRI© 2006, Royal Museum for Cental Aca T3,Lemeneneeneg 3080 Tervuren (Begum) ‘wowaliccmtern bs ISBN: 90.7048. Legal Deposit: Dy2005/0254)07 Core: Nadine Van Noppen (pctre) and Sony Van Hoedks, RMCA (aout) agout Piing: Peters Belgas, Aight reserve No part ofthis publation maybe reproduce, stored na eres system or ransmited ing form or any mean ceetroni, mechanical photocopying, ecoding or otherwise without eae ‘len permission of he Royal Museum for Cereal Ae Preamble Foreword o Acknowledgements Iroducton, Spiders and their interest. Morphology and terminology Glossary = Keys — “The main sections. 1. Mygalomorphae 2 Cribelaes. 3 Lessthan 8 ges. 4 2eaws 5. Sclavs Key to spider webs, Phylogeny of spiders Family descriptions. 1. Actinopodidae. ssn 2 Apelenidae 3. Amaurobiidas 4 Ammoxenidae 5. Amphinecidae 6. Anapidae 7. Anwodiactdae. 0 8. Anyphaenidae 9, Araneidae. oo 10. Archacidse T1. Atgyronetidae. 12. Agypidae. : 13. Austrochilidae 14, Barychelidae . 15, Caponiidae — 16, Chummidse ~ 17, Githaeronidae 18, Clubiondae... = 19. Corinnidae 20. Crenidae a 21. Ctenizidae 22, Cyatholipdaes nn 23. Cyelocrenidae _ 24, Gyremicheniidae 25. Deinopidae Contents Desidae no Dietyniae Diguetdae Dipluridae 116 Deymsids, us Dysderidae Eresidae Piistacdae Gallienilidae Gaphosidse Gradungulidae Habniidae 132 Hers 4 Hevatheldae 136 Holarchacidae oe Homaloayehidae. 140 Hisetonidae a2 Hypochiidae 44 Iopidae C6 Lamponidae on 48 Lepronetie 190 Linge Liocranidae - Liphisidae. Tycosidae a 138 Malkaridse 160 Mecieoborhidae... 162 Mecysmaucheniidae 166 ‘Micropholeommatidae. 166 ‘Mierostigmatidae 168 Migidae 70 Mimetidae im Miturgidae m7 Mysmenidae non Nemesiidae - Nephilidae _ Nestcdae [Nicodamidae sv sovenennn 184 Ochyroceraidae coo 186 ecebhie anne 188 onopidae. S190 Orsolobidae ST 92 Oxyopidae oars Palpimanidae os Pararchavidac I 19sParatropididae, Periegopidae Philodromidse Pholeidae Phyxlididae Pimotdae Pisaurdae Plecteuridac Prodidomidae Psechridae Saliidae Seytodidae ‘Segestrida, Selenopidse. Senoculidae Sicaidae.-- Sparassidae Stenochildse Stiphididae Symphycognathidae Synaphridae 200 202 204 206 208 210 212 214 224 240 Synotaridae.... Telemidae Tengen ‘Tetrablemmida Tetragnathidae.. Theraphosidze Therididae ‘Theridiosomarida ‘Thomisae Ttanoecidse ‘Trechaleidae. ‘Trochanreridae Uloboridae Zoxhatiidae Zoridae ‘Zoropsidae Bibliography Colour plates Preamble The Royal Muscum for Cental Aiea has long tradition with arachnological studies and was ‘one ofthe frst esearch centres in the world with a specialization in systematics of Afrotropial spiders. The earliest such papers were published almost half centuty ago and articles in the same fei are stil being produced today. In this context, a very large collection of arachnids from ‘Central Aftica was established, la completed with high numbers of specimens from East and West Ata. This provided the senior author of tse present book an ideal opportunity to collborate with ‘Anna Dippenzar-Schoeman, an expert arachnologist from South Africa, The collaboration resulted in the production of manual forthe identification of Afroropical spiders. The vastnest ofthe Aocropical realm indeed renders the creation of manual on its animal groups a daunting task fora single author. But Dippenaar Schooman and Jocqué combined their knowledge on i ferene parts of Aftica and withthe help ofthe complementary collections they were able to com pile the knowledge on the spiders ofthe African continent a chad accumlted until te end of the previous century “This fruitful collaboration based on ther balanced knowledge was a perfect starting point to complete the effort and write an overview ofthe sper families ofthe world. As Africa harbours more than two thirds ofthe word's spider families te task might have appeared reasonably m= ple buran extended effort lasting fr almost ren years was necessary to compete it.The rapidly ‘hanging situation inthe field of systematis, certainly in a group like that of spiders for which the expertise must be considered as the tal-blazing methodology for studies on hyperdiverse arthropod groups, surely complicated the undertaking “This book may stand a8 an example fr intense and constructive callaboration between experts from diffrent continents. Iemay alo be the final ilestone inthe epoch of morplakgieal paler studies tthe onset ofthe era of molecular systematics But perhaps exactly hecause ofthat, the ‘book will remain an interesting document. Although the primary audience for this hook isthe imerested scientist, the publica large, whichis fascinard by the beauty and variation of orga isms, may continue to find the morphological approach easier to appreciate “The Royal Museum for Central Africa gratefully acknowledges the support received for this research from the Belgian Federal Seience Policy Office, and from the Belgian Directorate General for Development Cooperation, which facilitated the collaboration with our African research partners, Guido Gryseels Director General Royal Mascum for Central AfricaForeword Spiders have inhabited Eset for some 400 milion years and are the most profuse and diverse predators on land today. They are familiar animals to everyone. Spiders inhabit almos al teerestil and some aquatic habitats, and many species have developed quite extraordinary methods co ive in extreme environments Understanding the importance of spiders in ecosystems is esendl forthe ‘development of procedures for pes contelin agriculture. Furthermore, because some species which fre venomous man have synanehropic habits, some countries have severe problems wich spiders fas pests. Even where spiders pose no threat, mat people ae aficred with a tangible phobia inter presence. Readers ofthis hook wil, ofcourse, be aware ofthe significance of spiders, be captivated ln thee best an ingen, ad havea sincere deseo Jean more about these fascinating animal “The complete epider fauna is known for nly avery few, small areas ofthe globe, thanks t0 the dedicated efforts over many years of enehusiasicarachnologsts, For most of the world, how ‘ever, out knowledge of che diversity of spiders is imperfect, deficient or negligible. A great deal more information woul hecome available if those who visie and collect spiders in areas where ‘our knowledge fs impoverished were abl to easily identify, even to Family or genus kev the spiders they encounter In Britain, interest in ue spider fauna enjoyed a real boost following the avail abit of the Fist field guide; and the recen plication of Spidersof North American identcation ‘maaal (ick al, 2005) previded an excellent ineoduction to idensifying North American spiders Some other parts of the world have excellent field goes or general books which introduce the reader to the arsncofiuna, bit there is no book which brings together all world spider families Ihetween ts cower. In my oven research on fos spiders, I regularly need o obtain basic infor mation about a range of spider families, many of which L might nover have encountered ‘in the flesh and many arachnologists havea deseo learn about spider families from other partsof the world out of simple curiosity Ts will be obvious co many eaders that this book fllows up onthe authors succesful African pers: An denifcaion Manual (Agricultural Research Counc, South Afia, 1997). The Afican book opened pa whole new world to those unfamiliar withthe spiders of this region. For many, like myself, who sought concise descriptions of a mumber of spider families with clear ilustations and up-to-date references, the African manual was a godsend. Oh to have a similar book covering allehe word families! Well, the authors have aken hee of our entreaties and produced this volume ‘Tae ered-and.tested format ofthe text and illustrations from the Aftican spider book have been followed and extended toa further 40 or so families. Simple keys are provided for those who ike to fall them, and inchde one for webs. Clear line drawings were an indispensable feature ‘ofthe African book, nd continue tobe valuable here. An aasional bonus inthis new book isthe incision of some 150 high-quality colour photographs illustrating represenatves ofeach family “This book will beusefl to professional and amateur arachnologists the word over-Ieis hoped that twill appeal noe only to confirmed arachnophiles but also ro others looking fora general ‘overview ofthe work! spider fauna. Armed with cis volume, willbe raach more sragheforward to track down unidentified spiders throughout the world. As Lit this, news is breaking of many new species across all terrestrial proups being discovered ina previously unexplored highland for ‘stareain New Guinea, One wonders howe many new specie, genera ~even families? ~of spiders ‘exist there? We have a great deal more to learn, Paul A. Selden Past President, International Society of Arachnology Department of Palaeontology, Natural History Museum, London 7Acknowledgements “This book woud probably never have seen the Tight of day without the gencros help of Jon Murph anited nal sages of ts preparation, Tso a he gave ste cpporanity 0 oan {peimens from his emarlaby diverse collection, He allowed the use ofthe ravings made by IM Roberts for is book on Soth East Asa and efeed fee access othe amating colon of phos by his ate ie Frances And ontop of tat he pve comforablesecommodaon co he {Eni ethor one mero vis we pa i. Tune materia sonia forth work nd we te gratfl > Chitin Rolla, Norman Pam Lou Sorkin, Rober Raven, ark Hare, Leon Baer and son Bond who al povided re any colleagues kindly read he text on families that can be considered their Ril of special tuaon,patcalrly Charles Griswold, Reber Raven, Gustavo Horngs, Marin Ramice 2 Maret Runner no provide sme ofthe corrections. nthe same context we aso thank Mark Alors Fernando Aiarer-adll, ose Guidance Bernard ber, Peter ager Christ jen rot Lara opardo, Milan Rersé and Michael Ric Sail Zachoe was very pl ith the prpaaron ofthe key owes yur from the photos by Frances Murphy, an important clltion of pctres was made vay James Cokendelper for which we are ery rate. We ao received pictures rom ‘ast Aldershot Amlaparamil Jason Bond Mil Deer Chates Gris ‘Man Hendk, Castro Hori, Nora Larsen, Astana oi Ley, Demy Mie rer Miler Les Ones, Robert Raven, Hannah Wood and August Verbrogen copyright ARABEL, “ouesy en Van Kee) Samael Zschoke provided some nike spss o complete the web Pecaps te moss oineating work ware ur bythe artis who ad to cope with the odlexcomectos tothe raving Asis Regland Nadine Van Noppen proved emp eo ‘ed with pence’ and chat was ery mach appreciated Some of he rings made by Ea Von Nicer were taken rom our Aiea Spier Book. Jn Boel also provided ws with some of tis fine drawings for which we are most grate ‘We are vary mich indeed Ms Li Heol and Drs No Dien ith for their vale ng suggestions. and Tony Russll- Introduction ‘Over the lst few years the taxonomic study of spiders has witnessed substantial progress. “The fauna parts of che northern hemispheres now well known (eg Heimer & Nenewig, 1991 Paguin & Dupérré, 2003; Roberts, 1998; Ubicke al, 2005; Yaginuma, 1986) and enough know ‘edge has accumulated co allow the production of some local atlases (Harvey et al, 2002), For the restof the world, however, the inventory isin ts infancy. Until ecenly, che literature dealing with the spider faunas of Arca, tropical Asia, Latin America and large part of Oceania was scattered land relatively inaccessible However, with the compilations of Dippenaar Schoeman 8 Jocque (1997) for Africa, Murphy & Muephy (2000) and Deeleman Reinhold (2001) fr eropical Asia, Song etal. (1999) For China, nd keys for Australia by Davies (1986) and Raven & Bache (1997), and Forster & Forster foe New Zealand (1999), the sicuation has changed profoundly. For all of these major regions, almost ll txonomic and mach other published information on spiders has ben summarized in comprehensive books. The majo publications among these provide keys to families, some with descriptions of supra-specific taxa narrowed down to subfamilies (Dippenaar Schocman & Joo, 1997) a genera (Murphy & Muephy (2000), Songet a (1999)), Some provide ‘complete lists of genera with short descriptions of their distributions. The appearance ofthese publications has greatly facilitated the stay of spider in these regions of high biodiversity The availability of a global regularly updated catalogue of spiders onthe internet (Platnick, 2008) has arguably heen even more influential than the recent appearance of regional sper books. “This catalogue provides a complete list of al spiders described to date, together with theit syn- ‘nym, coneise account of thei dstibations and a complete bibliography of the taxonomic itr ste dealing withthe Araneae since Clerc (1757), thus even predating Linnaeus (1758), Ie further ‘ives the number of genera and species per family as well 25 the rota number of species, gencra nd families inthe Aranese. By the time this book is published, almost 40,000 spiders wll have ‘heen described in more than 3,500 gener, distributed over 109 families. Progress has also been nade in uncavelling the systematics of the Araneae. Thorough analy ses have been carried out fr several subgroups such as dhe Gaaphosoidea (PLtnick, 1990), the orb ‘weavers (Scharff & Coddington, 1997) and several families such as the Sakcidae (Maddison, 1985, 1996), the Zodariidae Jocqué, 1991) andthe Cyatholpidae (Griswold, 2001). Coddington K Lev (1991), Coddington eral. (2004) and Coddingron (2005) produced acadogram of all the spider families. However, some taxa, such as the Amauzobioidea, require corroboration, and some pats of the tre remain unresolsed, such as the ‘Dionychae’ ‘A large gap remains, namely the absence ofan illustrated overview of the spider Fails. Although complete list ofthese families is avalabe in Planck's (2005) on-line spider catalogue, no one has compiled a comprehensive account with descriptions and lustrations of represents tives ofthe diffrent families. After having produced the identification manual African Spiders (Dippenaar-Schoeman & Jocgué, 1997) in which 71 of the then 109 families were listed and described, we believed that s was feasible to complete an overview and produce a book dealing ‘wih all anlics, inching che 38 cha had nos been previously covered. “Thas the present book appeats a a sequel wo our manual on African spders."The format is similae and much of the information and many of che illstatons are taken from the previous ‘book. The task was far more complex than we had anticipated. Noc only were new descriptions and illustrations required for the 39 addtional families (though a few are synonymized here), the descriptions ofthe African families requited adjustment to accoune for cheir variability wodldwide Te was clear from the beginning that it would be impossible to produce a dichotomous key with the same detail a that inthe ‘spider mana’ Inthe key in that volume, we included all pessible °‘exceptions tthe ‘normal’ family patterns, sch asthe Zadarids with wo claws, gnaphoids with pseuddosegmcated legs and eyeles prodidomids, to mention bua few. The key we present heres hot as detailed Because we anticipate che appearance soon of family keys on the Internet (smile tothe one for the Australian spider fauna by Raven & Bach), which will hve the advantage of ‘being mach more flexible and allow continuous improvements we decided not to invest too much time in creating 2 dichotomous key that includes all possible excepsions. Our key is therefore a ‘asic one inthe sense thar tony includes those axa that present the basic patern for the Family towhich they belong, It il therefore, be impossible wo find the family to whic, for example, an «eyeessycosid belongs, or the taxonomic rank ofa linyphiid lacking chelceralstrdulation ridges. The main sim of his book is to provide a quick reference tal spider families and an overview ‘ofthe recent literature. Ie might he asked whether now isan opportune time to write a book ot the world's pider fails, asthe axonamy of spiders is bound 0 change rapidly once the ATOL (Araneae Tree OF Life) project has been implemented (see Hormiga et al. 2004). Tis lnge-seale project sims to complete a phylogenetic analysis ofall spider Families by the creation ofa character {ltabase combining morphologial and molecular characters, rsuling ina matrix with approx rately ewenty millon ells. Because the molecular characters wll outnumber the morpbologial ‘mes, it islikely char the position of many taxa wil change inthe near Future. As s already vious in many othe taxonomic groups (eg. Turbellaria; Willems eta, 2006), spider systematies will bbe increasingly based on molecular, ie. ‘invisibe, characters. ris tobe expected that spiders with similar morphology that are now considered toe closely rated coukl be assigned to taxa from diferent evolutionary lineages owing o new molecular evidence. However, there are disciplines, ‘uch as paleontology, in which molecular data is of litle use. Our book may well be the last ‘verve based entialy on morphological characters, butt could tll eof use in disciplines that rely entirely on morphology long after the hasies of spider systematics have been altered by new molecular insights. Tha certain sense this book was conceived with the following question in mind: what infor marlon wold palaconssloyn eyuire to tdentiy« oni spider based os naracter ta rely tarbe preserved? Thisunderlies our lack ofemphsison details that are wnlikely robe observed in 4 fossilized specimen. The degree of detail ofthe spinneret spigot armature is an excellent exam- ple. For many spider families thee are now detailed deseripions ofthe spigots om each spinneret (Penick, 1990; Plaick ct al, 1991), bur only a few ofthese are relevant wo the positioning of families in che phylogeny ofthe Araneae, Only those spigots that have to do with the placement ‘of taxa are mentioned in this book “The example above illustrates the modest scope ofthis book The most complete overview of spider morphology and sjstematics published to date, the monumental monographs of Simon (1892-1003), dates back to 1897. His work summarized almost everything known at chat time bout the systematics and morphology of spiders. Today, similar undertaking would require she cooperation ofa tam of specialists, a task that could be atempted in the coming years. In the inet, however, this hook provides a concise overview of the efferent spider fails, summa tizes the relevan information that soften scateredacros= many papers inthe iterature and sheds Tight on the position of tata within the phylogenetic system that isin use today Contents of book “This book contains the following: + Mastrated hey to the families thee diagnostic and descriptive characters, notes on thelr taxonomic position, and ther lifestyles and distribution. ‘+ Apart from the key to fails, a key to spider webs is provided. This is the ies time that \wehave wied to inci his ina book, and while doing so we realized the enormous varity ‘of spider webs. The key is far from complete bu this irs attempt at providing informa- tion about webs may form the hass for creating more complete keys i future. + A series of phocographs of living spiders, mainly from the collections of James Coken- dllpher and the late Frances Murph, i also provided. + Inthe appendices a general phylogeny of spiders is presented nostic characters foreach clade thalliseoF the main diag- As mentioned inthe Introduction, the keys will only ead to the identification to family level of those specimens that match the diagnostic characters. Certain pooely defined families suchas the Stiphilidae and Desida have been exchded fom the keys Family descriptions ‘The family name in Latin followed by the name of te author and bya veenacular name forthe family: In some cases several names are provided because the family is referred to indifferent ways In some instances we had to coin vernacular names, and ie was not always possible to find an appropriate vernacular name for those Fails that are poorly dened and contain a great varety of forms. ‘A the beginning ofeach description the type genus i given along with information about the sizeof the family (numberof genera and species). Ths was often based on information inthe n-ne spider catalogue (Patnick, 2005) The complete list of generais nly provided if thet num her does not exceed ten, ‘This fllowea by alist of diagnostie characters. It summarizes those characters that define the family Strangely enough, ic became apparent that several families lack elear diagnostic fea- tures. Some are based on one ora series of pesiomorphie characters (e@ Gallienelidae), which is workable, bur others are in fat defined through a process of elimination This is the case with large families thar were pli in several pars sacl as the Clubionida. Mo fries hat were spit ‘off (eg. Selenopidae, Sparasidae) are how well defined, alchough among these some da not sm tw form monophyletic ents (¢ Liocranidae, Stiphididae), But the main problem is with what ins ofthe original taxon, in tis case the Clubiondae, for which no sfnapomorphies have ‘been ecognized. Some families ae temporarily retained simply for historical reasons (Desidae), Following the list of diagnostic characters, an overview i given of important external car acters. Considering the variability within some Families, i was often impossible ro describe the entire range of variation in certain characters, and we consequently attempted t is he different character states accoeding to subfamilies In some case an approach athe gens level would have been necessary toillstrate he envire range of variation within a Family, but we considered this co bbe buyond the scope of our sid Under the heading taxonomic status, she present status ofthe family is summarized and, where relevant, ashore historical aecoune of ts systematic position is given. For the smaller fam: ilies, te avallbiliy of evsions of particule genera is mentioned “The distribution of family may be important because t wil indicate whether it has been found ina particular region. However, one ofthe main purposes ofthe book isto enable a person to deny taxa that may be encountered outside thet known geographic range Under the heading lifestyle, a hei account is provide of what kindof predator the spider {sand in what habitats it may be found. Each family account ends with list of relevant Heeracure, havi, the books and articles on which the information in the text is based. Many ofthese sources requted meticulous analysis oF lists of characters, matrices and cladograms, often in crypte form, to retrieve useful infoemstion.‘The significance of and fascination with spiders ‘Spiders form one ofthe largest groups of invertebrate animals, nearing 40,000 known species (Platnick, 2005). They are disteibued worklwide, oecur on all continents except Antarctica, and fare found in every conceivable terrestrial habitat, including caves, snow-covered tundea, high mountains and intertidal zones. One species, Argyroneta aguaica, has even adopted an aquatic Iesyle, All spiders ae carnivorous, and in principle al are hunters, although many of them use webs to catch their prey (Shear, 1986; Dippensar-Schoeman & Jocqué, 1997). Despite the simi larity in their biology, deals ofthe festylesof spiders vary greatly. Their enormous rangein size provides hin ofthis variacon, For example, the hody length of spiders ranges from a large 120mm {Therapose blond 1 somes ess than 0:4 mm (Pata dig), and there fas mac variation in ‘heir biology asin their size or each family, we provide 2 short summary of the ifestyl, mentioning only the most com ‘mon types of behaviour Far more detailed account have been published elsewhere; for example ‘Shear (1986) provides comprehensive information on webs and how they are used, The classic ‘work by Bristowe (1958), snd the Fist spider peblieation to contain spectacular colour photo graphs by Kullman & Stera (1981), cootain more information on this topic. The volumes edited by Witt & Rovner (1982), Barth (1985) and Nentwig (1987), and the works by Foclx (1992) and ‘Bauch (2001) focus on many dilferene aspects of spider biology. Jackson & Pollard (1996) focus ‘on behaviour These works ar ‘compulsory reading for all those who wish tp faniliasizethem- ‘selves with the lives of spiders ‘More general information on the lifestyles of spiders from various continents s availabe, Dippenaar-Schoeman & Jocque (1997) deal with African spiders, Forster & Forster (1999) ‘with spiders fiom New Zesland, Murphy 8 Murphy (2000) with South-East Asis, and Ubick ‘ral. 2005) with North America, These books are general works, There is also a wealth of Tocal books on spiders from many countries, which usually contain chapeers on biology and behaviout ‘Spiders are excellent models fo the study of sexual selection. Their exposed male copulatory congans ange in complexity froma simple intromittent'thorn'to along whip-like structure accom panied by amazingly complex supporting sppendages. The variety and complexity ofthese organs fnd the accompanying variety in Secondary sexual characters and courting behaviour, all part of ‘what i called the ‘mating module’ Jocque & Seats, 2001), provide suitable and often preferred models forthe study of sexwal selection (Huber, 2005c). Te wealth of iterate on spiders, mentioned above, leads ws tothe conclusion thar spiders belong to larg taxon with avast array of adaptations ranging from geheralist to those narrowly adaped to particular ecological circumstances. ‘Spiders have other‘ peoperties' that make them idea subjects forthe study of biodiversity in gener and the evaluation of natural habitats in particular Tn contrast to other mega-iverse terrestrial groups with large numbers of species such as the Acari, Nematoda and Collembola, spiders ar relatively large animals. Because their external ‘copulatory ongans are species specifi, i fey simple to identify chem. No time-consuming dissections ae requited for thir ientfcaion, and sorting spiders by morphospecies is thus fairly ‘staightforward, Surveys of spiders are, therefore, he most simple to conduct, and spiders appear to be the ideal animals co use for rapid biodiversity assesment. Simple identification, ease of ‘collection and fineuned isribuions make these animals study objects par exellence fr decison- tnakets who require information about the intinsic biological value of any particular habitat. Spiders provi such information aboue the vale of any pardcula abicar beter than higher plants 2 ‘or vertebrates (Mittermeier el, 1999), and thus offer small-scale data for the selection of bio- Aivesiy “hor spots tha is required for eige purposes “And theres more! Jacq (1981) and Volrath (1999) showed tha size in spiersis dependent ‘on the quality of the habitat: Monitoring thir size can, therefore, be considered an cay system revealing changes in habitat quali svarning- Tn assessing the quality of rainforest patches, the preferred indicator axon is Ctenidae. The density of occurrence ofthese spiders can be rial ‘estimated by night collecting using the light froma headlamp chat reflects off the taper in th back oftheir eyes (Jcqué eta, 2005). ‘Spiders are important in another way. They are vital component of mos erestrial ecos}s- ‘ems, not leas of agricultural systems. They strongly affect the density of insect populations and have heen shown to limit insect pests inthe agricultural environmene. Wise (1993) summarized the research on tis subject.Morphology and terminology “The body of spider is divide into two major regions: the eephalochorax (prosoma) and abdomen. (episthosoma, connected by anarrew pedicel (igs 12, b). The following morphological details se considered inthe family discussion: cephalothorax: carapace, sternum, eyes, celicerae, ‘mouthparts appendages: egs and alps: abdomen: dorsum and veter of abdomen, spinnerets and genitalia. ‘Carapace (ig. 1a): cam be divided into the cephalic and thoracic regions. In some species the Alivnton is clearly demarcated by the cervical groove. Behind the cervical groove a depression, [known as te fovea, i present in mos families Ir serves as an attachment for the dorsal muscles ofthe sucking stomach and the muscles to each cliera. The shape ofthe fovea canbe logit ‘inal or transerse. When transverse ican be straight, pro- or recurved. The shape and station fof the carapace are important taxonomic tats, The venom gland is loeated inthe cephalothoras “nd consist of along, cylndrieal portion and an adjoining dct that terminates ar the tip ofthe ‘eliceral fang. ‘Steemum (figs 1b, the undivided sternal plat (sternum) lis on the ventral side ofthe cara- pace. Anterenly the stermim i marked by a distinet groove, the lablosternal junction, but it can be fase in some families (eg. Filsatidae). Inthe Mygalomorphae, the sternum bears sgl, smal licular impressions on the sternum that vary in pesition, shape and number between families Precoslscletite are sometimes present between the sternum and coxae and interconal selertes ‘between the coxae Eyes (igs 2h, c: most spiders have eight simple eyes, while some have sis, fou, eo or none. "The ees are arranged in fons or groups. The most common arrangement isin two rows that can be stright, procurved or recurved. The eyes are named according to their postion on the cara paceanterior median eyes (AME), anterior lateral eyes (ALE), posterior median eyes (PME) and posterior lateral eyes (PLE), The median acular quadrangle (MOQ) isthe area of te Four median jen The eyes are sometimes situated on an eye tubercle, carina, kels oa protuberance. A tape tm is sometimes present inthe secondary eyes (AME, PLE, PME), which then appear pale in colour and purportedly allow nocturnal vision. [Cheticerae (figs 2s, ,d) cach chelicera consist of a stout basil ection (paruron) anda smaller, movable distal section, the fang. The chelicera are either Free or fused ( g, Symphytognathida). "The fang usually rests ina groove, the cheliceral furrow. One ofboth sides ofthe furrow are often armed with teeth (promargnal and retomarginal eth). Spiders with such teeth mastcate thelr prey, while spiders without teeth suck the fluid out oftheir prey. The cheicerae in some groups fre chelate closing down on a toothike process. Others beat laminae or keels (eg. Gaaphost ae), peg teeth (e- Archacidae) or stridlatng files (Linyphiidae). The fangs can be shore and stout (Zoxaridae), very long (Desidac) or provided with kel (Migidae). The movement of the che Ticerae is either paranal or diaxal. The anterioe portion of the paruron is usually provided wich strong spines in moet of the burrowing mygalomorph spiders. These spines, collectively known as a vasellry, are used to excavate burrows “Mouthparts figs 1b, c, 23) the basal segment (cox) ofthe pap is enlarged to form the chewing rmouthperts, the endites(gnathocoxae). Inthe Mygalomarphae (fg 1c) the endtes are broadened ig. Exteel morphaog- 4, Amon dn ves Anema er i Myge ‘hte, eal em 1s~ “ Sees | S005 a nl atm ~. je conjte coum — Fig 2, External morphology a. mouthparts and pl, etal ews bchliere and ge fontal ew 98 pricem dona sews celts, rl ew showing arm pine, Jel view ALE serie itl ‘les AME” aco tan ee: PLE ponte era yes PME: pero malian ees 16 laerally. In most spiders the rm ofthe ene hears a.caicula, serrated ridge known asthe serrul This is used ina sale fashion to cut pre. The promargas ofthe endites are feinged with seop ‘la, 2 dense cover of seta used to filter the liquefied fod. In between the edie isthe Ibium, ‘whic is fee from the sternum but sometimes fused ri (eg Filistaidae). In afew families the lista end ofthe labiam i thickened and strengthened, i. rebordered (cg Linyphiidae). Inthe Mygalomorphue the endizes an labium frequently beat cuspules (ig. 1). ‘Legs (ig, 12): four pars, each consisting of seven leg segments, the legs are usually conered with seta, spines, various sensory setae and receptors, Distinct sensory setae ate the fin, fae ike seize sct vertically in conspicuous socket, the trichobothria. Some groups have dense, shor stiff setae, the scopula, positioned ventrally on one or more ears or even tnettarsi. All spiders hhave at least two claws om each earns. In most of the web-sing groups 3 smaller third claw is present (fig, 394). The eworclawed spiders usually have a dense brush of setae, (Gig. 75g) below each claw. In some species the claws are situated on an extended part ofthe ta sus, the onychium. In some three-lawed spiders, serrated bristles known as accessory claw are present. Inthe Theriiidae and Nesticidae, tarsus [V bears a comb of serrated bristles (fig. 104). In cribellate spiders a calamistrum, consisting of « comb of setae is present on metatarsi IV (1108) he claw tuft Papi (igs 22, 32-: these are leglike appendages consisting of sx segments (compared to the seven leg segments). The papal mecatarsts slicing, In females che pap is simple and usvally bears single carsal caw In adult males, some segments ofthe palpate modified into a second: ary copalstory ongan and can either be simple in structare (haplogyne pap, fg. 3c) or complex (aelegyne pap, figs 3b) The tibia and sometimes the patella and femur ae provided with one ‘ormore apophhyses (dorsal ental or retrolareral), In the entclegyne typeof pap the tarsus ust ally provided wih a bow-shaped cavity, the eymbium, and a genital bub. In some families the {ymbium bears a basal appendage, the paracymbium (eg Linyphidac). The genital bulb consists ‘oF more a less sclerorized tegulum, with the winding spesm dts frequently discernable, ad an inromitaneargan, che embolus. The embolus usually estson alate or membranous appendage, the conductor. The elu x sometimes previded with a sclerotized appendage witha memlra- ‘os insertion, the median pophyss. Abdomen (figs 12+) the ablomen is joined tothe cephalouhorax bya thin pedicel trough which the circulation and feeding eystems are eanalized The exoskcleton of che abdomen fe much thin rer than shat of the cephafothorax and this allows great expansion ofthe abdomen when prey boeing fed upon, or when a large numberof eggs are being formed inthe female. The abdomen is quite variable in size and configration, In many spiders iis eliptca, oval or globose and ether Soto covered with scleries knoxin as seuta. Many species, however, have distinct abdominal shapes, decorated with projections and protuberances, The hearts frequendy visible as long tudinal mark through the integument. The dorsum can also be decorated with patterns consisting of for example spots, bands, chevrons oa fliam. Spinnerets (fig. 24): most spiders have three pairs of spinners, the anterior, median, and pos- terior pairs, situated in font of che anal opening The spinneretsate very mobile and well provided with muscles. The sinning glands, consisting of ampullate aciniform, cuuliform, aggregate, p= iform or fagelform glands, terminate as small spigots onthe sueface of each spinneret. The posi tion, dhickness and number of spinneret segments and the number and shape ofthe spigots arc ‘characters used at generic level. the cribellate group a sieve-ike plat, the eibellar, i presen “The special type of silk emisted by this organ is combed out by the calamistrum on meeatts V ”a » men ocptyse esa eponre Fig 3. External and internal morphology apap (ete) tral ve: bs male pap (mene) ‘bunt venser male pal Cuplogyne bert ew ema gerald ee eae ena cig tran dsl iw {many famities lacking acibellay, a small conical appendage, the coll, present a the ase ofthe anterior spinnercts Respiratory system (igs Ib): most spiders possess two kinds of respiratory systems: one or to pairs (Mygalomorphae) of baoklungs and one or two pars of tubular tracheae. The external ‘of the hookkang ze present on ether side ofthe epigastric re tubular rachese ‘open through the tracheal spsace. The eacheae can branch Uhroughout the body and vary i size and pattern of distribution. Inthe Mygalomorphae, four booklungs ae present. ‘Genitalia (figs 3, c: close to the pedicel, the epigastric groove runs transversely across the abdomen, The external openings ofthe genizala are located inthe middle ofthis furrow. I the Females of the entclegyne group a sclerotized, special copulatory organ, the epigyne, i present in frone of the genital opening. Is absear inthe haplogyne group and Mygalomorphae.Glossary singola; p= pla Abdomen: aso called opsthosoma: posterior par ofthe body of spider. Accessory claws: mf setae athe tip ofa only presenti web bulding spiders sed rogether tvth areal claws to secure grip om threads. Alveolation: a small but Jeep rounded depression, ‘Anal tubercle: small uberlesbovespinetets through which the snus opens ‘Anterior lateral eyes (ALE): er situated at each end oF anterior ee row [Anterior lobe: anterior part of ene, often elongated in Mygalomorphae [Anterior median eyes (AME): mile pair of ees in anterior ey row. ‘Apex: disap. “Apomorplhy derived character ‘Apophysis (p= apoplhyses): an excrescence or appendage changing the general cylindrical or global ‘dpe of asclerice ost often sed for description of male pal. Arancophagous: feeding on spider. ‘Atrium (p= atria internal chamber ateneanceof copulation tract ema haplegyne spiders. Autospasy: casting of limb. Many spiders have a weak spot or aerack in a parscuar place on the igs ha acilaesauospasy [Bidentace: provided with wo roth, Biserially dentate: provided with two rows of teeth Booklings: respiratory organs situated infront of epigastric furrow on ental side of bomen, open ing through natrow sis. ‘Branchial operculum: (p = oprcul) 3 pair of plates (two pars im Mygalomorplae), often pale or ‘orange in colour on ventral side of abdomen, jst in font of pigasvic fla slit posterior end opens {neo booklungs ‘Bulbs (p= bull) or bab: comple part of male pap iserted on usualy hollow veneralsde of ey ba Capue (p CCalamistram (p= clams}: combslke row’ or rows, o oval are, of modfed setae oa mea Sus of eel per, wed to comb sk produced by the crbll pita: another name for cephalic region ofcephalothoras CCarapace: dots slerit covering cphalothora, she anterior pat of body. Cervical groove: shallow U-shaped groove, sparaing cephalic region from thoracic region of cara mre, (Cephalothorax: sso called the prosoma, anterior part of body covered by the carapace, beating 6% lcgrand mouthparts composed of two regions: cephalic region in front separated from thoracic region Inhind cervical groove Cephalic region: frontal par of esphalothorax dclimied by cervial grooves (Chelate: sid of calicerae in which the fang doses doven on 3 tooth ke proces. 2 Cheticera ‘Cheliceral furrow: shallow groove on basal portion of chaicera accommodating the fang, usually provided with reth on ts pomarpin and retomargin, eicerse):insered at front of carapace, consisting of large basal porion anda Fang CCheliceral lamin (p= laminae): chicnous ridge extending distally on prolaterl edge ofeach che- Teens. CChelicera ech: thor-like extensions af heicral furrow promargin (ter oF dorsal margin) and _eromargi (inner oF vente margin) (Chita (p= cia small sericea base of cisicer, ust under Aypews (Claws se tarsal cis, (Claw tuft: a dense group of etc under the paired tarsal laws usualy well veloped in hunting spiders CClypens: part ofthe carapace hetwcon yes and anterior main CCollariform with one curved ie CColutus:short median Conductor: pur of bulbus, accompanying and supporting the embolus (Condy: laeral boss a base of chliceae. CCopulatory opening: double opening in epigye through which the embolus inserted (in enelgynes) fr sngle slit ftough which make pal ongane are inomied in Baplogymes). rotuberance in ron of spinners, considered 2 modification ofthe cibellum, Coa: (p = cose) see legs and pap. (Crenulate: with numberof longitudinal sides, CCribetlate provided witha cibellm, Cribetium: sieve-ike spinning plate in fron of pinseets CCeyptozoie: ving a concealed ie CCaspules: smal spiny warts on endtes and labium of Mygalomoepbae (Cusps: shore, mosdy blunt spines Cymbium: (p= ymbia) dorsal par of mae papa tarsus, most often holo, carrying the usualy com ple bulbs Dentate: toothed. entitle: small oth. Desmitracheate median tracheal trunks with many Branches passing into he prosoma (se aplo sracheate. iad: group of wo, a psi, Diaxiak sid of chelicerae extending downside with fangs dosing towards midline Dionychous possessing tw claws on tarsus, Dorsum: dors side of sdomen, ‘Bcribellate without a eibellum and calansrum dentate: without teth, Embolus:inromitted par of male pspal bulbus, wsualy slender, sharp-ipped and strongly sleo- tized, carrying terminal part of sperm duc. Endlice: basal segment of lp lso called the maxis or goathocon. a Nee ee)Endive heck: innermost extremity of end adjacent to anterior tp of abi, Endosternit:interal slit, a remnant of abdominal segmentation Entelegyme: refers to spiders with n epgyme, wih separate dct for sperm transport daring nse ‘naton towards spermathecas) and ferlization(fowards were): see haplogye. Entrance ducts ducts in eatlegyneepigynes leading fom copulatory opening to spermathecae Eplandrous: structure on the abdominal veatr in the male, inthe Viiy ofthe epgasre furrow EEplgastrie furrow: a tansvers slit on anterior pat of ventral side of abdomen, The anterior pa of bookings opens a edge of furrow as do the onopores. Epigyne (p= epigyas)a chiinous plate on ventral side f female abdomen in which the genital open ings are loeaed only fly develope nad Females of ensegyne spider; spiders belonging “Mygalomorpae and ‘haplogyne” Arancacmorpie do not have an epgyne. Eye formula: poston of eyes i often expressed by digits separated by colons (eg: 22-22 = ees in four ems comprising eo cys each; G2 ~ een in wo roms, fa ow With i, second with C0 eye). Fang, distal part of thechlicerae. Femur (p femora}:sce eg a palpi Fertilization ducts duct inentelegyne cpigynes leading from spermathecse to uses. Fissidemtate:tvth having more than oe point olism (p= fli) lea'shaped pattern on dorsum of abdomen. Fovea (p= foreae}acemval depression on cepslothoras, often reduced wo alngicadinal drksripe, corresponding with an internal ge o which muscles ave atached Pased chelicerae:chelicerae fused togeher,somesimes only at base or along entire length Genial opening: in female che opening of uterine dct i epigatrc Farrow batween books males the opening of duct fom estes infarc between booklangs Gnathocoxa (p= gnathocoxse: basal segment of pap also called the mails or ent. GGumfoot web: space web with stands ef sik provided with sticky droplets near distal tachment point Hiaplotracheate: median trunks of tracheae unbranched, confined to the abdomen (see desmirs- chest Haematodocha (p= huematodochae); membranous, inlable par of bulbus in male pap ‘Haplogyne: refers to spiders lacking an epigyne and thus having only one pa f ducts for eanspore ‘oF sperm fom ste to spermathecae ding insemination and back o eer fr feriztiony see ‘emtlegyne. Hinged hn: long dark sea with hinged base giving it unlike ordinary setae, flexible dvection, Hirsute: hry Intercoxa slerites small slits between conaeof legs, sometimes connecting sternum and ca pace Labiosternal junction: junction betes Ihium and sterem, sometimes sbscne when chee slo rites ar fused. Labium (p= labia: slerite stated between enites infront of sternum, [Labrum (p= labra}: ape ip: mouthpsrt concealed hy cheer Labral spar: shor: shri eteasioa of labrum, 2 {Lamina (p laminae} lat, mos often translucent, slerotizeexcrescence, om cera in some spiders. comarginofche Laerigrade: refers to spiders that move like 2 crab, with legs directed wo the side (opp. prograde) Legs: consist of seven segment: coxa, trochanter, femur, acl, ii, metatrsus and tarsus Legare numbered frm the front 1 Leg formula: reaive length of legs epresented by Four numbersin sequence of longest shortest eg 4123 fourth legis longest nd hid eg shortest. Male palp: refers vo the mouied tars ofthe pal in male spiders copulation organ, notin diese ‘onnestion with exes. Consists ofan enlarged and hollow tarsus (yb), smeines plc nt (cymbisns and paracymbium), bearing the sexual organs. These vary greatly in shape and complesity {nd ae the mose important characters fr species identification i ale spider. “Mating spur: strong spur usally presen on eg of males, used daring mating Manilla (p= manila) se ences ‘Median apophysis:.n apophysis of male papal bulls, usualy with membranous insertion Metatarsus (p = meats) segs. “Monophyleti: refs to a systematic grouping which is consiered tobe natural contrast ply pyle ‘Median ocular quadrangle (MOQ): quadrangle limited by four median ees “Median seprum: longitudinal serve on Noor of epigynal atrium. Notched: describes trochamtes with a ventral indentation on distal magn, "Nubbin: nonfunctional spigot, usually nob shaped often a vestige ofa igo hat was functional carlierin evoluion.Ampullatespigots are moce numerous in ancestral Araneomorphae andendt0 Be ‘eplaced by mubbina batore they comply dieppese ‘Onychium (p= onychia): ventral extension of tp of asus bearing the das Opisthosoma: sce abdomen (Ostia (= ostium) slight dilatations with openings ofthe main hemolymph vessel in the opsthosors Palea (p » plea distal 2 east parilly membranous part of tegulum, wel separated from rest of serie Palp (p= palp): als called peipalp; second appendage of ephalothor in front oe composed of ene rochanter, fem, patel, ia an tarwis. This appendage is modified into a capulstory ‘ongun in mae spiders, aracymbium (p = poracymbia) appendage of eymbium on male pap, may bea separate seer araxial: suid of cheicerse extending forwards, wih fngs closing toward abdomen. Patella (p= pte) se legs a pl aturon: basil segment ofthe chelicerae. Pedice: marrow connection between cepalothorax and abdomen. Pedipalp: soe pap eget cher ethin sek coming of sb, stot spins on pomarginor roman of Petiolus:sce pie Plesiomorphy: ancestral character a ee‘Pleural bars: nacrow, horizontal slertes between conae and carapace, sometimes connecting inter. coma certs Plagiognathy:fngs directed obliquely ( apposed ro orhognathy or lsbidognathy) Pluridentate: having move than on toch Polyphyletic: ers to an artical grouping of axa (in contas to monophyei, which designates 2 aturl groping) Porrect describes celicerae directed forwards, Posterior lateral eyes (PLE): ests at each end of posterior 1% Posterior median eyes (PME) so incermedist yes in poserier row reconal sclerites: small scrife extensions berween stem and eg cote. Preening brash: dens clister of setae near ventral ip of poster mecatars reening combs 3 tramserse row of tifsetz at vena tp of posterior metaaes Procarsus: 4 rrm restricted to Pholeidae where i used t indicate the strongly developed, often ‘lindo paraymbium, Procurved: describe structure a 20s in which outer edges at in fron ofthe central part Prograde: dscbes spiders wih og dected forward (LI) and backwards (1 and IV) (opp et grad) Promargin: dorsi margia of celica furrow Peosoma: anerior part of spider including cephalothorss and ts appendages. Radial anascomosis: rai of we join before reaching the ub, seal (p = rast): aie structure arexremityofchalicere in Mygatomorphae ofan rece ta few song spines used fr burrowing, [Rebordered: with thickened edge; describes assert in which the margins thicker than the main pare Recepeaculum (p= ceceptacul) se spermathecie Recurved:desrins struct or ow in which over edges are behind che ental part -Reeromargin: venta or posterior margin of chelicea furrow ‘Sclrite: a single slevotzed part of external, hardened tegument SScapus (p=scpi):an elongate marrow lingsiform (eongue-shaped) appendage of he epigyne. Scopala (p= scope: 2 brush of setae om either promargin of eicera, distal end of ents or the ‘ental side of terminal leg segmenss: improves grip on substrate or pre. Scutum (p =scuta: sclerotized place on abdomen of some spiders. Secondary eyes: three cj pairs anterior lateral eyes, posterior malian eyes and posterior lateral eyes ‘used primarily movement detectors. ‘Semichelate:chticera in which dhe fang has restricted mobil Seerala (p= srrulic): 2 ow or chaser oftiny teeth along anterior margin of end often ony visible asa dark line Serrated bristle: typeof sta that slightly curved, bearing serrations aloag one se Seta p= setae): baile, capered and feb rcs on legs and body ef spine andtrchobesbriam). ‘Sperm duct: duct in reglum of mal plp for storing sperm, connected wo embols. ‘Spermatheea (p= spormatheca)-biadr-ikestrucares with gland ining in vulva of females for bring sper afer nseminston, Sigila (= sila): circular impressions on sternum of some Mgslmoephae andl dorsum in some “Arancomorphse, crtespending With mera muscolar tachment Spigot: tiny cusps or elindvcal excrescences a apex of spinneret from which silk merges Spine: pone, rigid srvcureon body and legs, usually aticulating Spinnerets: appendage of abdomen, arranged in several pairs but reduced to three in most spiders amterior lateral, posterior median and poster leral (here ofcn called resp. antrio, melt and posterior spinners). Provided with small pigs from which the silk emerges, The anterior median Spinmeres are icher replaced by cebu, acollus or ae totaly absent 5 les shore spines, almost as thick ae they are long. Spiracle: ee trachea spire, Spur: cuicular appendage hevier than a spine. ‘Squamate: wich small scales in connection wih tegument) Seabilimentum (p= stabilimenta}:band of dense silk ina web, Seermum: large selerce on ventral sde of ephaloshora situated between leg coxae and behind the habia, Seria (p= striae): paired depressions; usually three prs of darkened stripes adtng rom fore, Stridulating organ: a series of thin ges on sevice, forming a sedating le corresponds with 2 Series of shor si setae another part ofthe body that can be scraped slong the fie to make a Seridustory sound ‘Sustentacuum: thick macroseta with ben tp stuaced behind the accessory claws. om tars IV ‘Synapomorphy: shared derived characters “Tapetum (p= apo) ight reflecting ser n secondary cyes(ALE, PME and PLE); yes app pale sn colour assumed o be used for noctumnal isin, “Tarsal claws: situated at tip of asus ether a single pir, often concealed in aca efter pair and a thi single cla, whichis much smaller ad situated under the paired sualypecinate claws (= 0 ‘ded witha row of san-like tet). “Tarsal organ: a sensory receptor, most ofen a tiny depression on dorsal side of asus in some fais well developed projecting above surrounding tegument and clearly discernible, “Tarsus (p= tarsi: see legs, “Tartipores: races ofthe insertion of pits presenting the poson ofthe spigot ding the previous ‘Tegulum: par of the bulbs housing sperm duct which ends in embolus. “Tegument: external cial skin, "Tenent sotae: tubular sete with expand tps present om ars toast in gripping. “Terminal apophysis: most apclslerite that inserts nto embolus vs distal haematodachs ‘Thoracic region: posterior pat of eepalothoran “Tibia (p= eines legs and paps. “Tibial apophysis an apophysis on iia of male pap in some spiders Me