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iS HARVARD UNIVERSITY -^ Library of the Museum of Comparative Zoology (US ISSN 0027-4100) The Systematics of Neotropical Orb-weaving Spiders in the Genus Metepeira (Araneae: Araneidae) WILLIAM H. PIEL MCZ LIBRARY JUL 3 2001 ( HARVARD UNIVERSITY HARVARD UNIVERSITY VOLUME NUMBER 157, 1 CAMBRIDGE, MASSACHUSETTS, U.S.A. 8 JUNE 2001 . (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952- bulletin 1863- Memoirs 1865-1938 JoHNSONiA, Department of Mollusks, 1941-1974 Occasional Papers on Mollusks, 1945- SPEGIAL PUBLICATIONS. 1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere- dinidea (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, 1973. Morphology and Evolution of Blastozoan Echino- J., derms. 284 pp. 4. Eaton, R. 1974. A Flora of Concord from Thoreaus Time to the J., Present Day. 236 pp. 5. Rhodin, A. G. and K. Miyata (eds.), 1983. Advances in Herpetology J., and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 725 pp. 6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes ofthe Gulfof Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. {Bulletin ofthe M. C. Z, Vol. 108.) Reprinted 1971. Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings ofthe First In- ternational Symposium on Natural Mammalian Hibernation. {Bulletin ofthe M. C. Z, Vol 124.) Orinthological Gazetteers ofthe Neotropics (1975-). Peter's Check-list of Birds ofthe World, vols. 1-16. Proceedings ofthe New England Zoological Club 1899-1947. (Complete sets only.) Proceedings ofthe Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum ofComparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. Thispublicationhasbeenprintedonacid-freepermanentpaperstock. ©ThePresidentandFellowsofHarvardCollege2001 THE SYSTEMATICS OF NEOTROPICAL ORB-WEAVING SPIDERS IN THE GENUS METEPEIRA (ARANEAE: ARANEIDAE) WILLIAM H. PIEU CONTENTS Abstract _ ___. 1 21. Metepeira gosoga Chamberlin and Ivie Introduction 2 59 Acknowledgments 2 22. Metepeira ohnec New Species 60 Materials and Methods 3 23. Metepeira comanche Levi 62 Collections Examined 3 24. Metepeira pimungan New Species 62 Locality Data Storage and Manipulation 4 25. Metepeira triangidaris (Franganillo) .. 63 Examination and Illustration 4 26. Metepeira arizonica Chamberlin and Metepeira F. O. P.-Cambridge 5 Ivie 66 Key to Female Metepeira 12 27. Metepeira atascadero New Species .... 67 Key to Male Metepeira 17 28. Metepeira incrassata F. O. P.- Metepeirafoxi Group 19 Cambridge 68 1. Metepeira datona ChamberHn and Ivie 20 Metepeira ventura Group 71 2. Metepeira desenderi Baert 21 29. Metepeira ventura Chamberlin and 3. Metepeira grandiosa grandiosa Ivie 71 Chamberlin and Ivie 23 30. Metepeira revillagigedo New Species 73 4. Metepeira grandiosa alpina 31. Metepeira celestun New Species 74 Chamberlin and Ivie 24 32. Metepeirauncata F. O. P.-Cambridge ... 76 Metepeira vigilax Group 26 33. Metepeira crassipes Chamberlin and 5. Metepeira cajahamba New Species .... 26 Ivie 77 6. Metepeira glornerabilis (Keyserling) ... 28 34. Metepeira chilapae Chamberlin and 7. Metepeira vigilax (Keyserling) 30 Ivie 78 8. Metepeira rectangula (Nicolet) 32 Metepeira ininima Group 80 Metepeira labyrinthea Group 33 35. Metepeira petatlan New Species 80 9. Metepeiraspinipes F. O. P.-Cambridge .. 34 36. Metepeira minima Gertsch 82 10. Metepeira lacandon New Species 37 37. Metepeirapacifica New Species 84 Metepeira nigriventris Group 38 38. Metepeirajamaicensis Archer 86 1112.. MMeetteeppeeiirraa tniagrraipvaecnatriNse(wTaScpzeacnioewssld) 4308 LIintdeerxature Cited 8918 13. Metepeira calamuchita New Species .. 42 14. Metepeira galatheae (Thorell) 43 Abstract. Ofthe 39 species and three subspecies 15. Metepeira karkii (Tullgren) 46 Metepeira conipsa Group 47 ofthe orb-weaver genus Metepeira in the Americas, 16. Metepeira compsa (Chamberlin) 48 36 species and two subspecies are known to occur 17. Metepeira roraima New Species 53 outside of the U.S. and Canada. Yet, despite their 18. Metepeira gressa (Keyserling) 54 conspicuous webs, diurnal foraging, and relatively Metepeira incrassata Group 56 common presence, the taxonomy of Metepeira is 19. Metepeira maija New Species 56 poorlyunderstood, probablybecausethegenitaliaare 20. Metepeira inca New Species 58 smallanddifficultto distinguish. Infact,manynames for species south ofthe U.S. were, at some time, in- correctly synonymized with the name Metepeira la- ' Museum of Comparative Zoology, Harvard Uni- byrinthea. In this paper, 14 new species are named versity, Cambridge, Massachusetts 02138. Current (Metepeira atascadero, M. cajahamba, M. calamuchi- address: Institute ofEvolutionaiyand Ecological Sci- ta, M. celestun, M. inca, M. lacandon, M. maya, M. ences, Leiden University, 2311 GP Leiden, The ohnec, M. pacifica, M. petatlan, M. pimungan, M. re- Netherlands; [email protected]. villagigedo, M. roraima, M. tarapaca); 11 newjunior Bull. Mus. Comp. ZooL, 157(1): 1-92, June, 2001 1 2 Bulletin Museum ofComparative Zoology, Vol. 157, No. 1 synonyms are reported (M. acostai, M. bani, M. dom- [1989]), but this practice can lead to trou- inicana, M. grinnelli, M. latigyna, M. perezi, M. san- ble. In one case, the behavior of several tnae,nsMis.);saflievie,cMas.esseodfiteirorsao,neMo.uslvyausryineoonryummi,zeMd.nvaimrgeis- different species was initially studied un- are reversed; 22 species and two subspecies are re- der the false assumption that they all be- described (M. arizonica, M. triangidaris, M. chilapae, longed to the same species (e.g., Uetz et M. Comanche, M. compsa, M. crassipes, M. datona, al., 1982). Clearly, a strong taxonomic sMo.gad,esMe.ndegrria,ndMi.osgaalaaltphienaae,,MM..ggrlaon7dneiroasbailigsr,anMd.iosgao,- foundation is important for further biolog- M. gressa, M. incrassata, M. jamaicensis, M. karkii, ical work. M. minima, M. nigriventris, M. rectangula, M. spi- Ultimately, the relatively small, indis- nipes, M. uncata, M. ventura, M. vigilax); and a key tinct genitalia and the relatively homoge- to allMetepeira speciesispresented. Inaddition,sev- neous abdominal patterns are to blame for eral ecological and life history observations are re- the weakness in our knowledge of Mete- ported forvarious species. peira taxonomy. Many ofthese species are INTRODUCTION undoubtedly hard to distinguish, and this fact has surely intimidated arachnologists The absence of a comprehensive revi- from taking on the painful task ofrevising sion of Neotropical Metepeira has left the the group. In the absence of good distin- taxonomy of this group in shambles. Over guishing characteristics, the catalogs of the years, a fair number of species have Bonnet (1957) and Roewer (1942) synon- been named, particularly by A. F. Archer, ymized the names of many Neotropical R. V. Chamberlin, and W. Ivie. However, species with the name Metepeira lahyrin- these efforts have been sporadic and, for thea. Levi's (1977) revision of Nearctic the most part, scant. For example, the de- species observes that M. labyrinthea is ac- scription of Metepeira dominicana (Ar- tually limited to the eastern United States. cher, 1965) provides little information oth- One task in this revision consists of reas- er than "form typical of Metepeira in all serting the names ofspecies that were im- respects," a few measurements, and two properly synonymized and clarifying the unrecognizable figures. Evenwhen species diagnostic characters that are needed to are properly described they have far less identify them. taxonomic value when published alone, in ACKNOWLEDGMENTS the absence ofa full comparative revision. i The poor understanding of Metepeira This paper is part ofmy Ph.D. thesis for taxonomy has persisted despite great eco- the Department of Organismic and Evo- logical and behavioral interest in this ge- lutionary Biology, Harvard University. I ain nus. Indeed, many species are obligate or indebted to many people for their help, facultative social species and offer excel- assistance, and encouragement in this pro- lent models for investigating genetic and ject. I am especially thankful for the ded- environmental factors that influence colo- ication and support of my advisors, Her- ny formation (e.g., Uetz and Cangialosi, bert W. Levi and Edward O. Wilson. I am 1986; Uetz et al, 1987). The monumental grateful that my colleagues in the Depart- work carried out over manyyears by G. W. ment of Invertebrate Zoology provided Uetz has made great strides in our under- such a pleasant place to work: Edward standing of gregarious social behavior in Cutler, Ardis Johnston, Laura Leibensper- spiders and in risk-sensitive foraging the- ger, Damhnait McHugh, Diana Sherry, ory in general (e.g., Uetz, 1996). Still, in Van Wallach, and Dee Woessner, among the absence of solid taxonomic literature, others. behavioral ecologists have been forced to Field collecting and new specimen ac- apply informal names to their study ani- quisitions were made possible with the mals (e.g., Metepeira "atascadero" in Uetz help ofGita Bodner, Fundacion Capacitar, [1989] or Metepeira "Species A" in Viera Tim Coonan (CINP), Fred Coyle, Dawn Metepeira • Piel Fitzpatrick, Germania Jacome, Antdnia turelles de Belgique, Brussels, Monteiro, Tila Perez, George Putnam, Belgium; L. Baert Linda Rayor, Grace Smith (NAWF), and JAK A. Kochalka, Ciudad Univer- J. George Uetz. I am particularlyindebtedto sitaria, Paraguay George Uetz for his assistance and corre- JEC Carico, Lynchburg, Virginia, J. spondence. United States I am thankful for the —comments by JMM Maes, Leon, Nicaragua those who read this paper especially to MACN JM.useo Argentino de Ciencias the members on my thesis committee: H. Naturales, Buenos Aires, Argen- W. Levi, N. E. Pierce, and E. O. Wilson. tina; E. A. Maury, C. L. Scioscia I am also indebted to Kathy Horton for MCN Museu de Ciencias Naturais, her help in formatting and preparing the Fundagao Zoobotanica do Rio manuscript and to the Colles Fund for de- Grande do Sul, Porto Alegre, fraying the costs of publication. Curators Rio Grande do Sul, Brazil; E. H. at various institutions who lent me speci- Buckup, M. A. L. Marques mens are listed in the Materials and Meth- MCZ Museum of Coniparative Zool- ods section. I cannot overstress the value ogy, Harvard University, Cam- ofmuseum collections andexpertcurators, bridge, Massachusetts, United without which research in taxonomywould States; H. W. Levi not be possible. Museum collections are MECN Museo Ecuatoriano de Ciencias the most important tools available for un- Naturales, Quito, Ecuador; Ger- derstanding biodiversity. mania Estevez Jacome MEG MATERIALS AND METHODS M. E. Galiano, Buenos Aires, Argentina Collections Examined. The taxonomic MLJC Maria Luisa Jimenez, Centro de revision was carried out on specimens bor- Investigaciones Bioldgicas del rowed from the following collections. The Noroeste, La Paz, Mexico abbreviations correspond to those listed MLP Museo de Universidad Nacional, with each record after eveiy species de- La Plata, Argentina; R. F. Arro- scription. I am grateful to the museums, zpide, C. Sutton curators, and staff that graciously loaned MNRJ Museu Nacional, Rio de Janeiro, the material. Brazil; A. Timotheo da Costa ADC A&M MNSD Museo Nacional de Historia A. Dean, Texas University, Natural, Santo Domingo, Re- College Station, Texas, United publica Dominicana; Felix Del States AMNH AHimsetroircya,n NMeuwseuYmorko,f NUantiutreadl MUSM MMounsteeo de Historia Natural, Universidad Nacional Mayor de States; N. Platnick, L. Sorkin BMNH Natural History Museum, Lon- San Marcos, Lima, Peru; D. Silva don, England; P. Hillyard MZSP Museu de Zoologia, Universida- CAS California Academy of Sciences, de de Sao Paulo, Sao Paulo, SP, eSdanStFartaensc;iCs.co,GrCiaslwioflordnia, Unit- MZUF BMruazsielo; P.ZoVoalnozgoilicnoi,dJe. L".LaLeSmpee- CV Carlos Valderrama A.; Bogota, cola" Universita di Firenze, Colombia Florence, Italy; S. Whitman FSCA Florida State Collection of Ar- NRMS Naturhistoriska Riksmuseet, thropods, Gainesville, Florida, Stockholm, Sweden; T. Krones- United States; G. B. Edwards tedt IRSNB Institut Royal des Sciences Na- PAN Polska Akademia Nauk, Warsza- 4 Bulletin Museum ofComparative Zoology, Vol. 157, No. 1 DMAAG ONG wa, Poland; Proszynski, A. Slo- ing contour maps, such as J. jewska, W. B. Jedryczkowski aeronautical maps; in most cases, elevation REL R. E. Leech, Edmonton, Alber- was estimated using NOAA data with an ta, Canada on-line database server (http://phylogeny. SMF Forschungsinstitut Sencken- harvard.edu/~piel/find.htiril). berg, Frankfurt am Main, Ger- The enhanced locality database was many; M. Grasshoff used to reveal ecological and life history SR Susan Riechert, Knoxville, Ten- traits. Seasonality ofspecies was expressed nessee, United States by plotting a circular histogram showing USNM National Museum of Natural the relative amount of collecting activity History, Smithsonian Institution, per 5-day interval (Figs. 300-337). While Washington, D.C., United locality dates alone cannot control for the States; Coddington, S. F. seasonal activityofhuman collectors, these J. Larcher data at least provide an estimate ofspider ZMB Zoologisches Museum der seasonal abundance, if only approximate. Humboldt Universitat, Berlin, Some syiupatric species show incongruous Germany; M. Moritz seasonal abundance, which is at least some ZMUC Zoologisk Museum, Gopenha- evidence that seasonality of spider collec- gen, Denmark; H. Enghoff, N. tors does not unduly overshadow the sea- Scharff sonality of the spiders themselves. ZSM Zoologische Staatssammlung, Examination and Illustration. Speci- Munich, Germany mens were examined under 80% ethanol in a dish with light and dark sand grains Locality Data Storage and Manipula- for specimen support. Digital photographs tion. Localitydata from eachcollectionvial of preserved specimens were taken were entered into a database designed us- through a Nikon SMZ-10 photomicro- ing Glaris FileMaker Pro®. Geographic scope using a Panasonic WV-CL320 GGD coordinates were added to locality data video camera, chosen for its high sensitiv- that lacked them using maps, USBGN gaz- ity to light. Video images were captured etteers, and on-line databases (http://164. using a Quicklmage®24 digitizer and ed- 214.2.59/gns/html/ and http://mapping. ited on a Quadra 700 Macintosh® com- usgs.gov/www/gnis/). Occasionally locality puter. The computer allows relatively in- information was illegible or unknown or expensive pictures to be printed rapidlyon one of several homonymous sites. In such a 1,200 dpi Xante® Accel-a-Writer 8200 cases a reasonable, educated guess was laser printer. Digital pictures were used to inade and a "[?]" designation was append- help sort out individuals to species, to cre- ed to the locality. In some cases the itin- ate publishable pictures of gross dorsal erary ofa collectorwas reconstructedfrom and ventral markings, and to aid in the il- other known records, and the ambiguous lustration ofgenitalia. As an aid in illustra- locality was assigned a coordinate halfway tion, the digital pictures functioned as a between the previous and followingknown camera lucida because they assured accu- collection sites. The locality database racywhen drawing the proportions ofgen- worked in concert with the mapping pro- ital parts and sclerites. Usually a digital gram Atlas Pro® to generate thematic picture was laid over carbon paper and an maps on the fly. These maps helped in the outline of the genitalia was transferred to process of delimiting species and discov- coquille board underneath. The illustra- ering cryptic species. tion continued on the coquille board using Elevation (in meters) was estimated for a Staedtler OmniGhrom® pencil and a each locality that lacked this information. drafting pen with India ink and then was In some cases, elevation was estimated us- scanned at 600 dpi on a LaGie Silverscan-