ie Shi gee Tae ficeniberiitie tt Lan ik civ? Eeeenerat se WN tai Tecan Tiesdates pee erie Sopot ease HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology The Systematics of Neotropical Orb-weaving Spiders in the Genus Metepeira (Araneae: Araneidae VOLUME 157, NUMBER 1 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 Jounsonia, Department of Mollusks, 1941-1974 OCCASIONAL PAPERS ON Mo.Luusks, 1945— SPECIAL 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, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4. Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology 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 of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the 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 of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1947. (Complete sets only.) Proceedings of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2001. THE SYSTEMATICS OF NEOTROPICAL ORB-WEAVING SPIDERS IN THE GENUS METEPEIRA (ARANEAE: ARANEIDAE) WILLIAM H. PIEL' CONTENTS FANS trac tees See LEER reed PA ae LY, 1 Imtrocductionie 2... 2e5 ao ae ek ee 2, Acknowledgments) #2 esses en ei ee neat ae 2} Materials and Methods __..- 3 Collections Examined __ 3 Locality Data Storage and Manipulation ___ 4 Examination and Illustration 4 Metepeira F. O. P.-Cambridge ~............--.-.-.-- 5 Key stoplemale Metepeind, yo sss. se seen ee 12 Key ston MialesMetenet ia te saws. se ee 2 lef IMICLenetnauoxts Group les saree mele. ewes Is) 1. Metepeira datona Chamberlin and Ivie 20 2. Metepeira desenderi Baert ___........ ail 3. Metepeira grandiosa grandiosa Chamberlin and Ivie 23 4. Metepeira grandiosa alpina Chamberlin and Ivie _ 24 Metenevrawicilasc: Group ese elo cian 26 5. Metepeira cajabamba New Species _.. 26 6. Metepeira glomerabilis (Keyserling) .. 28 7. Metepeira vigilax (Keyserling) —_... 30 8. Metepeira rectangula (Nicolet) 32 Metepeira labyrinthea Group ........-.-------------------- 33 9. Metepeira spinipes F. O. P.-Cambridge . 34 10. Metepeira lacandon New Species Oil Metepeira nigriventris Group —.........--.--------------- 38 11. Metepeira nigriventris (Taczanowski) 38 12. Metepeira tarapaca New Species _..... 40 13. Metepeira calamuchita New Species. 42 14. Metepeira galatheae (Thorell) 43 15. Metepeira karkii (Tullgren) 46 iMeteneinascompsa Croup =: at en eee 47 16. Metepeira compsa (Chamberlin) —__ 48 17. Metepeira roraima New Species —-...... D3 18. Metepeira gressa (Keyserling) — 54 Metepeira incrassata Group —.2--- 56 19. Metepeira maya New Species __.......... 56 20. Metepeira inca New Species -............... 58 ‘Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts 02138. Current address: Institute of Evolutionary and Ecological Sci- ences, Leiden University, 2311 GP Leiden, The Netherlands; piel@rulsfb.leidenuniv.nl. 21. Metepeira gosoga Chamberlin and Ivie Atcha Cadre Some iligteeetye tale SON ceed Siete teeta Te 59 22. Metepeira olmec New SESS 60. 23. Metepeira comanche Levi ____------- 62 24. Metepeira pimungan New Species ..... 62 25. Metepeira triangularis (Franganillo) _ 63 26. Metepeira arizonica Chamberlin and [vie tir ee aie Peasy sehen as 6 Ahead Bry) 66 27. Metepeira atascadero New Species ~.. 67 28. Metepeira incrassata F. O. P.- Cambridge yrs ene. pales ee ee 68 Metepeira ventura Group 20.0222 71 29. Metepeira ventura Chamberlin and vi See Renee el ET ae eee re: ee Bk 71 30. Metepeira revillagigedo New Species 73 31. Metepeira celestun New Species __... 74 32. Metepeira uncata F. O. P.-Cambridge _. 76 33. Metepeira crassipes Chamberlin and [hvalevnael Scie oe RM Den OLB ayiee elem ccd RELY a 34. Metepeira chilapae Chamberlin and vac jos epee ae Wee ae ee eee ee 78 Metepeira minima Group ~....--..--.------------------------- 80 35. Metepeira petatlan New Species —...... 80 36. Metepeira minima Gertsch _............-.- 82, 37. Metepeira pacifica New Species 84 38. Metepeira jamaicensis Archer 86 [Eaten abinen (Cue) 88 Ds Gl x es ee sae RE ree | Eu 91 ABSTRACT. Of the 39 species and three subspecies of the orb-weaver genus Metepeira in the Americas, 36 species and two subspecies are known to occur outside of the U.S. and Canada. Yet, despite their conspicuous webs, diurnal foraging, and _ relatively common presence, the taxonomy of Metepeira is poorly understood, probably because the genitalia are small and difficult to distinguish. In fact, many names for species south of the U.S. were, at some time, in- correctly synonymized with the name Metepeira la- byrinthea. In this paper, 14 new species are named (Metepeira atascadero, M. cajabamba, M. calamuchi- ta, M. celestun, M. inca, M. lacandon, M. maya, M. olmec, M. pacifica, M. petatlan, M. pimungan, M. re- villagigedo, M. roraima, M. tarapaca); 11 new junior Bull. Mus. Comp. Zool., 157(1): 1-92, June, 2001 il 2 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 synonyms are reported (M. acostai, M. bani, M. dom- inicana, M. grinnelli, M. latigyna, M. perezi, M. san- ta, M. salei, M. seditiosa, M. vaurieorum, M. virgi- nensis); five cases of erroneously synonymized names are reversed; 22 species and two subspecies are re- described (M. arizonica, M. triangularis, M. chilapae, M. comanche, M. compsa, M. crassipes, M. datona, M. desenderi, M. galatheae, M. glomerabilis, M. go- soga, M. grandiosa alpina, M. grandiosa grandiosa, M. gressa, M. incrassata, M. jamaicensis, M. karkii, M. minima, M. nigriventris, M. rectangula, M. spi- nipes, M. uncata, M. ventura, M. vigilax); and a key to all Metepeira species is presented. In addition, sev- eral ecological and life history observations are re- ported for various species. INTRODUCTION The absence of a comprehensive revi- sion of Neotropical Metepeira has left the taxonomy of this group in shambles. Over the years, a fair number of species have been named, particularly by A. F. Archer, R. V. Chamberlin, and W. Ivie. However, these efforts have been sporadic and, for the most part, scant. For example, the de- scription of Metepeira dominicana (Ar- cher, 1965) provides little information oth- er than “form typical of Metepeira in all respects,” a few measurements, and two unrecognizable figures. Even when species are properly described they have far less taxonomic value when published alone, in the absence of a full comparative revision. The poor understanding of Metepeira taxonomy has persisted despite great eco- logical and behavioral interest in this ge- nus. Indeed, many species are obligate or facultative social species and offer excel- lent models for investigating genetic and environmental factors that influence colo- ny formation (e.g., Uetz and Cangialosi, 1986; Uetz et al., 1987). The monumental work carried out over many years by G. W. Uetz has made great strides in our under- standing of gregarious social behavior in spiders and in risk-sensitive foraging the- ory in general (e.g., Uetz, 1996). Still, in the absence of solid taxonomic literature, behavioral ecologists have been forced to apply informal names to their study ani- mals (e.g., Metepeira “atascadero” in Uetz [1989] or Metepeira “Species A” in Viera [1989]), but this practice can lead to trou- ble. In one case, the behavior of several different species was initially studied un- der the false assumption that they all be- longed to the same species (e.g., Uetz et al., 1982). Clearly, a strong taxonomic foundation is important for further biolog- ical work. Ultimately, the relatively small, indis- tinct genitalia and the relatively homoge- neous abdominal patterns are to blame for the weakness in our knowledge of Mete- peira taxonomy. Many of these species are undoubtedly hard to distinguish, and this fact has surely intimidated arachnologists from taking on the painful task of revising the group. In the absence of good distin- guishing characteristics, the catalogs of Bonnet (1957) and Roewer (1942) synon- ymized the names of many Neotropical species with the name Metepeira labyrin- thea. Levis (1977) revision of Nearctic species observes that M. labyrinthea is ac- tually limited to the eastern United States. One task in this revision consists of reas- serting the names of species that were im- properly synonymized and clarifying the diagnostic characters that are needed to identify them. ACKNOWLEDGMENTS This paper is part of my Ph.D. thesis for the Department of Organismic and Evo- lutionary Biology, Harvard University. Iam indebted to many people for their help, assistance, and encouragement in this pro- ject. I am especially thankful for the ded- ication and support of my advisors, Her- bert W. Levi and Edward O. Wilson. I am grateful that my colleagues in the Depart- ment of Invertebrate Zoology provided such a pleasant place to work: Edward Cutler, Ardis Johnston, Laura Leibensper- ger, Damhnait McHugh, Diana Sherry, Van Wallach, and Dee Woessner, among others. Field collecting and new specimen ac- quisitions were made possible with the help of Gita Bodner, Fundacion Capacitar, Tim Coonan (CINP), Fred Coyle, Dawn Fitzpatrick, Germania Jacome, Ant6nia Monteiro, Tila Perez, George Putnam, Linda Rayor, Grace Smith (NAWF), and George Uetz. I am particularly indebted to George Uetz for his assistance and corre- spondence. I am thankful for the comments by those who read this paper—especially to the members on my thesis committee: H. W. Levi, N. E. Pierce, and E. O. Wilson. I am also indebted to Kathy Horton for her help in formatting and preparing the manuscript and to the Colles Fund for de- fraying the costs of publication. Curators at various institutions who lent me speci- mens are listed in the Materials and Meth- ods section. I cannot overstress the value of museum collections and expert curators, without which research in taxonomy would not be possible. Museum collections are the most important tools available for un- derstanding biodiversity. MATERIALS AND METHODS Collections Examined. The taxonomic revision was carried out on specimens bor- rowed from the following collections. The abbreviations correspond to those listed with each record after every species de- scription. I am grateful to the museums, curators, and staff that graciously loaned the material. ADC A. Dean, Texas A&M University, College Station, Texas, United States American Museum of Natural History, New York, United States; N. Platnick, L. Sorkin Natural History Museum, Lon- don, England; P. Hillyard California Academy of Sciences, San Francisco, California, Unit- ed States; C. Griswold Carlos Valderrama A.; Bogota, Colombia Florida State Collection of Ar- thropods, Gainesville, Florida, United States; G. B. Edwards Institut Royal des Sciences Na- AMNH BMNH CAS CV FSCA IRSNB JAK JEC JMM MACN MCN MCZ MECN MEG MLJC MLP MNRJ MNSD MUSM MZSP MZUF NRMS PAN METEPEIRA ° Piel 3 turelles de Belgique, Brussels, Belgium; L. Baert J. A. Kochalka, Ciudad Univer- sitaria, Paraguay J. Carico, Lynchburg, Virginia, United States J. Maes, Le6n, Nicaragua Museo Argentino de Ciencias Naturales, Buenos Aires, Argen- tina; E. A. Maury, C. L. Scioscia Museu de Ciéncias Naturais, Fundagao Zoobotanica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil; E. H. Buckup, M. A. L. Marques Museum of Comparative Zool- ogy, Harvard University, Cam- bridge, Massachusetts, United States: H. W. Levi Museo Ecuatoriano de Ciencias Naturales, Quito, Ecuador; Ger- mania Eistévez Jacome M. E. Galiano, Buenos Aires, Argentina Maria Luisa Jiménez, Centro de Investigaciones Bioldgicas del Noroeste, La Paz, Mexico Museo de Universidad Nacional, La Plata, Argentina; R. F. Arro- zpide, C. Sutton Museu Nacional, Rio de Janeiro, Brazil; A. Timotheo da Costa Museo Nacional de Historia Natural, Santo Domingo, Re- publica Dominicana; Félix Del Monte Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru; D. Silva Museu de Zoologia, Universida- de de Sio Paulo, Sao Paulo, SP, Brazil; P. Vanzolini, J. L. Leme Museo Zoologico de “La Spe- cola” Universita di Firenze, Florence, Italy; S. Whitman Naturhistoriska Riksmuseet, Stockholm, Sweden; T. Krones- tedt Polska Akademia Nauk, Warsza- 4 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 wa, Poland; J. Prészynski, A. Slo- jewska, W. B. Jedryczkowski R. E. Leech, Edmonton, Alber- ta, Canada Forschungsinstitut Sencken- berg, Frankfurt am Main, Ger- many; M. Grasshoff Susan Riechert, Knoxville, Ten- nessee, United States National Museum of Natural History, Smithsonian Institution, Washington, D.C., United States; J. Coddington, S. F. Larcher Zoologisches Museum der Humboldt Universitat, Berlin, Germany; M. Moritz Zoologisk Museum, Copenha- gen, Denmark; H. Enghoff, N. Scharff Zoologische Staatssammlung, Munich, Germany REL SMF SR USNM ZMB ZMUC ZSM Locality Data Storage and Manipula- tion. Locality data from each collection vial were entered into a database designed us- ing Claris FileMaker Pro™. Geographic coordinates were added to locality data that lacked them using maps, USBGN gaz- etteers, and on-line databases (http://164. 214.2.59/ens/html/ and __ http://mapping. usgs.gov/www/gnis/). Occasionally locality information was illegible or unknown or one of several homonymous sites. In such cases a reasonable, educated guess was made and a “[?]” designation was append- ed to the locality. In some cases the itin- erary of a collector was reconstructed from other known records, and the ambiguous locality was assigned a coordinate halfway between the previous and following known collection sites. The locality database worked in concert with the mapping pro- gram Atlas Pro™ to generate thematic maps on the fly. These maps helped in the process of delimiting species and discov- ering cryptic species. Elevation (in meters) was estimated for each locality that lacked this information. In some cases, elevation was estimated us- ing contour maps, such as DMAAC ONC aeronautical maps; in most cases, elevation was estimated using NOAA data with an on-line database server (http://phylogeny. harvard.edu/~piel/find. html). The enhanced locality database was used to reveal ecological and life history traits. Seasonality of species was expressed by plotting a circular histogram showing the relative amount of collecting activi per 5-day interval (Figs. 300-337). While locality dates alone cannot control for the seasonal activity of human collectors, these data at least provide an estimate of spider seasonal abundance, if only approximate. Some sympatric species show incongruous seasonal abundance, which is at least some evidence that seasonality of spider collec- tors does not unduly overshadow the sea- sonality of the spiders themselves. Examination and Illustration. Speci- mens were examined under 80% ethanol in a dish with light and dark sand grains for specimen support. Digital photographs of preserved specimens were taken through a Nikon SMZ-10 photomicro- scope using a Panasonic WV-CL320 CCD video camera, chosen for its high sensitiv- ity to light. Video images were captured using a QuickImage™24 digitizer and ed- ited on a Quadra 700 Macintosh® com- puter. The computer allows relatively in- expensive pictures to be printed rapidly on a 1,200 dpi Xanté™ Accel-a-Writer 8200 laser printer. Digital pictures were used to help sort out individuals to species, to cre- ate publishable pictures of gross dorsal and ventral markings, and to aid in the il- lustration of genitalia. As an aid in illustra- tion, the digital pictures functioned as a camera lucida because they assured accu- racy when drawing the proportions of gen- ital parts and sclerites. Usually a digital picture was laid over carbon paper and an outline of the genitalia was transferred to coquille board underneath. The illustra- tion continued on the coquille board using a Staedtler OmniChrom™ pencil and a drafting pen with India ink and then was scanned at 600 dpi on a LaCie Silverscan- ner II™. The resulting digital image was edited in Adobe Photoshop™ and reduced in size to 1,200 dpi. The edited figures were finally arranged on plates using Can- vas®, External genital structures were manip- ulated with pins to reveal hidden parts. The terminal division on the male palp is hinged, so it had to be pried open to see the embolus and embolic apophyses prop- erly. In females, mating plugs had to be removed from epigynal openings using pins. Sometimes the entire epigynum was partly cut from the body so as to see it from a posterior view. Internal genital structures were studied by clearing them in clove oil and examin- ing them using an Olympus BH-2 com- pound microscope. Sketches were made directly on the computer in Canvas™ by aiming the camera lucida at the computer monitor. While internal genital structures helped in the process of delimiting spe- cies, they did not prove to be as useful as external genital structures in describing species; thus, these working sketches are not figured herein. Measurements of the spiders were tak- en using a Leitz stereo dissecting micro- scope with a calibrated reticule. Sizes of leg articles, eyes, and carapace, were per- formed on one specimen of each sex, for each species. The respective localities of the candidate specimens were indicated in the descriptions. This study placed little reliance on spider leg measurements be- cause they are not usually very useful in spider taxonomy, and because Metepeira species are notorious for their variability in size (Levi, 1977; Piel, 1996). All eye sizes were reported as a ratio of the posterior median eye diameters to the diameter of every other eye type. For ex- ample, in the case of “ratio of eye diame- ters: posterior medians and anterior me- dians 2.0, anterior laterals 0.5, posterior laterals 1.0,” the reader should interpret the anterior medians to be half the size of the posterior medians, and the anterior lat- erals to be twice the size of the posterior METEPEIRA ° Piel 5 medians. Eye separations were expressed in terms of their own diameters, or in terms of the anterior lateral eyes when be- tween eyes of different types. Oval eyes were measured as an average of the lon- gest and shortest lengths. In parallel with the last revision of Me- tepeira (Levi, 1977), leg measurements were made on each article distal to the tro- chanter for the first leg and on the com- bined lengths of the patellae and tibiae for all remaining legs. Variation in total body size was provided as an average, minimum, and maximum of the total lengths from a number of mature specimens, usually cho- sen from a wide geographic spread. Metepeira F. O. P.-Cambridge Metepeira F. O. P.-Cambridge, 1903: 457. Type spe- cies by original designation M. spinipes F. O. P.- Cambridge 1903. The name is feminine. Diagnostic Abstract. Web combines bar- rier or scaffolding structure surrounding a classic araneid orb with a retreat suspend- ed in air (Fig. 1). Like a raccoon with its facial colors reversed, the eye region is lighter than any other part of the carapace (Fig. 2). The venter has a wide median white line set on a black background that, with only some exception, extends anteri- orly on the sternum (Fig. 3). With one ex- ception, the total lengths of distal leg ar- ticles (metatarsus and tarsus) exceed that of the middle articles (patella and tibia). The median apophysis has two distinctive flagella (F in Fig. 5) and, in some species, an easily recognizable keel (K in Fig. 5). The dorsal abdominal markings (the foli- um) look like an inverted fleur-de-lis, al- lowing easy recognition of the genus in the field (Fig. 2). Description and Diagnosis. For field ecologists, the most obvious and distinctive feature of Metepeira is the combination of orb and barrier web (Fig. 1). The barrier web forms scaffolding around an almost vertical orb and supports the spider's re- treat, which is thus suspended away from any substrate. In contrast to most araneids, the cara- 6 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 pace of Metepeira is lightest in the eye re- gion. However, this distinctive feature varies within the genus: in the case of M. rectangula (Nicolet, 1849), the lighter re- gion takes up almost half the carapace (Fig. 65); in the case of Metepeira F. O. P.-Cambridge, 1903, the lighter region is usually limited to the anterior edges of the carapace (Fig. 2). White, downy hairs of- ten cover the carapace but are especially white and conspicuous on the lighter parts of the carapace outside the eye region. In some species, such as M. spinipes, these hairs make the carapace look gray or sil- very when the spider is alive, but dark brown when the spider is in ethanol. The eyes of Metepeira are not particu- larly unusual. Eye separations relative to eye diameters increase with spider size: larger spiders tend to have relatively great- er eye separations. In either sex, the pos- terior median eyes are between 1.1 and 1.7 times the size of the anterior medians, and the separation between posterior me- dian eyes is between 0.4 and 0.7 of that between anterior median eyes. The sepa- ration between the anterior median eyes and the anterior lateral eyes is between 1 and 3.7 times the size of anterior median eyes in males and between one and five times the size in females. The diameter of the anterior median eyes exceeds the height of the clypeus. The shape of the female abdomen rang- es from wider than long and rhomboid (M. datona, Fig. 12) to roundish (e.g., M. de- senderi, Fig. 20; M. rectangula, Fig. 65), to longer than wide and oval (e.g., M. inca, Fig. 169). The dorsal folium has a recog- nizable white fleur-de-lis pattern, usually on a dark background, its edges shaped by a wavy, zig-zag white outline (Fig. 2). The dorsum of live spiders is often more red- dish—a pigment that rapidly dissolves in alcohol. Somewhat less common among other araneids is the median white line on the venter of the abdomen (Fig. 3), which is present (though shortened) even in the most darkly pigmented species. However, unique among araneid genera is the com- bination of median white line on the ven- ter and median white line on a black or brown sternum. Some Metepeira species lack a complete white line on the sternum, but even those, such as M. datona, that usually have an entirely black sternum nonetheless show hints of white markings in some specimens. Characteristics found in the carapace, abdomen, and sternum of Metepeira are also found in Araneus koepckeorum Levi, but this last species lacks the white line on the venter. With the exception of M. datona, and in some cases, M. desenderi, all Metepeira species have a combined metatarsus and tarsus that is longer than the combined pa- tella and tibia. This feature is unusual among araneids and is not found in Kaira O. P.-Cambridge or other likely relatives to Metepeira (Levi, 1977; Piel and Nutt, 1997). In most species the leg articles are ringed, usually with brownish black on the distal and dorsal surfaces of each article, except for the patellae and tarsi which are usually entirely dark. In mainly tropical and high-altitude species, the coxae are mostly black (e.g., Fig. 75), but in desert/ mesquite species they appear yellowish white (e.g., Fig. 28). Unlike many other araneids—and _ per- haps because of the small male size—the coxa on leg I of male Metepeira lacks the hook and corresponding groove typically found on femur II. In addition, males lack a tooth on the lateral side of the endite, and they lack a basal tooth on the palpal femur. The phylogenetic analysis of Scharff and Coddington (1997) incorrectly codes Metepeira as having a tooth on the endite. However, had the authors coded this character as absent, they would have decreased the length of their preferred tree because the nearest relatives hypoth- esized for Metepeira (Kaira, Zygiella, and Singa) also lack this tooth. Macrosetae usually concentrate on arti- cles that contact other spiders during mat- ing or grappling. In contrast to most gen- METEPEIRA ° Piel ri Barrier Web White Dorsal Light Eye Region / Fleur-de-Lis Pattern Median White Line on Sternum See \\\ , * if _ < We HUAI ROT SES Sa \ Median White it BS SVANS TS 7 D 3 Line on Black Venter Orb Web nigriventris incrassata minima labyrinthea compsa ventura Eee " —— — Flagellae Dy Lae \Y7 ‘on Stalk (+) : DEA (-) K on MA (-) F, Teeth on Face of K (+) DEA (+) | North America “ SC fEnlarged TA (+) : ; [) South America datona (1) Figure 1. Web of immature Metepeira grandiosa alpina from Chihuahua, Mexico. Figure 2. Dorsum of adult female Metepeira crassipes. Figure 3. Venter of adult female Metepeira tarapaca new species. Figure 4. Hypothetical phylogenetic relationships among Metepeira species groups. Shaded branches indicate species groups that live in South America; open branches indicate species groups that live in North America, Central America, and the Caribbean. Abbreviations: DEA, distal embolic apophysis; K, keel of median apophysis; TA, terminal apophysis; (+), character state gain; (—), character state loss. Figures 5, 6. Male palpus. 5, mesal view, Metepeira compsa. 6, ventral view of distal embolic division, Metepeira labyrinthea (Hentz). Abbreviations: BEA, basal embolic apophysis; C, conductor; DEA, distal embolic apophysis; E, embolus; F, flagellum on median apophysis; K, keel of median apophysis; MA, median apophysis; TA, terminal apophysis; TD, terminal division. Figures 7-13. Metepeira datona Chamberlin and Ivie (sp. 1; 17°53’N, 76°19’W). 7, male palpus, mesal. 8, epigynum, posterior. 9, epigynum, ventral. 10, male, dorsal. 11, male, ventral. 12, female, dorsal. 13, female, ventral. Scale bar: dorsum and venter figures 1.0 mm. 8 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 era related to Araneus, Metepeira has con- centrated macrosetae on femur I instead of tibia II (Scharff and Coddington, 1997). Female Metepeira have between two and five macrosetae on the anterior side of the femur, and between zero and seven on the anteroventral side. Males typically have more setae than their conspecific females: four to nine on the anterior side and two to nine on the anteroventral side. Variation in the number of macrosetae appears to correlate with body size. In most species, the male palpal tibia and patella each have two strong macrosetae (Levi, 1977, fig. 8). Compared with other araneid genera, Metepeira have rather small and similar genitalia, which on the one hand makes the genus easy to recognize, but on the other hand makes species tough to iden- tify. The small epigynum is fleshy, variable in shape, and weakly sclerotized. Unlike Araneus, Metepeira’s scape never has a pocket but always ends with a pointed tip (e.g., Fig. 31). The cleared epigynum— and in many cases the uncleared epigyn- um—reveals a pair of sclerotized spherical structures where the embolus is inserted, as well as ducts to pass semen to the larger, spherical seminal receptacles. In some species, these spherical structures are wide apart (e.g., Figs. 16, 17), in others they are tubular (e.g., Figs. 39, 40), but in many, they are closer together (e.g., Figs. 93, 94). Frequently the deeper, large seminal re- ceptacles can be seen through uncleared tissue (e.g., Figs. 201, 295). The male palp is more distinctive. In particular, the median apophysis (MA in Fig. 5), while not always a good character for separating closely related species, is ex- cellent when it comes to identifying the genus. Two flagellae (F in Fig. 5) grace- fully curve off the base of the median apophysis, and in some species, a toothed or smooth keel (K in Fig. 5) extends in the opposite direction. This design is also seen in Kaira, Aculepeira, and Amazonepeira, but none of these have flagellae that ap- pear so integral to the base structure. The terminal division on the Metepeira palp is very similar in almost all species. | When this structure is pulled up, a basal embolic apophysis (also known as an em- bolar lamella) can be seen in the shape of a club or spatula (E in Figs. 5, 6). Some- times a distal embolic apophysis can be seen if it is not hidden from view by an overhanging terminal apophysis. When the terminal apophysis is large and_sclero- tized—which is the case in all but the Me- tepeira foxi species group—it has a rec- ognizable toothed notch, like the mouth on a wrench (Fig. 6). Virgin males have a cap on the embolus that remains in the epigynum after mating and presumably serves as a barrier to subsequent mating (Levi, 1977). The shape of the embolus cap varies from tiny (e.g., Fig. 178) to short but wide (e.g., Fig. 199) to large and winged (Fig. 46). Finally, the terminal di- vision lacks a stipes—a sclerite between the radix and the embolus that is frequent- ly found in other genera related to Ara- neus (Scharff and Coddington, 1997). Natural History. All Metepeira species build a unique web that combines an orb with a barrier web (Levi, 1977; Lopez, 1993). As with Cyrtophora Simon or Me- cynogea Simon (Levi, 1997), the retreat of Metepeira hangs in the air, away from sub- strate, and is suspended by a scaffolding structure created by the barrier web (Fig. 1). The spider detects vibrations in the web and gains quick access to the hub us- ing a signal line that runs from the retreat to the center of the orb (Fig. 1). Tan col- ored egg sacs are strung together, usually above the retreat, and the most recently laid eggs are nearest to the spider. In some species the egg sacs and retreat are deco- rated with insect parts (e.g., M. spinipes); in other species they are carefully wrapped by leaves and woven together (e.g., M. da- tona). Unlike the webs of Cyrtophora and Mecynogea, the orb web of Metepeira is oriented vertically, and the number of radii and sticky spirals are more typical of other araneines. In some species, such as M. pimungan (personal observation) and, to a lesser de- gree, M. incrassata (G. Uetz, personal communication), juveniles and adults without eggs will live on webs lacking a suspended retreat. Instead, the spider sits on a white disk-shaped stabilimentum in the center of the hub. Of 110 M. pimun- gan specimens observed on San Miguel Is- land, about 40% occupied webs of this type. In two cases the disk stabilimenta were partly separated from the hub by barrier web lines and were further bent over to form a partly covered protective retreat for the spider. This observation makes it possible to imagine that the disk stabilimentum seen in M. pimungan re- sults from the fusion of the suspended re- treat with the hub. When food supplies are plentiful, spi- ders of all kinds show an increased toler- ance for one another and an increased ten- dency to aggregate (e.g., Gillespie, 1987; Rypstra, 1986). The suspended retreats and barrier webs of Metepeira, Cyrtopho- ra, and Mecynogea may actually further fa- cilitate in the formation of aggregations by easing dependency on substrate availabili- ty and by providing a common support sys- tem (Burgess and Witt, 1976; Uetz, 1986). In any case, colony formation is known to occur in: all three genera (e.g., Rypstra, 1979), but especially in Metepeira. Small colonial aggregations of two to 10 individ- uals occur in M. datona (Spiller and Schoener, 1989), M. minima (personal ob- servation), M. glomerabilis (R. Baptista, personal communication), and M. atascad- ero new species (e.g., Uetz and Hodge, 1990). Medium-size colonies of 10 to 30 individuals occur in M. pimungan (person- al observation), M. gressa (Viera and Cos- ta, 1988), M. nigriventris (L. Rayor, per- sonal communication), M. tarapaca (VN. Roth, locality label), and M. spinipes (e.g., Uetz, 1988a). Large colonies, sometimes in the thousands of individuals, commonly occur in M. incrassata (e.g., Uetz and Hodge, 1990). Near rivers and in other lush habitats, M. tarapaca colonies can reach 200 individuals (M. Roy, personal communication). These cases of social be- METEPEIRA ° Piel 9 havior, broadly spread across seven differ- ent species groups, may mean that aggre- gation is a frequently lost and relatively old trait, or it may mean that species are prone to converge and evolve the same behavior independently. Either way, much research has focused on elucidating the selective forces behind colonial behavior in Metepeira. In partic- ular, Uetz (1988a,b, 1996) has provided strong support for the hypothesis that Me- tepeira forage using a risk-sensitive strat- egy. He suggests that spiders in abundant habitats seek to minimize individual vari- ance in prey capture by aggregating in col- onies, whereas spiders in poor habitats seek to maximize variance by living soli- tarily—perhaps in a risky attempt to find areas of local prey maxima. The diversity of social tendencies among species is therefore commensurate of the diversity of ecological habitats that they inhabit. Indeed, Metepeira species thrive in a wide array of habitats, though often they are quite harsh. These include wet, mon- tane cloud forests in Mexico and Panama (M. incrassata, M. olmec); tropical and wet agricultural areas (M. uncata, M. vigilax, M. glomerabilis, M. roraima); high-eleva- tion pine forests (M. lacandon, M. nigri- ventris, M. grandiosa alpina); Canadian bogs (M. grandiosa palustris); deciduous forests in the eastern U.S. (M. labyrin- thea); Caribbean coastal shrubbery (M. da- tona, M. minima, M. triangularis, M. ja- maicensis, M. maya, M. celestun); Mexican mesquite grasslands (M. atascadero, M. chilapae); Patagonian dunes and scrub (M galatheae) and pampas grass (M. karkii); dry Californian buckwheat and sage (M. crassipes, M. ventura, M. foxi, M. grandio- sa grandiosa); and arid and semiarid de- serts (M. arizonica, M. inca, M. ventura, M. crassipes). Although some species (e.g., M. galatheae, M. spinipes, M. compsa) cov- er vast geographic areas and live in many different habitats, many species are more biogeographically restricted. In fact, sev- eral species follow narrow ecological zones that decrease in elevation with distance 10 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 from the equator (e.g., M. rectangula, M. vigilax, M. cajabamba, Fig. 36; M. arizon- CG, See 2i3)): Close cohabitation with different inter- and intrageneric species is not uncommon. Colonies of M. incrassata are known to contact webs of Nephila clavipes Linnaeus (Hodge and Uetz, 1996) and Mecynogea ocosingo and Gasteracantha cancriformis (personal observation). Often M. crassipes, M. ventura, M. foxi, and M. grandiosa grandiosa are collected together (Levi, 1977), as are M. minima and M. celestun (personal observation). Species that have been collected from identical localities, though not necessarily at the same time, include: M. chilapae and M. spinipes; M. chilapae and M. atascadero; M. karkii and M. galatheae; M. calamuchita new species, M. gressa, and M. galatheae; M. rectan- gula, M. calamuchita, and M. galatheae; M. compsa and M. gressa; M. vigilax and M. compsa; M. glomerabilis and M. vigilax; M. compsa and M. glomerabilis; M. comp- sa and M. nigriventris; M. conypsa and M. inca; M. datona and M. jamaicensis; and M. datona and M. triangularis. Despite the wide biogeographic ranges of M. compsa (Puerto Rico and south to Argentina, Map 8) and M. datona (His- paniola and north to Florida, Map 1), they nonetheless come geographically close to one another but do not overlap. It is hard to imagine that the hurricanes that fre- quently pass through the Caribbean, as well as the homogeneous island environ- ments, would not gradually cause these two species distributions to overlap. Per- haps these abrupt, disjunct distributions are a rare example of competitive exclu- sion in Metepeira, which in other species is not thought to be an important factor (Wise, 1983). Sphecid wasps are predators on Mete- peira. Locality labels indicated that M. pa- cifica has been found in the nests of Try- pargilum nitidum, T. tenoctitlan, and _T. bensoni. Jiménez and Tejas (1994) report that M. crassipes is the most frequent prey item in the nests of Trypargilum triden- tatum. Colonial spiders, such as M. incras- sata, are especially vulnerable to wasps, other spiders, sarcophagid flies (e.g., Ar achnidomyia lindae, A. rayorae), and hummingbirds (Hieber and Uetz, 1990; Lopez, 1989; Rayor and Uetz, 1990). Species Groups. Nearctic Metepeira were divided into two species groups: the M. labyrinthea group and the M. foxi group, based on the pattern on the ster- num and the shape of the median apoph- ysis (Levi, 1977). Baert (1987) questioned the taxonomic usefulness of the M. foxi species group (M. foxi, M. grandiosa, M. datona) because he found that M. desen- deri has both a keel on the median apoph- ysis and a white sternal line (Figs. 15, 21)—a combination that is incompatible by Levi's scheme. Nonetheless, the geni- talia of M. desenderi closely ally this spe- cies with the M. foxi group, so I am re- defining the M. foxi group based on purely genitalic characters. This is likely to be a basal, paraphyletic group (Fig. 4) (Piel and Nutt, 1997). Seven additional species groups are dis- tal to the M. foxi. These remaining species are united by sharing a large terminal apophysis that is sclerotized and _ usually studded with teeth or denticles. The M. vigilax group (M. vigilax, M. cajabamba, M. glomerabilis, M. rectangula) are united by large emboli with long scooplike basal embolic apophyses (Fig. 60). Unlike the remaining species, the terminal apophysis in this group—albeit large—does not ac- tually overhang or hide the embolus. In addition to an overhanging terminal apophysis, the remaining taxa are also united by a distal embolic apophysis that either protrudes (Fig. 76), curves off (Fig. 185), or is secondarily lost (Fig. 264). The M. labyrinthea group (M. labyrinthea, M. lacandon, M. spinipes) share a toothless, smooth keel on the median apophysis (Figs. 67, 69). The M. nigriventris group and the M. compsa group together share a median apophysis with teeth on the face of the keel (Figs. 92, 149). The M. incrassata “al e datona S = desenderi Map| Map 3 METEPEIRBA ° Piel 11 \ = grandiosa alpina ¢ e grandiosa grandiosa ~~ Se @ vigilax | + rectangula / \e ns j My f Se Map 4 i 2 Maps 1, 2. Metepeira foxi species group. 1, M. datona, M. desenderi. 2, M. grandiosa grandiosa, M. grandiosa alpina. Maps 3, 4. Metepeira vigilax species group. 3, M. glomerabilis, M. cajabamba. 4, M. vigilax, M. rectangula. group, the M. ventura group, and the M. minima group all lack a keel on the me- dian apophysis (Figs. 164, 222, 293). How- ever, both the M. compsa group and the M. incrassata group have epigyna with similar oval or round sclerotized rims (Figs. 151, 166), so it is likely that these are paraphyletic and consist of species leading up to a major North American (without a keel) and South American (with a keel) phylogenetic split (Fig. 4). The South American branch includes the M. compsa group (M. compsa, M. ro- raima, M. gressa) and, more distally, the M. nigriventris group (M. nigriventris, M. tarapaca, M. calamuchita, M. galatheae, M. karkii). This latter group is united by a distinctive and derived scape, which pro- jects out and down, creating a noticeable arch and overhang (Fig. 86). The remaining species all lack a keel on the median apophysis, and with one ex- ception (M. inca), they live exclusively in North America (from the Caribbean and Panama to Nevada). The M. incrassata group (M. gosoga, M. maya, M. inca, M. comanche, M. olmec, M. atascadero, M. ar- izonica, M. incrassata, M. triangularis, M. pimungan) are very likely paraphyletic. The epigynum on each species seems au- tapomorphic and difficult to unite with any others. Some species (M. gosoga, M. maya, M. inca) have a pointed or projecting distal embolic apophysis (Fig. 171). Others have 1 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 a distal embolic apophysis that curves off sharply but does not project forward (Fig. 185). Finally, others have a distal embolic apophysis that curves off gently, almost to the point of hiding the existence of an apophysis (Fig. 206). The M. minima group (M. jamaicensis, M. minima, M. pacifica, M. petatlan) and the M. ventura group (M. uncata, M. ven- tura, M. celestun, M. chilapae, M. revilla- gigedo new species, M. crassipes) are unit- ed by derived characters: both have thin flagellae arising from a thin base on the median apophysis (Fig. 264), and both share the secondary loss of the distal em- bolic apophysis. The M. minima group is clearly monophyletic; its species all have their flagellae further set off from the me- dian apophysis on a separate, narrow stalk (Fig. 286). KEY TO FEMALE METEPEIRA 1 Epigynal openings strongly sclerotized, round but tilted so that they appear oval from a ventral view (Figs. 40, 48, 55) pe ee IE EY eR eck. \od OL ONE earn 9} = Epigynal openings weakly sclerotized (e.g., Fig. 208), and if round, they are not tilted and do not appear oval from a ventral view (Fig. 131) Epigynal openings shaped like the en- trance to a snail shell (Fig. 55); His- paniola, Brazil, Argentina (Map 4) Be ER ee eee ee (7) vigilax = Epigynal openings tubular (Figs. 40, 48) Epigynal openings strongly tilted (Fig. 40) Pexul(Mapro)s = a (5) cajabamba = Epigynal openings weakly tilted (Fig. 48); Colombia to Brazil (Map 3) ———--- Sia EL We SSN Smee nee dat REO (6) glomerabilis Epigynal openings large and gaping, cre- ating large atria inside (Figs. 61, 62); Chile and Argentina (Map 4) ———— Seer Se eS ee (8) rectangula = Epigynal openings not gaping and not creating large atria (e.g., Fig. 69) ______.. 5 Weak posterior lobes on the epigynum create a single, wide epigynal depres- sion (Figs. 9, 17, 24, 31) or epigynum with crescent-shaped sclerotized open- ings on either side of a thin scape (Levi, 1977, fig. 87). Dark, sclerotized spheres below epigynal openings are greatly separated (Figs. 8, 16) 6 = Stronger posterior lobes on the epigynum create separate, smaller epigynal de- pressions that are not crescent-shaped, or if crescent-shaped, the scape is thick and puffy (e.g., Figs. 86, 102, 143, 166, 187, 208). Dark, sclerotized spheres below epigynal openings are closer to- gether (eseeehigs ss lOlen 42) sa eeeenes 11 Sternum with longitudinal white line (Fig. 21); Galapagos Islands (Map 1) yall ak ISN ae Aen al (2) desenderi ~ Sternum entirely black or brown (Figs. Igy OAS wis) Abdomen wider than long (Fig. 12); Flor- ida to Hispaniola (Map 1) _- (1) datona = Abdomen longer than wide (Figs. 27, 34) Coxae as black as sternum (Levi, 1977, fig. 98); Canada—U.S. border (Levi, ESO Ch suey o\\))) akin ee Delia (Levi, 1977: 212) grandiosa palustris = Coxae yellow or orange and lighter than Stemmum(Higsy28.5.3.0) a) as eee 9 Epigynum with crescent-shaped sclero- tized openings on either side of the scape (Levi, 1977, fig. 87); western U.S. and Canada (Levi, 1977, map 2) Pesce ses Ayr Os (Levi, 1977: 210) foxi = Epigynum with wide, transverse depres- sion (Figs. 23, 24, 30, 31) Scape wide and stubby (Fig. 24); Baja California north to Canada (Map 2) — BSI 5 Shad 2 (3) grandiosa grandiosa — Scape triangular (Fig. 31); in mountains from north-central Mexico to Canada (Mapa?) eee aes (4) grandiosa alpina Scape thickness equal to greater than width of epigynal depressions (e.g., Figs. 86, 119, 131, 143, 180, 201, 266) 9(8) 10(9) 11(5) = Scape narrower than epigynal depres- sions, or epigynal depressions in the shape of longitudinal slits (e.g., Figs. D3 8¥ 2520230) iain cies ks Se eae ana 35 12(11) Base of scape originates anteriorly and projects ventrally before curving pos- teriorly. This projection creates an overhang and a noticeable gap between the scape and the genital openings (e.g., Figs. 85, 86, 101, 102, 119, 143) = Scape does not create a noticeable gap or overhang (e.g., Figs. 77, 78, 130, 131, NGS GGMO6 SIN Veit te eine ee eens 18 13(12) Rim of epigynal depressions slightly sclerotized and oval-shaped (Fig. 143); northern Brazil, French Guiana, and Colombia (Map 8) (17) roraima - Epigynal depressions without distinct rim (Fig. 123) or not sclerotized (e.g., Fig. OD )\eees 3 = aR A a ee 14 14(13) Sternum black, coxae mostly black (Fig. A calamuchita g@ karkii @ nigriventris # compsa ® roraima @ gressa mi Skangse A PN as, Rg METEPEIBA ° Piel 13 Maps 5, 7. Metepeira nigriventris species group. M. calamuchita, M. karkii, M. nigriventris, M. galatheae, M. tarapaca. Map 6. Metepeira labyrinthea species group. M. spinipes, M. lacandon. Map 8. Metepeira compsa species group. M. compsa, M. roraima, M. gressa. 91), and carapace without lighter me- 15(14) Stermum brown to black with parallel dian mark (Fig. 90); high altitudes in Bolivia and Peru near Lake Titicaca (IMfayp 35) hy Wetec ee AE aol ae (11) nigriventris = Sternum with median white line (Fig. 99), or if sternum is black, then either the carapace has a median lighter ar- row-shaped mark (Fig. 113) or the cox- = ae are mostly yellow (Figs. 114, 128) lines on either side of median white line on venter: the parallel lines are thicker anteriorly than posteriorly (Fig. 128). Lower lip on epigynum thick and bulbous (Fig. 123); southern Argentina and southern Chile (Map 5) —_ (15) karkii Sternum with median white line, or if sternum is entirely brown to black, then parallel lines on either side of 14 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 white line on venter are either absent (Fig. 114) or equally thick anteriorly as posteriorly (Fig. 99). Epigynum with- out thickened lower lip (Figs. 94, 102, AUMIES Seite? (0) PRs. CA eee oS, ee 16 16(15) Dark, sclerotized spheres in epigynal openings small (Figs. 119, 120). Ante- rior lip of epigynum rounded off, sometimes with openings shifted pos- teriorly and wrinkled portion of the scape’s hood shifted more to the epi- gynum proper (Fig. 118); Chile and Argentina (Map 7) -............-- (14) galatheae = Dark, sclerotized spheres in epigynal openings large, and anterior lip of epi- gynum not rounded off. Openings al- ways located midway down the epigyn- tural (ESI 9.539 yp) 2) eee ae eee ee ee 17 17(16) General shape of epigynum is triangular with posterior width greater than an- terior width. Scooped-out depressions project more posteriorly than laterally (Fig. 102); north-central Argentina (MIEN oe) ieee a (13) calamuchita — General shape of epigynum square. Scooped-out depressions project more laterally than posteriorly (Fig. 94); northern Chile and southern Peru (Miapiii)) ea Sot ke Se ee (12) tarapaca 18(12) Epigynal openings almond-shaped, not noticeably sclerotized, and at a 40° to 60° angle from axis of spider (Figs. 69, cS) ise ene LES Saat. CE Awaba aut aie 19 = Epigynal openings not almond-shaped (eg, Figs. 180, 187, 201), or if al- mond-shaped, slightly sclerotized and at an angle from spider's axis of 80° to 100° (larress, IMSL ISIS IGi6) 2) 19(18) Almond-shaped openings created by de- pression where scape arises from epi- gynum (Fig. 173); California, Arizona, northwestern Mexico (Map 9) Se a = Se ee eee a (21) gosoga — Almond-shaped openings created by membranes inside depressions and not associated with scape (Figs. 69, 78) __ 20 20(19) Distinct C-shaped depression created where the scape arises from the epi- gynum (Fig. 78). Black marks inside al- mond-shaped openings not cross-eyed in appearance (Fig. 78). Black ster- num, usually with white spot in center (Fig. 83); mountainous regions of Chia- pas, Mexico (Map 6) _....... (10) lacandon — Indistinct depression created where the scape arises from the epigynum (Fig. 69). Black marks inside almond-shaped openings cross-eyed in appearance (Fig. 69). Sternum black, with or with- out a median white line. If with only a portion of a median white line present, usually only at the posterior end of the sternum (Fig. 75). Never with only one GOUCS) KYOTOLE TH SIME 21 21(20) Because of interspecific variability and polymorphism, females of the follow- ing two species are almost impossible to separate reliably without molecular sequence data. Small ribosomal sub- unit (12S) mtDNA sequence data has the following diagnostic markers. Base 14261: ACGGT: base 14285: ATTTT;: base 14361: ACTAC; base 14394: CTTAT: base 14412: ATTA. (Base numbers refer to homologous sites in the mitochondrion of Drosophila yak- uba, as reported by Clary and Wolsten- holme [1985].) One quarter of scape extends below lower lips of epigynum (Levi, 1977, fig. 14); New England to Florida and west to eastern Texas (Levi, 1977, map 1, but not including points appearing in Mexico) ———--—— Spe cee tah Se lsP PEs (Levi, 1977: 196) labyrinthea = For 125 mtDNA sequence data, the fol- lowing sequences are diagnostic. Base 14261: ACGAT;: base 14285: ATCTT: base 14361: ACCAC: base 14394: CTAAT: base 14412: TTTA. One third of scape extends below lower lips of epigynum (Levi, 1977, fig. 21; Fig. 69); Mexico City north to California (Map CH Rpereereeeeee CAE wi ROM mM INC n ls ns ou" (9) spinipes 22(18) Epigynal openings small, round, and sclerotized (Fig. 131). Openings some- times hidden by wide scape (Fig. 134); Puerto Rico to Argentina and Chile (Miayp.S)) 2 ia eee) Sees (16) compsa = Epigynal openings not small, round, and SCIETOLZE Ci, ee Wan ees nee ie eee 23 23(22) Rim of epigynal openings sclerotized and in an oval or teardrop shape (Figs. 151, 5 OSA GGE AUST srsstaet 6 es Rien ee ee 9A - Rim of epigynal openings not sclerotized in an oval or teardrop shape (Figs. 180, NOAL 9 Oilise2, 08) Ls Ai eee OAT 24(23) Epigynal openings oval, small, and partly hidden by scape (Fig. 151); northern Argentina, Uruguay, and southern Bra- Zilt (Miaps8)\ see ener t re ied (18) gressa — Epigynal openings teardrop-shaped (Fig. 187) or large and oval but not hidden loyy Sroatoxsy (laress, ISS), TGS) 25(24) Epigynal openings teardrop-shaped (Fig. 187); northeastern Mexico to Texas (Kevin oiieemapeleMap >) == ee (23) comanche = Epigynal openings oval-shaped (Figs. IL) OS SG. G)) sieesas. hs Cen eae ee eee SUE 26 26(25) Lower lip of epigynum pointed (Fig. 166), abdomen white (Fig. 169); north- emmytipyorkerul (Mapa lil) james (20) inca METEPEIRA ® Piel 15 + arizonica ~~ @ gosoga ce Bal ais @ pacifica % inca olmec incrassata mg petatlan @ pimungan Bre Map 9 Map 10 Map 11 ta @ uncata w ventura @ crassipes a 4 celestun g revillagigedo ® chilapae @ jamaicensis B minima # atascadero ® triangularis Map 14 Maps 9, 11, 14. Metepeira incrassata species group. 9, M. arizonica, M. gosoga, M. incrassata, M. pimungan. 11, M. comanche, M. inca, M. olmec. 14, M. maya, M. atascadero, M. triangularis. Maps 10, 15. Metepeira minima species group. 10, M. pacifica, M. petatlan. 15, M. minima, M. jamaicensis. Maps 12, 13. Metepeira ventura species group. 12, M. uncata, M. crassipes, M. revillagigedo. 13, M. ventura, M. celestun, M. chilapae. — Lower lip of epigynum thickened but not or, if sclerotized, not in shape of sun- pointed (Fig. 159), abdomen dark (Fig. glasses) (ce: Higs1208.( 224) ieee ee 28 162); southern Mexico and Belize to 28(27) postenion epigynal lobes converge behind Costa Rica (Map 14) ___.... (19) maya the scape so that epigynal depressions 27(23) Rim of epigynal openings sclerotized and appear closed off from ventral view shaped like a pair of sunglasses (Fig. (HigswilS08 208521160224) aia a 29 201); eastern Cuba and Hispaniola = Posterior epigynal lobes end before they (Mapp rll) ees re hh. (25) triangularis disappear behind the scape so that epi- = Rim of epigynal openings not sclerotized gynal depressions appear open poste- 16 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 riorly from ventral view (Figs. 194, DAD. Oko 6 6) ea eran ie eed ne A a 32 29(28) Scape relatively long and thin, epigynal depressions large and round, large black spheres take up almost all the space in the depressions (Fig. 180); southern Veracruz to Panama (Map 11) BAS Mt 60 Aine on Bere em (22) olmec = Scape relatively short, fat, and fleshy. Epigynal depressions not perfectly round; black spheres do not take up most of the space in the depression (Figs. 2082268224) ee eens 30 30(29) Epigynum puffy, scape so thick that de- pressions on either side of scape ap- pear crescent-shaped (Fig. 208); south- western U.S. to central Mexico (Map Oe tie ee SOR. EME eel eels (26) arizonica = Epigynum not puffy, scape not so fat that depressions become crescent-shaped (Bigs DIGHOOA) ea tah tine ven 31 31(30) Very social. Epigynal depressions large, disk-shaped, with shiny-smooth scler- otized inner surfaces and thin posterior lips (Fig. 224). Sternum mostly black, sometimes with anterior white marks. Venter without U-shaped mark circum- scribing median white line posteriorly. Coxae mostly dark brown (Fig. 228); mainly in southern Veracruz (Map 9) fete Biri tithe Serene eRe ey it's CoP (28) incrassata = Mostly solitary. Epigynal depressions small, oval, with reduced shiny-smooth sclerotized surfaces and thick posterior lips (Fig. 216). Sternum with median white line or only white mark at pos- terior end. Venter with U-shaped mark circumscribing median white line pos- teriorly. Coxae mostly yellow (Fig. 221); central Mexican plateau (Map IAS) mrs rae eee een Re ee (27) atascadero 32(28) Small square-shaped epigynal depres- sions on either side of scape (Figs. 259, DGG) pte vttts eased ook kale Reisen oar Sire end? cate i) = Large, more rounded epigynal depres- sions (Fig. 194) or with straight to S- shaped edges mostly covered by scape (Bigs 245) iernot darstinvie Ni oi8 cella 34 33(32) Dark spheres inside epigynal depressions appear slightly walleyed. Scape rela- tively narrower at base: about the width of the depressions (Fig. 266); southeast and south-central Mexico (Mia ppl) isiat as eee ae ee (34) chilapae — Dark spheres inside epigynal depressions appear slightly cross-eyed. Scape rela- tively wider at base: about twice the width of the depressions (Fig. 259); northwestern Mexico and California (Map. 1:2) fs teas a At se ties (33) crassipes 34(32) Large, rounded epigynal depressions with large black spheres inside (Fig. 194). Sternum black with posterior dewdrop-shaped mark. Pair of short parallel lines on either side of ventral median white line (Fig. 198). Carapace with large anterior white region (Fig. 197); San Nicolas Island off southern California (Map 9) (24) pimungan Epigynal depressions with straight to slightly S-shaped edges, mostly hidden by a wide triangular scape (Fig. 245). Sternum with wide median white line. No parallel white lines on either side of ventral median white line (Fig. 249). Carapace with small anterior white re- gion (Fig. 248); Yucatan Peninsula (Mato 1:3) ate en career (31) celestun Epigynal depressions wider than long (Figs, 23015059) 1b oN Wiel cobae 36 Epigynal depressions longer than wide (e.g., Figs. 238, 273, 280, 288) 37 Black comma shapes inside epigynal de- pressions (Fig. 231). Sternum with me- dian white line (Fig. 235). Dorsum lightly pigmented (Fig. 234); north- western Mexico and coastal California (Miao: S))ant= es ONE eee (29) ventura Black S-shaped marks inside epigynal de- pressions (Fig. 252). Sternum black with dewdrop mark at posterior end (Fig. 256). Dorsum darkly pigmented (Fig. 255); Guatemala and Costa Rica (Mayon 12) 2a else besornei ty byee (32) uncata Epigynal depressions indistinct anterior- ly. Black comma-shaped marks inside depressions and covered by translucent membranes (Fig. 238); Isla Socorro of the Archipiélago de Revillagigedo (Mapel2)ie 2s ee (30) revillagigedo Epigynal depressions distinct anteriorly. Black spheres shifted laterally and lo- cated outside the depressions (e.g., Figs. 273, 280, DOS) gree See eae bat ees: 38 Epigynal depressions slit-shaped and usually narrower than scape (Figs. 280, 281): northwestern Mexico and Yuca- tan Peninsula (Map 15) ___.... (36) minima Epigynal depressions oval and wider than Scapen(Hicsh:27/3 92.05) yaeeeeann aaeeeaennens 39 Dark spheres larger than epigynal open- ings (Fig. 273); west coastal Mexico (Mapytl 0) ees ree a (35) petatlan Dark spheres smaller than epigynal open- Ingsa(MICSHZO S29 5) pees een ernene 40 V-shaped ridge under the scape. Dark spheres located behind the junction where the ridge meets the lateral edge of the epigynal depressions (Fig. 288); Honduras to Costa Rica (Map 10) —— se te et See Pe (37) pacifica Straight ridge under the scape. Dark spheres located laterally and outside of the junction where the ridge meets the lateral edge of the epigynal depressions (Fig. 295); Cayman Islands, Jamaica, and Haiti (Map 15) -.......... (38) jamaicensis KEY TO MALE METEPEIRA Terminal apophysis thin, small, fleshy, without teeth or sclerotized parts (Figs. OES ES SOR OAS)) | kara Rai tae Lae Meee. 5 RE 2, Terminal apophysis enlarged, meaty, with teeth or sclerotized parts (e.g., Figs. 3 SMO ORSAT a9 9) eee ee ee 7 Terminal apophysis narrow; embolus curled clockwise like the tip on a cork- screw (Figs. 7, 15) Terminal apophysis wide; embolus tilted up and L-shaped (Figs. 22, 29) 4 Curled embolus with a ridge on the up- per surface and raised on a pedicel; basal embolic apophysis enormous (Fig. 15). Sternum with wide median white line (Fig. 21); Galapagos Islands (AWE T opal) Sree eee een es (2) desenderi Curled embolus smooth on its upper sur- face and not raised on a pedicel; basal embolic apophysis hardly noticeable (Fig. 7). Sternum entirely black (Fig. 11); Florida to Hispaniola (Map 1) SS ha ae iene NT (1) datona L-shaped embolus at an acute (< 90°) an- gle (Levi, 1977, figs. 91-93); western U.S. and Canada (Levi, 1977, map 2) Be ies robes i (Levi, 1977: 210) foxi L-shaped embolus at angle of 90° or preate;rs (igs# 22-229) Wate Se ees 5 Median apophysis with rounded projec- tion on dorsal side; jagged posterior edeejoiikeel (Bigsk 220029), sas 6 Median apophysis without projection— flat on dorsal side; rounded posterior edge of keel (Levi, 1977, fig. 105); along the Canadian and U.S. border, north to Nova Scotia and British Co- lumbia, south to Maine and North Da- kota (Levi, 1977, map 2) eid es (Levi, 1977: 212) grandiosa palustris Lower, transverse part of L-shaped em- bolus longer than vertical part (Fig. 29): mountains from north-central Mexico to Canada (Map 2) —-—---— pee De Sets Sees See ene (4) grandiosa alpina Lower, transverse part of L-shape em- bolus shorter than or equal to vertical part (Fig. 22); Baja California north to Canada (Map 2) ~ (3) grandiosa grandiosa Terminal apophysis does not entirely overhang the embolus. Embolus long and robust, with a long and thin gap created between the embolus and the 11(7) 14(13) 15(12) METEPEIRA ° Piel 17 basal embolic apophysis (Figs. 38, 46, 53, 60) Terminal apophysis often overhangs the embolus, covering it from view. Em- bolus not long and robust, without long, thin gap between embolus and basal embolic apophysis (e.g., Figs. 67, 79, 92, 121, 141, 185) Longer flagellum as thick as shorter fla- gellum. Keel short, slim, and feather- shaped. Embolus as wide as base of median apophysis (Fig. 53); Hispanio- la, Brazil, Argentina (Map 4) —. (7) vigilax Longer flagellum thicker than shorter fla- gellum. Keel absent (Fig. 38) or wide and arrowhead-shaped. Embolus thin- ner than base of median apophysis (HiGS38. 465.60) eee dt clio 9 Keel absent or greatly reduced (Fig. 38); Peru. (Map 3) ae (5) cajabamba Keel present (Figs. 46, 60) Large embolus as long as basal embolic apophysis (Fig. 60). Normal embolic cap; Chile and Argentina (Map 4) —— es ee ieee RENE aSAL) (8) rectangula Small embolus shorter than basal embol- ic apophysis, often seen with winged embolic cap (Fig. 46); Colombia to Brazile(Map! 3) 2. ses (6) glomerabilis Median apophysis with keel (e.g., Figs. QONIAT py SEs cele Rok aan ee Seer ee 12 Median apophysis without keel (e.g., Figs. 157, 178, 286); all North Ameri- can or Caribbean species except for M. UVC CAEN SRS et eS 29; Keel without teeth; smooth (e.g., Figs. 67, 76); North America _........---------- 13 Keel with teeth on face; rough (e.g., Figs. 84, 92, 100); South America 15 Distal embolic apophysis sleek, pointed, and feather-shaped when viewed from underside of terminal division (Fig. 70); Mexico City north to California (OMEN op 6) ak te eee ees (9) spinipes Distal embolic apophysis spoon-shaped (Fig. 6) or widened with bump (Fig. 79) Distal embolic apophysis spoon-shaped (Fig. 6). Sternum reddish brown with median white line. New England to Florida and west to eastern Texas (Levi, 1977, map 1, excluding points in Mexico) -....... (Levi, 1977: 196) labyrinthea Distal embolic apophysis widened with bump (Fig. 79). Sternum black with or without faint white mark in center (Fig. 81); mountainous regions of Chia- pas, Mexico (Map 6) _...... (10) lacandon Distal embolic apophysis a simple exten- sion that projects forward, parallel to the embolus. Keel usually rounded 18 16(15) 17(16) 18(17) 19(18) 20(19) 21(16) Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 (Fig. 129); Puerto Rico to Argentina and Chile (Map 8) (16) compsa Distal embolic apophysis raised up or projected away from the embolus. Keel usually pointed or jagged (Figs. 84, 92, TOO, WSLS WAL TANS) ee 16 Distal embolic apophysis points up and away from the embolus. Keel usually jagceda(Euigsael 4k 4Q) ieee eee 21 Distal embolic apophysis wide and raised up on boss. Keel usually pointed (Figs. SARAO DSi OO) eater oad eh a eee 17 Dewlap extension under embolus (Fig. 100) curves under, narrowing the gap between the embolus and the basal embolic apophysis by one-half (Fig. 103); north-central Argentina (Map 5) ESSE RRO Soe i Tes (13) calamuchita No such dewlap (Figs. 84, 92, 110, 121). Gap between embolus and basal em- bolic apophysis narrow by less than one-half the widest distance (Figs. 87, Ot Meili QA) Meant telat en dete cle Outside edge of distal embolic apophysis gently rounded when viewed from un- derside of terminal division (Fig. 124); southern Argentina and southern Chile (Map 5) (15) karkii Outside edge of distal embolic apophysis with distinct bump when viewed from underside of terminal division (Figs. Sie) Spell ee loa ta aE te Outside edge of distal embolic apophysis with rounded bump that is sclerotized and has a distinct line rising up from embolus (Fig. 117); Chile and Argen- tina (Map 7) (14) galatheae Outside edge of distal embolic apophysis with pointed bump, less sclerotized than embolus proper and without a dis- tinct line rising up from the embolus (ESS pS i159.) ee aie eee Berchet rah hele Thick neck joining embolus and _ basal embolic apophysis. Bump on distal em- bolic apophysis peeks out from under the terminal apophysis (Figs. 92, 95). Sternum usually with white spot or me- dian white line (Fig. 97); northern Chile and southern Peru (Map 7) osc enone ena (12) tarapaca Thin neck joining embolus and basal em- bolic apophysis. Bump on distal em- bolic apophysis does not peek out from under the terminal apophysis (Figs. 84, 87). Sternum always black, general col- oration dark (Figs. 88, 89); high alti- tudes in Bolivia and Peru near Lake Titicaca (Map 5) (11) nigriventris Distal embolic apophysis thinner than embolus near the junction of the two. Embolus tip is curved gently (Fig. 20 ES bo ey) 149): northern Argentina, Uruguay, and southern Brazil (Map 8) _. (18) gressa Distal embolic apophysis the same size as embolus near the junction of the two. Embolus tip is curved abruptly (Fig. 141): northern Brazil, French Guiana, and Colombia (Map 8) (17) roraima Distal embolic apophysis a projecting bumpy (hig ste oi 64 5oli/fll) eeneeern na 23 Distal embolic apophysis curved off (Figs. 178, 185, 199, 214), rounded (Figs. 192, 206), or absent (Figs. 264, DASE 29 Bhs. NEA Rates. Sel eae teat AE Distal embolic apophysis raised up from the embolus and pointed (Fig. 171); California, Arizona, northwestern Mex- Tato) (NYY oy 0) a ee (21) gosoga Distal embolic apophysis not raised up, but projecting forward and rounded off (Bigs.057 164). 0 a Wa Main flagellum on median apophysis thin and initially as thick as base. Embolus and embolic cap shortened (Fig. 164); northern tip of Peru (Map 11) —. (20) inca Main flagellum on median apophysis thick, but initially thinner than base. Embolus and embolic cap elongated (Fig. 157); southern Mexico and Belize to Costa Rica (Map 14) (19) maya Embolus shaped like the nib on a foun- tain pen, with bump near tip on op- posite side of distal embolic apophysis (Figs. 214, 217); central Mexican pla- 25 24 teaua(Mapel4) aa (27) atascadero Embolus without such bump near tip (Ces AhIE R222) AGS eee ee 26 Distal embolic apophysis abruptly ends in sharp curve and flagellae not set off on a narrow stalk (Figs. 178, 185, 199, DOO Ni tht S rule Sean tibet ee SOURS eaters een see ik Distal embolic apophysis gently curved off (Figs. 229, 236, 243), rounded (Figs. 192, 206), or absent (e.g., Fig. DO BN) (52! wae ake betel erred aa ee Flagellae very thin and equal in length. Embolus long and arching, with wide but short embolus cap. Overhanging terminal apophysis covers only a distal portion of the sclerotized part of the embolus (Fig. 199); eastern Cuba and Hispaniola (Map 14) _. (25) triangularis Flagellae normal in thickness and usually of different lengths. Terminal apophy- sis centered above the entire sclero- tized portion of the embolus (Figs. 178, 185, 222) Not known to be very social. Height of embolus plus distal embolic apophysis just before the latter curves off sharply is equal to or greater than length of embolus tip distal to this point (Figs. 178, 185). Sternum with median white Markie sa (IGS gl S25 GQ) asec e ns 29 = Highly social species. Height of embolus plus distal embolic apophysis just be- fore the latter curves off sharply is less than the length of the embolus tip dis- tal to this point (Fig. 222). Sternum mostly black; sometimes with anterior white marks. Coxae mostly dark brown (Fig. 226); mainly in southern Vera- cruz, Mexico (Map 9) -........ (28) incrassata 29(28) Darker, sclerotized portion of the embo- lus does not extend over the hump of the distal embolic apophysis. Base of embolus narrower than widest part of the first flagellum (Fig. 178); southern Veracruz, Mexico to Panama (Map 11) oP NOS eee (22) olmec = Darker, sclerotized portion of the embo- lus extends over the hump of the distal embolic apophysis. Width of embolus base the same or greater than widest part of the first flagellum (Fig. 185); northeastern Mexico to Texas (Levi, lOv@eemape "Mapa ie es (23) comanche 30(26) Distal embolic apophysis rounded off to form convex curve (Figs. 192, 206) _..... 31 = Distal embolic apophysis gently falls off to form concave shape (Figs. 929, 236, 243)! or absent (e:¢., Fig.) 293) i222 32 31(30) Embolus S-shaped (Fig. 192). Sternum black with posterior white mark. Ven- ter without white anchor shape mark (Fig. 196); San Nicolas Island off southern California (Map 9) —-—-——— See eee ee a ES (24) pimungan = Embolus convex on upper surface, straight on lower surface (Fig. 206). Sternum black with median white line. Venter with faint white anchor-shaped mark posterior to median white line (Fig. 210); southwestern U.S. to cen- tral Mexico (Map 9) _........ (26) arizonica 32(30) Flagellae on median apophysis set off on separate, narrow, stalk (e.g., Figs. 278, ORNG Nee eae ee Sa ae eee 38 ~ Flagellae on median apophysis not set off on a separate, narrow, stalk (e.g., Figs. 99959 GA) et 2 eRe OU) aa eer 33 33(32) Larger flagellum twice as wide as smaller flagellum (Fig. 236); Isla Socorro of the Archipiélago de Revillagigedo (Map UT: ip ater esate Bene fe ttt sede a (30) revillagigedo = Larger flagellum less than twice as thick as smaller flagellum (e.g., Figs. 229, GALEN Re Tienes Beter Mie Laut, aan eee 34 34(33) Flagellae on median apophysis relatively (dawn) (URakas PABKO), PAGV)) Le ee 35 - Flagellae on median apophysis thicker (Figs. 2.2 QUOTAS UES 517) es seemed Me ake EERE 36 35(34) Embolus with sharp bend near the tip METEPEIRBA ° Piel 19 (Fig. 264). Sternum with median white line (Fig. 268); southeast and south- central Mexico (Map 13) -_.... (34) chilapae = Embolus with gentle bend further from the tip (Fig. 250). Sternum mostly black with small white mark at poste- rior end of sternum (Fig. 254); Gua- temala and Costa Rica (Map 12) ——— DDS 5 RE (32) uncata 36(34) Embolus with sharp bend near the tip (Fig. 229); northwestern Mexico and coastal California (Map 13) _. (29) ventura = Embolus with gentle bend further from thestipa(Hics 32430 2517) mass ears wees 37 37(36) Sclerotized portion of embolus about as long as the longer flagellum (Fig. 243); Yucatan Peninsula (Map 13) ———— tee ata Rot Die 2 oa Salen Kom SB (31) celestun = Sclerotized portion of embolus shorter than the longer flagellum (Fig. 257); northwestern Mexico and California (Miers i) ea eee ae Rea (33) crassipes 38(32) Embolus thick and tapering to a point (Figs. SATA OAT S)) |peecmeretias Wit S MeN ESE hy pee ROE 39 = Embolus thin and needlelike (Figs. 286, DAC HN a heee tee eee en Moen nee Se ea Oe, 40 39(38) Longer flagellum greater than half the length of the cymbium (Fig. 271); west coastal Mexico (Map 10) _.... (35) petatlan = Longer flagellum less than half the length of the cymbium (Fig. 278); northwest- ern Mexico and Yucatan Peninsula (Map ells) ies: sere pe Stee (36) minima 40(38) Flagellae set off on long, thin stalk (Fig. 293); Cayman Islands, Jamaica, and laity (Miappilis)) a seee aes (38) jamaicensis = Flagellae set off on short, thicker stalk (Fig. 286); Honduras to Costa Rica (Miay sli) gcse Be. A ewes ote 8 (37) pacifica Metepeira foxi Group The M. foxi species group (sensu Levi, 1977) and M. desenderi share very similar genitalia, especially among males. Conse- quently, I am expanding the M. foxi group to include M. desenderi, but with the ex- clusion of the black sternum as a diagnos- tic character. The M. foxi group (sensu lato) includes males with an embolus that lacks a distal apophysis. The embolus curls almost 180° clockwise around a reduced or fleshy terminal apophysis (Figs. 7, 15, 22, 29). In other species groups the embolus is straighter and does not curl (e.g., Fig. 60), and the terminal apophysis is large, overhanging the embolus, and often scler- otized with teeth (e.g., Fig. 121). Epigynal 20 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 features uniting females in the M. foxi spe- cies group are harder to discern. Generally speaking, the epigynum has a_ stubby, shorter, often triangular scape and weaker posterior lobes that permit a wider view into the epigynal openings (compare Figs. OF 7. 2473 witha Figs. 143, 252), 1. Metepeira datona Chamberlin and Ivie Figures 7-13, 312; Map 1 Metepeira datona Chamberlin and Ivie, 1942: 68, fig. 196, 2. Female holotype from Daytona Beach, Florida, USA, in the AMNH, examined. Levi, 1977: 208-210, figs. [S=8 OO Rn Brignoli, 1983: Dp Ds Metepeira inerma Bryant, 1945: 378. Female holo- type from Cap Haitien, Haiti in the MCZ, exam- ined. First synonymized by Levi, 1977: 208-210. Description. Female from Morant Point, Mammee Bay, Jamaica. Carapace light aroma eyes with short lateral poste- rior extensions, sometimes with thin lon- gitudinal white line (Fig. 12). Legs ringed. Femur I with row of two macrosetae on anterior side; three light setae on anter- oventral side. Anterior half of dorsal foli- um white, posterior half black, margined by white; two halves separated by trans- verse white line. Abdomen widest in cen- ter (Fig. 12). Venter with a longitudinal white line. Pair of white spots on either side of spiracle (Fig. 13). Sternum usually entirely black, though sometimes with thin anterior and posterior marks, suggestive of median white line (Fig. 13). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.1. Anterior median eyes separat- ed by 1.4 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 1.6 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 4.2 mm. Carapace 1.7 mm long, 1.4 wide. First femur 2 mm, patella an tibia 2.4, metatarsus 1.5, tarsus 0.8. Second patella and tibia 1.9 mm, third 1.1, fourth 1.6. Male from Morant Point, Mammee Bay, Jamaica. Carapace yellowish brown, light around eyes. Median white mark anterior to thoracic furrow (Fig. 10). Legs same color as carapace, lightly ringed. Femur I with row of three macrosetae on anterior side, two on anteroventral side. Center of dorsum with transverse white line; poste- rior half darker than anterior half; thin me- dian black line; margin of folium white (Fig. 10). Venter, sternum as in female (Fig. 11). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separated by 1.8 diameters, posterior median eyes by 1.1, anterior me- dian eyes separated from anterior laterals by 1.5 diameters of anterior lateral eyes, lateral eyes separated by 0.5 their diame- ters. Total length 3 mm. Carapace 1.4 mm long, 1.1 wide. First femur 2.7 mm, patella and tibia 2.7, metatarsus 1.9, tarsus 0.8. Second patella and tibia 2.1 mm, third 1.0, fourth 1.3. Diagnosis. The dorsal folium and ab- dominal shape is distinctive in M. datona. Typically, the female’s abdomen is widest in the middle, forming a rhomboid in shape (Fig. 12), and both sexes have a transverse white line dividing the dorsal folium, with wide black markings on the posterior half (Figs. 10, 12). A ventral view of the epigynum in M. datona reveals a ridge under the scape that almost forms a straight line and which curves up at the ends (Fig. 9) instead of a V-shape (Figs. 17, 24, 31). The openings to the epigynum consist of small slits that flank a wide de- pression (Fig. 9), compared to larger openings that are relatively closer together (Figs. 17, 24, 31). The smooth embolus on M. datona is somewhat more compressed as it curls clockwise like the tip on a cork- screw, in contrast to the slightly ridged embolus supported on a stalk (Fig. 15) or the strongly ridged and tilted embolus (Figs. 22, 29). Like M. desenderi (Fig. 15) buat unlike M. grandiosa grandiosa (Fig. 22) and M. grandiosa alpina (Fig. 29), the posteroventral edge of the keel on the me- dian apophysis is rounded, as opposed to pointed (Fig. 7). Finally, M. datona has the thinnest and fleshiest terminal apophysis in the genus (Fig. 7). Variation. Average body length of 15 fe- males examined 3.8 mm, range 2.9 to 4.8 mm. Average body length of seven males examined 2.6 mm, range 1.8 to 3 mm. Natural History. Mature adults have been collected throughout the year (Fig. 312). According to locality labels, M. da- tona are found near the beach on cactus, palm, low scrub, mangrove, and bamboo. Females are known to wrap their egg sacs in dead leaves. Distribution. At sea level in Florida, Ba- hamas, British West Indies, and the Do- minican Republic (Levi, 1977, map 2; Map i) Records Examined. BAHAMAS Abaco Cays: Aba- co, 26°29'N, 77°5'W, 2.xii.1964 (W. B. Peck, MCZ); Allons Cay, 26°59'N, 77°40'W, 9.v.1953 (E. B. Hay- den, AMNH); Hopetown, Elbow Cay, 26°33'N, 76°57'W, 5.v.1953 (E. Hayden, AMNH); New Plym- outh, Green Turtle Cay, 26°50'N, 77°23'W, 7.v.1953 (G. Rabb, AMNH): Whale Cay, 926°43'N, 77°14'W, 121.1964 (W. B. Peck, CAS). Acklin’s Id.: Atwood’s Harbor, 22°13’N, 74°18'W, 15.ix.1958 (A. W. Scott Jr., MCZ): Salina Point, 22°13’N, 74°18’W, 15.viii.1958 (R. Robertson & A. W. Scott Jr, MCZ). Andros: Fresh Creek, 24°26'N, 77°57’W, 23.iv.1953 (L. Giovannoli, AMNH); Mangrove Cay, 24°15'N, 77°39'W, 26.iv.1953 (E. Hayden, AMNH); Nicolls Town, 25°8'N, 78°0'W, 14.iii.1967 (A. M. Nadler, AMNH). Berry Islands: Frazier’s Hog Cay, 25°24'N, 77°50'W, 29.iv.1953 (E. Hayden, AMNH); Little Harbor Cay, 25°34'N, 77°43’W, 1.v.1953 (Hayden & Giovannoli, AMNH). Crooked Island: North shore of Cripple Hill, 22°49'N, 74°16’W, 15.ix.1958 (A. W. Scott Jr, MCZ): NW end, Gordon (= Gun) Bluff, 22°50'N, 74°20'W, 15.viii.1958 (R. Robinson & A. W. Scott Jr, MCZ). Crooked Island Group: Long Cay, 22°37'N, 74°20'W, 7.iii.1953 (Hayden & Giovannoli, AMNH). East Plana Cay: 22°37'N, 73°33'W, 4.iii.1953 (E. Hayden, AMNH). Exumas: Musha Cay, 23°50'N, 76°15'W, 29.xii.1985 (A. Boutard, MCZ): Warderick Wells Cay, 24°22'N, 76°36'W, 9.i1.1953 (L. Giovannoli, AMNH), 11.i.1953 (Hayden & Giovannoli, AMNH). Grand Bahamas Island: Dundee Bay, 26°30'N, 79°15'W [?], 25.xii.1965 (L. Pinter, MCZ); near Fre port, pine-palmetto, 26°34'N, 78°27'W [P], 25.vii.1965 (L. Pinter, MCZ). Long Island: Clarence Town, 23°6'N, 74°59'W, 10.iii.1953 (L. Giovannoli, AMNH); Deadman’s Cay, 23°14’N, 75°14'W, 11.iii1.1953 (E. Hayden, AMNH). New Providence Is- land: 7 mi W. of Nassau, 25°5’N, 77°28’W, 4.1.1953 (Hayden & Giovannoli, AMNH). North Bimini: 25°44'N, 79°15'W, 25.i.1950 (C. M. Bogert, AMNB#), METEPEIRA ° Piel 21 6.vi. 1950 (M. Cazier & F. Rindge, AMNH), 15.v.1951 (W. J. Gertsch & M. Cazier, AMNH), 15.vi.1951 (M. Cazier, P. & C. Vaurie, AMNH), 15.vii.1951 (C. & P. Vaurie, AMNH), 13.xii.1952 (A. M. Nadler, AMNH), 28.xi.1959 (A. M. Nadler, AMNH). Rum Cay: near Port Nelson, 23°38'N, 74°50'W, 16.iii.1953 (Hayden & Giovannoli, AMNH). South Bimini: 25°42'N, 7T9°17'W, 12.vi.1950 (M. Cazier & F. Rindge, AMNH), 22.vi.1950 (M. Cazier & F. Rindge, AMNH), 15.v.1951 (W. J. Gertsch & M. Cazier, AMNHB), 15.vii.1951 (C. & P. Vaurie, AMNH), A viii.1951 (C. & P. Vaurie, AMNH), 4.xii.1952 (A. M. Nadler, AMNH), 25.iii.1953 (A. M. Nadler, AMNH); Gun Cay, 95°35'N, 79°20'W, 15.vi.1951 (AMNH). BRITISH WEST INDIES Caicos Islands: Long Cay, 21°28'N, 71°33'W, 10.ii.1953 (E. Hayden, AMNH);: South Caicos, from webs in upper beach zone, 21°31'N, 71°30'W, 3.iv.1973 (D. W. Buden, MCZ); West Caicos, 21°39'N, 72°28’W, 4.ii.1953 (Hayden & Giovannoli, AMNH), 5.ii.1953 (Hayden, Rabb, & Giovannoli, AMNH). Grand Cayman Island: 19°20'’N, 81°10’W, 15.ii.1960 (R. A. Lewin, MCZ). CUBA Oriente: Banes, 20°58'N, 75°43’W, 2.viii.1955 (A. F. Archer, AMNH); Ensenada Nispero, Ciudamar, 19°58’N, 75°52’W, 9.xi.1945 (P. Alayo, AMNH); Jur- agua, 19°56’N, 75°40'W, 1.x.1955 (P. Alayo, AMNH); Santa Fé [P?], 20°22'N, 75°53’W (A. F. Archer, AMNH). DOMINICAN REPUBLIC Barahona: Pla- ya Los Patos, 17°58'N, 7I°1LL'W, 31L.viii.1976 (J. A. Ottenwalder, MNSD). La Altagracia: Punta Cana, Isla Saona, 18°8’N, 68°34'W (Felix E. Del Monte & K. Guerrero, MNSD). HAITI Dept. du Nord: Cap- Haitien, 19°46’N, 72°13’W, 15.iii.1934 (E. Bryant, Utawana Exp., MCZ). JAMAICA Portland: between Boston and Blue Hole, 18°6’N, 76°37'W, 29.vii.1955 (A. F. Archer, AMNH). Saint Andrews: Hope Gar- dens, Gordontown, 18°2'N, 76°45’W, 27.vii.1955 (A. F. Archer, AMNH). Surrey: Morant Point, Mammee Bay, 17°53'N, 76°19'W, 14.x.1957 (A. M. Chickering, MCZ): Palisadoes, 17°56’N, 76°46’W, 11.xi.1967 (A. M. Chickering, MCZ); Palisadoes Area, 17°56'N, 76°46’W, 1.xi.1957 (A. M. Chickering, MCZ). U.S. VIRGIN ISLANDS Saint Thomas: Morant Point, 17°55'N, 76°10'W, 25.vii.1985 (G. B. Edwards, FSCA). 2. Metepeira desenderi Baert Figures 14-21, 307; Map 1 Epeira labyrinthea:—Banks, 1902: 60. Banks, 1924: 97. Misidentification. Metepeira sp.:—Roth and Craig, 1970: 116. Metepeira desenderi Baert, 1987: 145, figs. 16-21, 6, 2. Male holotype from Isla Pinzon, Galapagos, Ec- uador, in the IRSNB. Platnick, 1989: 341. Note. Holotype not examined because the figures in Baert (1987) are clear and because this is the only Metepeira species found on the Galapagos. Description. Female from east slope of Isla Santa Cruz, Galapagos Islands, Ecua- bo bo dor. Carapace yellowish brown, white around eyes, lighter median line (Fig. 20). Legs yellowish white, slightly darker an- nulations on distal ends of articles. Femur I with row of four macrosetae on anterior side: two to four on anteroventral side. Dorsal fleur-de-lis pattern broken into four white patches with anterior pair often larg- er than in most species. Posterior pair straighter than usual, forming a cross in center of folium (Fig. 20). Venter olive- brown with median white line surrounded by white U-shaped marking (Fig. 21). Pair of white spots on either side of spiracle. Sternum brownish black with wide, white line widening anteriorly (Fig. 21). Ratio of eye diameters: posterior medians and an- terior medians 1.0, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 2.3 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 5.5 mm. Carapace 2.3 mm long, 1.7 wide. First femur 2.7 mm, patella and tibia 3, metatarsus 2.2, tarsus 0.8. Second patella and tibia 2.4 mm, third 1.4, fourth 2. Male from same locality as female. Car- apace yellowish brown with wide white median mark (Fig. 18). Legs ringed like female. Femur I with row of four macro- setae on anterior side; four to seven on an- teroventral side. Dorsum and venter as in female, though median white line on ster- num sometimes broken (Figs. 18, 19). Ra- tio of eye diameters: posterior medians and anterior medians 0.9, anterior laterals 1.2, posterior laterals 1.1. Anterior median eyes separated by 2.2 diameters, posterior median eyes by 1.4, anterior median eyes separated from anterior laterals by 2.3 di- ameters of anterior lateral eyes, lateral eyes separated by 0.5 their diameters. To- tal length 3.2 mm. Carapace 1.6 mm long, 1.3 wide. First femur 2.6 mm, patella and tibia 2.6, metatarsus 2.3, tarsus 0.8. Sec- ond patella and tibia 2.2 mm, third 1.0, fourth 1.6. Diagnosis. Within the M. foxi species Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 8 © Females eX) @ Males ° Length of spider (mm) 2 -80 = -40 0 40 80 120 160 200 Collection day (since Jan 1) Figure 14. The length of mature M. desenderi collected on specific days of the year over a period of 84 years. There is a trend from smaller spiders early in the season to large spiders later in the season. Scale of abscissa: —80 = October 12 of the previous year; 40 = February 9; 160 = June 9. Symbols: Females (©), stippled regression line (fF = 40%); males (@), solid regression line (f° = 42%). group, the female abdomen shape of M. desenderi is closest to M. datona, though not quite as rhomboid (compare Fig. 20 with Figs. 12, 27, 34). The scape on M. desenderi is narrower and less triangular than other M. foxi species, and the de- pression under the scape forms a distinct U-shaped smile that is not seen in the oth- ers (compare Fig. 17 with Figs. 9, 24, 31). Like M. datona (Fig. 7) but unlike M. grandiosa grandiosa (Fig. 22) or M. gran- diosa alpina (Fig. 29), the posteroventral edge of the keel on the median apophysis is rounded as opposed to pointed (Fig. 15). Similar to M. datona (Fig. 7), the embolus of M. desenderi is curved like a slightly flattened corkscrew. Unlike M. datona, the embolus seems to be supported by a ped- icel off a large basal embolic apophysis (Fig. 15). In contrast, a pedicel is not ev- ident in M. datona, and the basal embolic apophysis is barely visible (Fig. 7). Variation. Average body length of 45 fe- males examined 5.8 mm, range 3.8 to 8.5 mm. Average body length of 13 males ex- amined 4.9 mm, range 2.9 to 6.5 mm. Natural History. Notes on the collection labels of Y. Lubin and R. Silberglied in- dicate that M. desenderi is active at night and that the retreat is composed of Opun- tia bark and croton leaves. According to Baert (1987), M. desenderi is found in large numbers in arid ecological zones on all islands. Although some mature speci- mens have been collected in November and August, most specimens are found be- tween January and June (Fig. 307), cor- responding to the “warm and wet” season in the Galapagos (van der Werff, 1978). The size of mature M. desenderi speci- mens appears to correlate with the collec- tion date: female spiders taken in August are, on average, 80% larger than those tak- en in November (Fig. 14). This trend may indicate that M. desenderi can vary the number of instars before maturity. Alter- natively, the intermolt growth may be greater for spiders with their antepenulti- mate or penultimate instars occurring dur- ing the warmer and wetter season. Distribution. Endemic to the Galapagos Islands (Map 1). Records Examined. ECUADOR Galapagos: Albe- marle, Tagus Cove, 0°16’S 91°22’W, 23.1.1899 (AMNH), 8.ii1.1899 (AMNH), 21.iii.1899 (AMNH), 23.iii.1899 (AMNH); Archipiélago de Galapagos, 0°0'N, 90°30'W (Williams Exped., 1923, MCZ); Ba- hia Conway, Indefatigable Island, 0°33'S, 90°32'W, 17.iii.1935 (Exline-Peck, CAS); Barrington Island, 0°49'S, 90°4’W, 1.viii.1929 (H. H. Cleaves, CAS); Campion, nr. Floreana (Santa Maria), 1°15'S, 90°27'W, 1.vi.1981 (Y. Lubin, MCZ): Charles Island, 1°17'S, 90°26'W, 10.v.1899 (AMNH): Indefatigable Island, 0°38'S, 90°23'W, 27.iv.1899 (AMNH), 18.vi.1929 (Pinchot South Sea Exp, USNM); Isla Abingdon, 0°35'N, 90°44'W, 25.vi.1899 (AMNH); Isla Albemarle, 0°30'S, 91°4’W, 13.i1.1899 (AMNH), 20.11.1899 (AMNH), 20.iii.1899 (AMNH): Isla Bin- dloe, O0°19'N, 90°29’W, 20.vi.1899 (AMNH): Isla Hood, 1°23’S, 89°39’'W, 18.v.1899 (AMNH), 26.v.1899 (AMNH); Isla Pinta, S Coast, 0°35’N, 90°44'W, 25.v.1964 (D. Q. Cavagnaro, CAS); Isla Pla- za, 0°35'S, 90°9’W, 7.iii.1970 (R. Silberglied, MCZ), 26.xi1.1973 (Y. Lubin, MCZ); Isla Santa Cruz, Acade- my Bay, 0°44'30"S, 90°17'30"W, 13.iii.1970 (R. Sil- berglied, MCZ); Isla Santa Cruz, E slope, 0°38’S, 90°23'W, 16.iv.1964 (D. Q. Cavagnaro, CAS); Isla Santa Cruz, Estacion Cientifica Charles Darwin, 0°44'S, 90°18’W, 24.1.1964 (D. Q. Cavagnaro & R. O. Schuster, CAS), 12.ii.1964 (Cavagnero & Schuster, CAS), 3.xi.1973 (Y. Lubin, MCZ), 24.xi.1973 (Y. Lu- bin, MCZ), 25.xi.1973 (Y. Lubin, MCZ); Isla Santa Fé, S coast, 0°50'S, 90°4’W, 30.i1.1983 (Y. Lubin, MCZ); James, 0°16'S, 90°42’W, 22.iv.1899 (AMNH); METEPEIRA ° Piel 23 Narborough Island, 0°25'S, 91°30'W, 28. iii. 1899 (AMNH); Sombrero Chino, Rocas Bainbridge, SE of Santiago, 0°21'S, 90°34’W, 31.iii.1970 (R. Silberglied, MCZ); Tower Island, Darwin’s Bay [?], 0°17’N, 89°59’W (AMNH): Tower Islands, 0°20’N, 89°58’W, 7.iv.1925 (N.Y. Zoological Society, AMNH); W coast of Albemarle Island, 0°11'’N, 91°21'’W, 9.iii.1935 (AMNBH). 3. Metepeira grandiosa grandiosa Chamberlin & lvie Figures 22-28, 321; Map 2 Metepeira grandiosa Chamberlin and Ivie, 1941: 17, figs. 24-26, 2. Female holotype from Ben Lo- mond, California, USA in the AMNH, examined. Metepeira palomara Chamberlin and Ivie, 1942: 72, figs. 200-204, 2, ¢. Female holotype and para- types from Mt. Palomar, California, in the AMNH, examined. First synonymized by Levi, 1977: 214. Metepeira grandiosa grandiosa:—Levi, 1977: 214, figs. 112-116, 2, 6. Note. Levi (1977) opted to collapse M. palustris, M. grandiosa, M. alpina, M. dakota, and M. palo- mara into three subspecies of M. grandiosa, with the caveat that more data may show the three sub- species to be distinct species. D. Buckle (personal communication) claims that his own recent obser- vations suggest that M. grandiosa alpina and M. grandiosa palustris are separate species. However, since the bulk of M. grandiosa specimens are out- side of the geographic range of this revision, and in the absence of molecular data, I will follow Levi's (1977) recommendation and leave these as separate subspecies. Description. Female from Los Angeles, California, USA. Carapace light around eyes with median white line extending to thoracic furrow (Fig. 27). Legs ringed. Fe- mur I with row of three to four macrosetae on anterior; one on anteroventral. Dorsum with usual folium, though lighter and more speckled than in most species (Fig. 27). Venter with a wide longitudinal white line. Pair of white spots on either side of spi- racle (Fig. 28). Sternum entirely black (Fig. 28). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.2, posterior laterals 1. Anterior median eyes separated by 1.9 diameters, posterior median eyes by 1.3, anterior me- dian eyes separated from anterior laterals by 3 diameters of anterior lateral eyes, lat- eral eyes separated by 0.4 their diameters. Total length 6.5 mm. Carapace 3.2 mm 24 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 long, 2.3 wide. First femur 3.5 mm, patella and. tibia o.7.) metatarsuseo 2.) tarsus lle Second patella and tibia 3.2 mm, third 1.9, fourth 2.8. Male from Parque Nacional Sierra San Pedro Martir, Baja California Norte, Mex- ico. Carapace as in female. Legs lightly ringed. Macrosetae on femur I variable— usually row of four macrosetae on anterior side, five on anteroventral side. Dorsum as in female (Fig. 25). Venter and sternum as in female (Fig. 26). Ratio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 1.5, posterior laterals 1.5. Anterior median eyes separated by 1.7 diameters, posterior median eyes by 0.8, anterior median eyes separated from an- terior laterals by 2.1 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 3.6 mm. Car- apace 1.9 mm long, 1.5 wide. First femur 2.9 mm, patella and tibia 2.8, metatarsus 2.5, tarsus 0.9. Second patella and tibia 2.3 mm, third 1.3, fourth 1.9. Diagnosis. The epigynal scape of M. grandiosa grandiosa is shorter and stub- bier than in other members of the M. foxi species group (compare Fig. 24 with Figs. 9, 17, 31). Unlike M. datona and M. de- senderi (Figs. 7, 15), the corkscrew em- bolus is tilted up with a heavy ridge (Fig. 22), but unlike M. grandiosa alpina (Fig. 29), it is more graceful and not as sharply bent. The posteroventral edge of the keel on the median apophysis in M. grandiosa grandiosa is not as pointed (Fig. 22) as in M. grandiosa alpina (Fig. 29) but not curved as in M. datona and M. desenderi (Bigss, (ela): Variation. Body length of females varies from 5.4 to 8.5 mm; males from 3.5 to 5.1 mm (Levi, 1977). Natural History. Mature specimens have been collected from March to Sep- tember (Fig. 321; Levi, 1977) in yellow pine forests and on Eriogonum fascicula- tum bushes. Elevations range from sea lev- el to 2,000 m. Distribution. Coastal mountainous re- gions from British Columbia to Baja Cal- ifornia Norte (Levi, 1977, map 2; Map 2). Records Examined. MEXICO Baja California Nor- te: Parque Nacional Sierra San Pedro Martir, 30°45'N, 115°13’W, 1.vii.1977 (C. E. Griswold, CAS). USA Arizona: Sycamore Canyon [?] Santa Cruz Co, 31°28'N, 110°42'W, 9.ix.1978 (G. F. Knowlton, MCZ). California: Los Angeles, 34°3’N, 118°15'W (Davidson, MCZ): Winchester, Double Butte, 33°42'N, 117°5'W, 20.iv.1974 (W. Icenogle, MCZ). 4. Metepeira grandiosa alpina Chamberlin and lvie Figures 29-35, 325; Map 2 Metepeira dakota Chamberlin and Ivie, 1942: 73, figs. 205-207, 2, 5. Male holotype and female paratype from Noonan, North Dakota, USA, in the AMNH, examined. Name synonymized by Levi, 1977: 212— 214. Metepeira alpina Chamberlin and Ivie, 1942: 74. Fe- male holotype and female paratypes from Fish Lake, Utah, USA, in the AMNH, examined. Metepeira grandiosa alpina:—Levi, 1977: 212-214, figs. 99, 100, 106-111, 2, 3. Brignoli, 1983: 276. Note. As first revisor, Levi (1977) chose to use the name M. alpina because its type specimen was collected closer to the center of the subspecies dis- tribution. Description. Female from Charcas, San Luis Potosi, Mexico. Carapace light around eyes with lateral posterior exten- sions and median white line extending to thoracic furrow (Fig. 34). Legs lightly ringed. Femur I with row of four macro- setae on anterior side; one on anteroven- tral side. Dorsum with usual folium, though lighter than in most species (Fig. 34). Venter with a wide longitudinal white line. Pair of white spots on either side of spiracle (Fig. 35). Sternum entirely black (Fig. 35). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.1, posterior laterals 1.1. Anterior median eyes separated by 1.7 diameters, posterior median eyes by 0.7, anterior me- dian eyes separated from anterior laterals by 2.8 diameters of anterior lateral eyes, lateral eyes separated by 1.1 their diame- ters. Total length 6.7 mm. Carapace 3 mm long, 2.4 wide. First femur 3.5 mm, patella and tibia 3.8, metatarsus 3.4, tarsus 1.2. METEPEIBA ° Piel 25 grandiosa alpina (4) Figures 15-21. Metepeira desenderi Baert (sp. 2; 0°38’S, 90°23’W). 15, male palpus, mesal. 16, epigynum, posterior. 17, epigynum, ventral. 18, male, dorsal. 19, male, ventral. 20, female, dorsal. 21, female, ventral. Figures 22-28. Metepeira grandiosa grandiosa Chamberlin and Ivie (sp. 3 [22, 25, 26] 30°45’N, 115°13’W; [23, 24, 27, 28] 34°3’N, 118°15’W). 22, male palpus, mesal. 23, epigynum, posterior. 24, epigynum, ventral. 25, male, dorsal. 26, male, ventral. 27, female, dorsal. 28, female, ventral. ; Figures 29-35. Metepeira grandiosa alpina Chamberlin and lvie (sp. 4 [29,32,33] 25°56’N, 105°22’W; [30,31 ,34,35] 23°8’N, 101°7’W). 29, male palpus, mesal. 30, epigynum, posterior. 31, epigynum, ventral. 32, male, dorsal. 33, male, ventral. 34, female, dorsal. 35, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 26 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 LYON ° vigilax = x rectangula & ° cajabamaba j= se 1000 > x zal 3 2 So} — 100 a -45 -35 -25 -15 -5 5 15 25 Latitude (Degrees North) Figure 36. The elevation of collection localities for M. vigilax, M. rectangula, and M. cajabamba at their corresponding lati- tudes. Species-specific altitudes appear to decrease with dis- tance from the equator. Elevations estimated from NOAA da- tabase of 5- by 5-minute geographic tiles. Regression line of M. vigilax does not include data points north of the equator. Symbols: M. vigilax (©), M. rectangula [%], M. cajabamba [@]}. Second patella and tibia 3.3 mm, third 1.9, fourth 2.9. Male from Santa Maria del Oro, Duran- go, Mexico. Carapace yellowish brown, light around eyes. Median white triangular mark anterior to thoracic furrow (Fig. 32). Legs same color as carapace. Macrosetae on femur I variable—usually row of four macrosetae on anterior side, five to six on anteroventral side. Dorsum, venter, and sternum as in female (Figs. 32, 33). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.4, posterior laterals 1.2. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 0.8, anterior median eyes separated from anterior laterals by 2.3 di- ameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. To- tal length 5.2 mm. Carapace 2.5 mm long, 2 wide. First femur 4.5 mm, patella and tibia 4.6, metatarsus 4.5, tarsus 1.3. Sec- ond patella and tibia 3.6 mm, third 1.8, fourth 2.8. Diagnosis. The epigynal scape of M. grandiosa alpina has the widest base of all members of the M. foxi species group, and the lobes surrounding the central depres- sion are somewhat fatter (compare Fig. 31 with Figs. 9, 17, 24,). Unlike M. datona and M. desenderi (Figs. 7, 15), the cork- screw embolus is tilted up with a heavy ridge (Fig. 29), but unlike M. grandiosa grandiosa (Fig. 22), it is thicker and with a sharper L-shape bend instead of a more curved L-shape. The posteroventral edge of the keel on the median apophysis in M. grandiosa alpina is pointier (Fig. 29) than in M. grandiosa grandiosa (Fig. 22) and not curved as in M. datona and M. desen- deri (Figs. 7, 15). Variation. Body length of females varies from 4.0 to 6.8 mm; males from 3.1 to 5.3 mm (Levi, 1977). Natural History. According to Levi (1977), mature specimens have been col- lected from June to August in meadows of bunchgrass, browsed aspen, oak, juniper, and sagebrush (Fig. 325). Elevations are at around 2,000 m. Distribution. North American Rockies from southern Alberta and Saskatchewan to Central Mexico (Levi, 1977, map 2; Map 2). Records Examined. MEXICO Durango: Santa Ma- ria del Oro, 25°56'’N, 105°22'’W, 28.vii.1947 (W. je Gertsch, AMNH). San Luis Potost: Charcas, moun- tain side, 23°8’N, 101°7’W, 7.vii.1934 (MCZ). USA. Colorado: Cimarron, 38°27'N, 107°33’W, 21.vii.1959 (H. W. & L. Levi, MCZ); Hayden Creek, Sangre de Cristo Mtns., 38°25'N, 105°35’W [P], 11.vii.1961 (H. W. & L. Levi, MCZ). Utah: SE shore of Bear Lake, 41°59'N, 111°20'W, 3.vii.1978 (G. F. Knowlton, MCZ). Metepeira vigilax Group Spiders in the M. vigilax group (Mete- peira cajabamba, Metepeira glomerabilis, Metepeira vigilax, Metepeira rectangula) are characterized by large, sclerotized epi- gynal openings, not unlike a snail’s shell (Figs. 40, 48, 55, 62), and long, robust, emboli with large scooplike basal embolic apophyses (Figs. 38, 46, 53, 60). 5. Metepeira cajabamba new species Figures 36, 38-45, 315; Map 3 Holotype. Male from Cajabamba, Cajamarca, Peru, 25.ix.1955, W Weyrauch, in CAS. The specific name is a noun in apposition after the locality. Description. Female paratype from Ca- jabamba, Cajamarca, Peru. Carapace dark brown, light around eyes with lateral pos- terior extensions (Fig. 43). Proximal two- thirds of femora white, remainder dark brown. Remaining articles lightly annulat- ed. Femur I with row of three macrosetae on anterior side; none on anteroventral side. Dorsal folium dark; white fleur-de-lis pattern reduced with thin branches (Fig. 43). Venter of abdomen black with wide white median line. Pair of small white spots on either side of colulus (Fig. 44). Sometimes the posterior end of median line ends in anchor shape (Fig. 45). Ster- num black, sometimes with (Fig. 44) or without (Fig. 45) median white line. Ratio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 1.4, posterior laterals 1.2. Anterior median eyes separated by 1.6 diameters posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 3 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 6.4 mm. Carapace 3 mm long, 2.3 wide. First femur 3.3 mm, patella and tibia 3.5, metatarsus 2.9, tarsus 0.9. Sec- ond patella and tibia 2.9 mm, third 1.7, fourth 2.7. Male holotype. Carapace dark brown, white around eyes with lateral posterior extensions and median white mark (Fig. 41). Proximal third of femora white, re- mainder dark brown. Remaining articles lightly annulated. Femur I with row of three macrosetae on anterior side; one on anteroventral side. Dorsal folium dark; white fleur-de-lis pattern indistinct with thin and broken branches (Fig. 41). Venter of abdomen black with median white mark. Pair of small white spots on either side of spiracle and colulus (Fig. 42). Ster- num black, often with median white line (Fig. 42). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 0.9, anterior me- dian eyes separated from anterior laterals by 2.3 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diame- ~ METEPEIRA ® Piel 2 ters. Total length 4 mm. Carapace 1.9 mm long, 1.5 wide. First femur 3 mm, patella and tibia 3, metatarsus 2.7, tarsus 0.9. Sec- ond patella and tibia 2.4 mm, third 1.2, fourth 1.8. Diagnosis. Female M. cajabamba and M. glomerabilis differ from the other spe- cies in the M. vigilax group (M. vigilax and M. rectangula) by the smaller and more tubelike openings to the epigynum (com- pare Figs. 40, 48 with Figs. 55, 62). Fe- male M. cajabamba differs Som M. glom- erabilis by having epigynal openings that are more oval (Fig. 40) than round (Fig. 48) when viewed ventrally, having the epi- gynal openings farther apart (compare Fig. 40 with Fig. 48), and by having the scler- otized tubelike openings more anteriorly directed (Fig. 40) than parallel to the epi- gynal groove (Fig. 48). Male M. cajabamba and M. glomerabilis differ from other spe- cies in the M. vigilax group by the smaller, thinner, and more graceful emboli (com- pare Figs. 38, 46 with Figs. 53, 60). Me- tepeira cajabamba differs from M. glom- erabilis by lacking a keel on the median apophysis, an only slightly fatter embolus, and having a normal embolus cap, in con- trast to a winged embolus cap (Fig. 46). A larger portion of the prosoma is dark in M. cajabamba (Fig. 43) as compared to M. glomerabilis (Fig. 51). Variation. Average body length of nine females examined 6.2 mm, range 5.4 to 7.5 mm. Average body length of four males examined 4 mm, range 3.7 to 4.9 mm. Natural History. With the exception of two specimens, mature M. cajabamba specimens have been collected in May through October (Fig. 315). Altitudes of collection localities appear to correlate steeply with latitude (Fig. 36). Median el- evation, about 500 m, with a range from near sea level to 3,500 m. Distribution. Ecuador and Peru (Map Sho Records Examined. PERU Ancash: Callejon de Huaylas, 9°10’S, 77°45'W, 15.viii.1988 (V. and B. Roth, CAS). Cajamarca: Cajabamba, 12.427, 7°37’S, 78°3’W, 25.ix.1955 (W. Weyrauch, CAS). La Libertad: 28 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Cerro Campana, La Cumbre, 8°1’S, 79°5’W, 10.x.1966 (A. F. Archer, AMNH). Lima: 20 km E An- cén, 11°47'S, 76°59'W, 1.xi.1953 (W. Weyrauch, CAS); 23 mi N Pativilea, 10°42’S, 77°47'W, 15.i.1955 (E. I. Schlinger and E. S. Ross, CAS); 5 km NW Chilca, 12°29'S, 76°48'W, 12.ix.1954 (E. I. Schlinger and E. S. Ross, CAS): between mouths of Rio Chillon and Ancon [?], 11°54'S, 77°7'W, 5.viii.1953 (M. Koepcke and Koepcke, MUSM); Cajacay [?], Rio Fortaleza, 10°40’'S, 77°52’'W, 6.iii.1956 (W. Weyrauch, CAS); Canta, Rio Chill6n, 11°28'12”S, 76°37'23’"W, 12.v.1951 (W. Weyrauch, CAS); Cerro Caracoles, 12°23'S, 76°45'W, 15.ix.1951 (W. Weyrauch, CAS): Lima, 12°3'S, 77°3’W (K. Jelski and Stolzman, PAN); Lomas de Iguanil [?] (Huaral), 11°29'51"S, 77°12'12’W, 14.vi.1986 (D. Silva, MUSM). 6. Metepeira glomerabilis (Keyserling) Figures 37, 46-52, 314; Map 3 Epeira glomerabilis Keyserling, 1892: 154, fig. 113, ?, 6. Male and female syntypes from Taquara, Rio Grande do Sul and Serra Vermelha, Rio de Janeiro, Brazil, in BMNH, examined. Male here designated lectotype. Araneus glomerabilis:—Petrunkevitch, 1911: 294. Roewer, 1942: 843. Aranea santa Chamberlin, 1916: 254, pl. 19, fig. 10, ?. Female holotype from Santa Ana, 3,000 m, Cuz- co, Peru, in MCZ, examined. NEW SYNONYMY. Metepeira santa:—Chamberlin and Ivie, 1942: 67. Platnick, 1993: 449. Metepeira glomerabilis:—Chamberlin and Ivie, 1942: 68, fig. 181. Metazygia glomerabilis:—Levi, 1991: 179. Platnick, 1993: 448. Erroneous transfer. Description. Female from Punapi, Tun- gurahua, Ecuador. Carapace brown; white arrowhead mark between eye region and thoracic furrow (Fig. 51). Legs dusky brown, lighter on ventral surfaces of fem- ora I and II; lighter on dorsal surfaces of patellae, tibiae, and metatarsi I and II. Fe- mur I with row of two to three macrosetae on anterior side; none on anteroventral side. Dorsal folium dark, speckled white, and white fleur-de-lis pattern with thin branches (Fig. 51). Venter black with short, wide, white median line ending in a T-shape; pair of small white spots on either side of spiracle (Fig. 52). Sternum black with median white line widening anteriorly (Fig. 52). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separated by 1.3 diameters, posterior median eyes by 0.9, anterior me- dian eyes separated from anterior laterals by 1.7 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their diame- ters. Total length 5 mm. Carapace 2.4 mm long, 1.7 wide. First femur 2.2 mm, patella and tibia 2.3, metatarsus 1.9, tarsus 0.9. Second patella and tibia 1.9 mm, third 1.2, fourth 1.8. Male from Pufapi, Tungurahua, Ecua- dor. Carapace as in female, except for a greater separation between white colora- tion around eyes and arrowhead mark (Fig. 49). Legs light tan, dark distally on femora. Femur I with row of three macro- setae on anterior side; four on anteroven- tral side. Dorsal folium and venter as in female (Figs. 49, 50). Sternum black with partly broken median white line widening anteriorly (Fig. 50). Ratio of eye diame- ters: posterior medians and anterior me- dians 1.1, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separat- ed by 1.8 diameters, posterior median eyes by 1.2, anterior median eyes separated from anterior laterals by 2 diameters of an- terior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 3.5 mm. Carapace 1.7 mm long, 1.4 wide. First fe- mur 2.5 mm, patella and tibia 2.4, meta- tarsus 2.3, tarsus 0.9. Second patella and tibia 1.9 mm, third 1.0, fourth 1.7. Diagnosis. Female M. glomerabilis and M. cajabamba differ from the other spe- cies in the M. vigilax group (M. vigilax and M. rectangula) by the smaller and more tubelike openings to the epigynum (com- pare Figs. 40, 48 with Figs. 55, 62). Me- tepeira glomerabilis differs from M. caja- bamba by having epigynal openings that are more round (Fig. 48) than oval (Fig. 40) when viewed ventrally, having the epi- gynal openings closer together (compare Fig. 48 with Fig. 40), and by having the sclerotized tubelike openings more parallel to the epigynal groove (Fig. 48) than an- teriorly directed (Fig. 40). Male M. glom- erabilis and M. cajabamba differ from oth- er species in the M. vigilax group by the smaller, thinner, and more graceful emboli (compare Figs. 38, 46 with Figs. 53, 60). Metepeira glomerabilis differs from M. ca- jabamba by having a keel on the median apophysis, an only slightly slimmer embo- lus, and a larger, winged embolus cap (compare Fig. 46 with Fig. 38). A larger portion of the prosoma is white in M. glomerabilis (Fig. 51), compared to M. ca- jabamba (Fig. 43). In addition, the mar- gins of the folium, particularly in the male, are whiter than in other species in the group (compare Fig. 49 and Keyserling [ 1892, fig. 113b] with Figs. 41, 56, 63). Variation. Average body length of 28 fe- males examined 5.1 mm, range 3.8 to 7.3 mm. Average body length of 10 males ex- amined 3.1 mm, range 2.4 to 4.3 mm. The base of the embolus varies from relatively thin (Fig. 46) to somewhat thicker, as in M. cajabamba (Fig. 38). Natural History. At first it would appear that this species is not seasonal—mature specimens have been collected throughout the year (Fig. 315). However on closer in- spection, there seems to be a seasonal shift with elevation: mature spiders are found at low altitudes (0-500 m) between March and October, at medium altitudes (500— 1,500 m) between August and March, and at high altitudes (1,500-4,000 m) between December and June (Fig. 37). In coastal regions, R. Baptista (personal communi- cation) reports that this species forms small aggregations of two to 10 spiders. Distribution. Colombia, Ecuador, Peru, Bolivia, Paraguay, and southern Brazil (Map 3). Elevations range from sea level to 4,000 m. Records Examined. BOLIVIA Chuquisaca: Mon- teagudo, 19°49'S, 63°59'W, 24.xii.1984 (L. E. Pefia, AMNH). BRAZIL Espirito Santo: Fazenda Santa Maria [?], Apiaca, 21°4'S, 41°25’W, 22.ix.1985 (R. L. C. Baptista, MZSP). Mato Grosso: Chavantina, 14°40'S, 52°21'W, 15.vi.1947 (J. C. Carvalho, MNBR)J): Fazenda Cervo, Trés Lagoas, 20°48’S, 51°43'W, 18.ix.1964 (Exp. Depto. Zool., MZSP); Utiariti, 13°2'S, 58°1L7'W, 15.viii.1961 (Lenks, MZSP). Minas Gerais: Lavras, 21°14'S, 45°0'’W, 20.x.1978 (W. Don Fronk, MCZ); Pedra Azul, 16°1'S, 41°16’W, 15.xii.1970 (F. M. Oliveira, AMNH); Peti Forest Res. [P] Santa Barbara, on bushes in cerrado, 19°56'S, 43°24'W, 28.vili.1986 (R. L. C. Baptista, MZSP). Pa- METEPEIRA ° Piel 29 @ Peru, Ecuador, Colombia ¢ Brazil, Paraguay, Bolivia Estimated Elevation (m) tN =} 6 6 50 100 150 200 250 300 350 400 450 500 550 Collection Day (since Jan 1) Figure 37. The elevation of collection localities for mature spi- ders of M. glomerabilis on specific days of the year between 1939 and 1990. There appears to be a shift in seasonal mat- uration times that corresponds better with elevation than with latitude. Elevations estimated from NOAA database of 5- by 5-minute geographic tiles. Scale of abscissa: 100 = April 9; 300 = October 27; 500 = May 15 of the following year. Symbols: Peru, Ecuador, Colombia [Ml]; Brazil, Paraguay, Bo- liva [@]. rand: Praia do Leste, Paranagua, 25°46'S, 48°31'W, 4.v.1967 (P. Biasi, MZSP). Rio de Janeiro: Guaratiba, 22°58'S, 42°48'W, 28. viii. 1976 (J. A. P. Dutra, MZSP); Ilha de Santana, Macae, 22°25'S, 41°44’W, 18.x.1986 (R. L. C. Baptista, MZSP); Itaipu, Niteroi, 22°56’S, 43°5'W, 20.iv.1985 (R. L. C. Baptista, MZSP). Rio Grande do Norte: Fazenda Canaan [?], Macaiba, 5°51'S, 35°21'W, 15.ix.1951 (M. Alvorenga, MZSP). Rondo6nia: Vila Rondonia, 10°52'S, 61°57'W, 9.ii.1961 (Pereira & Machado, MZSP). Santa Catarina: Nova Teutonia, 27°3'S, 52°24’W, 12.v.1949 (SMF). Sdo Paulo: Barueri, 23°31'S, 46°53'W, 13.iii.1966 (K. Len- ko, MZSP); Campos do Jordao, 22°44'S, 45°35’W, 15.iii.1945 (Wygod, MZSP); Instituto Oceanografico, Ubatuba, 23°26'S, 45°4'W, 15.v.1967 (P. Montouchat, MZSP); km 1 Rod, Rio Santos [?], Ubatuba, 23°26'S, 45°4'W, 12.x.1985 (R. L. C. Baptista, MZSP), 13.x.1985 (R. L. C. Baptista, MZSP); km 3 Rod, Rio Santos [?], Ubatuba, 23°26’S, 45°4’W, 6.1.1985 (R. L. C. Baptista, MZSP). COLOMBIA Cundinamarca: Sabana de Bogota, 4°43’N, 74°10'W, 10.xii.1990 (C. Valderrama, CV). ECUADOR Guayas: Guayaquil, 2°10'S, 79°54'W, 18.ii1.1942 (H. E. F & DL. E. FE CAS), 22.iii.1942 (Landis, CAS). Pichincha: 15 mi N Quito, 0°0'N, 78°30’W, 23.ii.1955 (E. I. Schlinger & E. S. Ross, CAS). Tungurahua: Bafios, 1°24'S, 78°25'W, 15.iv.1939 (W. C. Macintyre, MCZ); Pujiapi, 1°22'S, 78°28'W, 19.vi.1943 (D. L. F. & H. E. F CAS). PARAGUAY Alto Parand: Taguararaya [?], 25°30'S, 54°50’W (AMNH). Caazapa: Villa Pastoreo, 25°53’S, 55°45’W (D. Wees, MCZ). PERU Cajamar- ca: Cajamarca, 7°10’S, 78°31'W, 15.ii.1942 (W. Wey- rauch, AMNH). Lambayeque: 10 km S_ Chiclayo, 7°59'S, 77°17'W, 19.iii.1951 (EF. S. Ross & Michel- bacher, CAS). Piwra: Cerro Prieto, La Brea, 4°41'S, 81°6'W (CAS); Higueron (Las Lomas) [?], 4°19’S, 80°26’W, 29.vii.1941 (D. L. F & H. E. F CAS). VENEZUELA Sucre: 1 km S Villa Frontado, Rd. to 30 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Caripe, 10°27'N, 63°37'W, 12.ii.1984 (J. Coddington, USNM). 7. Metepeira vigilax (Keyserling) Figures 36, 53-59, 327; Map 4 Epeira vigilax Keyserling, 1893: 211, fig. 156, ¢. Male holotype from Taquara, Rio Grande do Sul, Brazil, in BMNH, examined. Araneus vigilax:—Petrunkevitch, 1911: 324. Roewer, 1942: 856. Metepeira dominicana Archer, 1965: 132, figs. 12, 18, ?. Female holotype from west of Bani, Dominican Republic, in AMNH. Holotype lost. Brignoli, 1983: 275. NEW SYNONYMY. Metepeira vigilax:—Levi, 1991: 180. Platnick, 1993: 449. Note. Although the type for M. dominicana is lost, the name has been identified by using Archer's (1965) description and illustration. Description. Female from Trujillo, west of Bani, Dominican Republic. Carapace light around eyes with lateral posterior ex- tensions (Fig. 58). Legs dark, light rings on proximal ends of leg articles. Femur I with row of four macrosetae on anterior side; three on anteroventral side. Dorsal folium darker than in most species; fleur-de-lis usually reduced to two white spots (Fig. 58). Venter brownish gray with lighter margins. Wide, short, median white line with pair of white spots on either side of spiracle (Fig. 59). Sternum brownish black with wide, white line widening anteriorly (Fig. 59). Ratio of eye diameters: posterior medians and anterior medians 1.2, anterior laterals 1.2, posterior laterals 1.1. Anterior median eyes separated by 1.8 diameters, posterior median eyes by 1.1, anterior me- dian eyes separated from anterior laterals by 3.5 diameters of anterior lateral eyes, lateral eyes separated by 0.1 their diame- ters. Total length 9.2 mm. Carapace 3.9 mm long, 3.2 wide. First femur 4.1 mm, patella and tibia 4, metatarsus 3.3, tarsus 1.2. Second patella and tibia 3.4 mm, third 223. tOUGt oe Male from same locality as female. Black carapace with white around eyes and extending posteriorly; white wedge mark in center (Fig. 56). Legs ringed like fe- male. Femur I with row of three macro- setae on anterior side; three on anterov- entral side. Dorsum and venter as in fe- male (Figs. 56, 57). Median white line may be limited to posterior end of sternum (Fig. 57). Ratio of eye diameters: posterior medians and anterior medians 0.9, anterior laterals 1.1, posterior laterals 1. Anterior median eyes separated by 1.6 diameters, posterior median eyes by 1.3, anterior me- dian eyes separated from anterior laterals by 2 diameters of anterior lateral eyes, lat- eral eyes separated by 0.2 their diameters. Total length 3.9 mm. Carapace 2 mm long, 1.5 wide. First femur 2.6 mm, patella and tibia 2.5, metatarsus 2.4, tarsus 0.9. Sec- ond patella and tibia 2.1 mm, third 1.2, fourth 1.7. Diagnosis. Female M. vigilax differ from those of other species in the M. vi- gilax species group by the shape of the epigynal openings: from a ventral view the openings are oval and angled inward pos- teriorly (Fig. 55); from a posterior view, the edges of the openings are more par- allel to the body (Fig. 61) as opposed to more perpendicular to the body (Figs. 39, 61). Male M. vigilax differ from other spe- cies because the embolus is larger and more robust (compare Fig. 53 with Figs. 38, 46, 60); the two flagella on the median apophysis are of more similar width (com- pare Fig. 53 with Fig. 38), and the keel on the median apophysis is slim and feather- shaped (Fig. 38), in contrast to arrow- shaped (Figs. 46, 60) or absent (Fig. 38). The dorsal folium differs from other Me- tepeira species by having a wide black me- dian stripe at the posterior end of the ab- domen (Fig. 58). In Brazilian and Bolivian specimens this stripe often extends all the way to the black anterior shoulders of the dorsum, forming a wide T-shape mark. Variation. Specimens from Argentina tend to be more lightly pigmented than those from more northern localities. White markings on the eye region of Brazilian and Bolivian specimens surround only the lateral eyes, in contrast to those on His- paniolan specimens, which cover the en- tire eye region. Natural History. Mature adults have METEPEIRBA ° Piel 31 lye vigilax ju PS (7) Figures 38-45. Metepeira cajabamba new species (sp. 5 [38-44] 7°37'S, 78°3’W; [45] 9°10’S, 77°45’W). 38, male palpus, mesal. 39, epigynum, posterior. 40, epigynum, ventral. 41, male, dorsal. 42, male, ventral. 43, female, dorsal. 44, female, ventral. 45, female, ventral. Figures 46-52. Metepeira glomerabilis (Keyserling) (sp. 6; 1°22'S, 78°28’W). 46, male palpus, mesal. 47, epigynum, posterior. 48, epigynum, ventral. 49, male, dorsal. 50, male, ventral. 51, female, dorsal. 52, female, ventral. Figures 53-59. Metepeira vigilax (Keyserling) (sp. 7 [53-57] 18°27'N, 72°17’W; [58,59] 18°17’'N, 70°22’W). 53, male palpus, mesal. 54, epigynum, posterior. 55, epigynum, ventral. 56, male, dorsal. 57, male, ventral. 58, female, dorsal. 59, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 32 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 been collected throughout the year except January, February, and March (Fig. 327). Spiders have been found among electric wires four meters above ground. Locality elevations range from near sea level to 1,400 m and follow an ecological zone that decreases in elevation with distance from the equator (Fig. 36). Over equivalent lat- itudes, M. vigilax lives at less than one- tenth of the elevation of M. rectangula (Fig. 36). Distribution. Hispaniola, Bolivia, Brazil, and coastal Argentina (Map 4). The dis- junct distribution between Hispaniolan and South American populations may be due to human-assisted migration. Records Examined. ARGENTINA Buenos Aires: Celsya—Pereyra [?], 34°50’S, 58°6’W (MACN); Ze- laya, 34°21'S, 58°52'W (MACN). BOLIVIA La Paz: Apolo, 14°43'S, 68°31'W, 10.viii. 1989 (L. E. Pefia, AMNH). Santa Cruz: Estaci6n Robore, above creek, 18°20'S, 59°45’'W, 27.ix.1955 (Azambuya, CAS). BRAZIL Espirito Santo: Fazenda Santa Maria [?], Apiaca, 21°4'S, 41°25'W, 14.v.1988 (R. L. C. Baptista, MZSP). Rio Grande do Sul: Sao Leopoldo, 29°46'S, 51°9'’W, 14.vi.1964 (Celia Valle, MZSP). DOMINI- CAN REPUBLIC Azua: El] Puerto, Majagual and Peralta, 18°34'N, 70°47'W, 10.xi.1979 (E. Marcano, MNSD). Prov. Trujillo Valdez: W Bani, 18°17'N, 70°22'W, 8.viii.1958 (A. F. Archer & E. de Boyrie Moya, AMNH). HAITI Departement de L’Ouest: Kenscoff, 18°27'N, 72°17'W, 15.xii.1929 (J. C. Myers, AMNH), 17.iv.1935 (AMNH). 8. Metepeira rectangula (Nicolet) Figures 36, 60-66, 306; Map 4 Epeira rectangula Nicolet, 1849: 500, female holo- type from Valdivia, Chile, in MNHN. Metepeira labyrinthea:—Petrunkevitch, 1911: 298. Roewer, 1942: 868. Bonnet, 1957: 2821. Erroneous synonymy. Metepeira rectangulata:—Chamberlin and Ivie, 1942: 71. Unjustified emendation. Note. The name was identified using drawings of the holotype (H. W. Levi, personal illustrations). Description. Female from Angol, Mal- leco, Chile. Carapace reddish brown with long white setae behind lateral eyes. An- terior third of carapace white, median white line reaching thoracic furrow (Fig. 65). Proximal halves of femora white, re- mainder black with distal white marks on dorsal surfaces. Patellae mostly black, re- maining articles white with black marks at base of setae. Femur I with row of three to five macrosetae on anterior side; three to four on anteroventral side. Anterior margin of dorsal abdomen black; dorsal fo- lium yellowish with brown speckles, white fleur-de-lis pattern with wide branches (Fig. 65). Venter black with wide white median line, flanked by pair of thin white lines; pair of white spots on either side of spiracle connected by white line (Fig. 66). Sternum black with posterior white mark (Fig. 66). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.1, posterior laterals 1.1. Anterior median eyes separated by 1.7 diameters, posterior median eyes by 1.0, anterior me- dian eyes separated from anterior laterals by 3.3 diameters of anterior lateral eyes, lateral eyes separated by 0.1 their diame- ters. Total length 8.4 mm. Carapace 3.7 mm long, 2.9 wide. First femur 4.2 mm, patella and tibia 4.1, metatarsus 3.7, tarsus 1.2. Second patella and tibia 3.4 mm, third Plesfountiags ule Male from Angol, Malleco, Chile. Car- apace reddish brown, anterior third white, median white line extending to thoracic furrow (Fig. 63). Proximal halves of fem- ora, white, distal halves black. Patellae black, remaining articles white with black spots at base of setae. Femur I with row of six to eight macrosetae on anterior side; seven to 11 on anteroventral side. Dorsal abdomen white, marbled, and speckled brown (Fig. 63). Venter dark brown with wide white median mark, flanked by pair of thin white lines; pair of white spots on either side of spiracle connected by white line (Fig. 64). Sternum dark brown with partly broken median white line (Fig. 64). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.2, posterior laterals 1.1. Anterior median eyes separated by 1.7 diameters, posterior median eyes by 1.1, anterior median eyes separated from anterior laterals by 2.9 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 6.1 mm. Carapace 3.2 mm long, 2.5 wide. First femur 5.2 mm, patella and tibia 5, metatarsus 4.9, tarsus 1.2. Second patella and tibia 4.1 mm, third 2, fourth Bode Diagnosis. Female M. rectangula differ from the other species in the M. vigilax group (M. vigilax, M. glomerabilis, M. ca- jabamba) by. the shape of the epigynal openings: from a ventral view they are larger, wider, and more gaping than in the other species (compare Fig. 62 with F igs. 40, 48, 55). Males differ from other spe- cies in the group by having an embolus that is relatively larger than that of M. glomerabilis and M. cajabamba, yet small- er than that of M. vigilax (compare Fig. 60 with Figs. 38, 46, 53). Among all members of the M. vigilax group, the keel on the median apophysis of M. rectangula is the largest and most robust, and tiel dorsal fo- lium has the lightest coloration (Fig. 65). Variation. Average body length of 11 fe- males examined 8.6 mm, range 5.8 to 10 mm. Average body length of six males ex- amined 6 mm, range 4.1 to 7 mm. Speci- mens from two localities in western Ar- gentina resemble M. vigilax, and may be hybrids. Natural History. This species appears to follow a narrow ecological zone that de- creases in elevation with increasing south- ern latitude (Fig. 36). Median elevation, about 500 m. Mature specimens have been collected January through April (Fig. 306). Specimens from localities south of the 36th parallel tend to be found in Jan- uary and February, whereas those north of the 36th parallel tend to be found in March and April. Distribution. Chilean Andes between 31° and 38° south (Map 4). Records Examined. ARGENTINA Cordoba: Cala- muchita, 32°4'S, 64°33’W, 15.iii.1954 Gi M. Viana, MACN). Mendoza: Mendoza, 32°53'S, 68°49'W, 30.iii. 1965 (H. W. Levi, MCZ). CHILE Bro-Bio: 4 km E road to Pinto, 36°42'S, 71°53’W, 4.i1.1976 (B. Mo- reno, AMNH): Road to Pemuco, Cruce del Carmen, 36°56'S, 72°4’W, 10.1.1976 (G. Moreno, AMNH). Co- quimbo: Illapel: Salamanca: Fundo Tahuinco, 31°44'S, 71°5’W, 30.iv.1946 (R. Doneso, AMNH). Malleco: Angol, 37°48’S, 72°43'W (D. S. Bullock, ie) ee) METEPEIRA ° Piel CAS), 10.iii.1945 (E. A. Chapin, USNM). Maule: 10 km S Curico, 35°4'S, 71°14’W, 15.iii.1968 (L. E. Pena, MCZ); Cordillera de Parral [?], 36°9’S, 71°50'W, 25.11.1956 (L. E. Pena, IRSNB): Linares, 35°51'S, 71°36'W (L. E. Petia, IRSNB): Miraflores, Pedag. [?], 35°55’S, 71°39’W (Toro, AMNH). O’Higgins: Fundo Millahue, Cunaco, 34°36'S, 71°16'W, 30.iv.1961 (AMNH). Regién Metropolitana: Melipilla, 33°42’S, 71°13'W (L. E. Pefia, IRSNB). Valparatso: Casablanca, 33°19'S, 71°25'W, 15.ii1.1955 (Edwin Reed, AMNH). Metepeira labyrinthea Group Levi (1977) described the M. labyrin- thea group very broadly—it included spe- cies with a longitudinal white line down the sternum and a short keel on the me- dian apophysis. Here, this species group is much narrowed to include only three North American species: Metepeira laby- rinthea, Metepeira lacandon, and Metepei- ra spinipes. Males of these three species are unique among Metepeira by having a toothless, smooth keel on the median apophysis. In addition, their distal embolic apophysis rises away (anteriorly) from the embolus proper and projects forward (ven- trally) until it is almost even with the em- bolus tip (Figs. 67, 76). In contrast, other Metepeira species with distal embolic apophyses have the embolus tip extend far beyond the projection of the apophysis (e.g., Fig. 171). The female epigynum has a characteristic shape. The scape is thick and fleshy and the epigynal openings have membranes that make them look distinctly (Fig. 69) or indistinctly (Fig. 78) almond- shaped. The epigyna of the M. labyrinthea species group are easily confused with that of the closely related species, M. gosoga (Fig. 173). Although most differences be- tween M. lab yinthea group and M. gosoga are only obvious in the epigynum, it should be noted that the dark marks inside the epigynal openings of the former ap- pear to look cross-eyed, but this cannot be said for the latter. In this work, only the species collected at localities south of the U.S./Mexico boarder are treated. 34 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 9. Metepeira spinipes F. O. P.-Cambridge Figures 67-75, 335; Map 6 Metepeira spinipes F. O. P.-Cambridge, 1903: 459, figs. 9, 10, 6, 2. Male holotype from Mexico City, Mexico, in BMNH, examined. Roewer, 1942: 868. Epeira labyrinthea grinnelli Coolidge, 1910: 281, °. Holotype from Palo Alto, California, lost. Araneus spinipes:—Petrunkevitch, 1911: 317. Aranea labyrinthea grinnelli:—Moles, 1921: 42. Metepeira douglasi Chamberlin and Ivie, 1941: 18, figs. 21-23, @. Female holotype from Santa Ana, California, in AMNH, examined. Chamberlin and Ivie, 1942: 66, figs. 169-170. First synonymized with M. labyrinthea grinnelli by Levi, 1977: 198. Metepeira labyrinthea grinnelli:—Roewer, 1942: 868. Metepeira labyrinthea:—Bonnet, 1957: 2822. Erro- neous synonymy. Metepeira grinnelli:—Levi, 1977: 198, figs. 21-27, 6, 2. NEW SYNONYMY. Description. Female from Huitzilac, Morelos, Mexico. Brown carapace with an- terior portion darker reddish brown, white behind lateral eyes (Fig. 74). Legs yellow- ish, femora reddish brown distally, other articles dark brown distally. Femur I with row of four to five macrosetae on anterior side: two to seven on anteroventral side. Anterior shoulders of abdomen black. Dorsal folium with usual Metepeira pat- tern, though largest branches of white fleur-de-lis shape usually widened into large spots (Fig. 74). Venter of abdomen black with wide white median line (Fig. 75). Pair of small white spots on either side of spiracle. Sternum black with posterior white mark that in some cases extends an- teriorly to the labium (Figs. 73, 75). Ratio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 1.0, posterior laterals 0.9. Anterior median eyes separated by 1.9 diameters, posterior median eyes by 1.3, anterior median eyes separated from anterior laterals by 2.9 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 9.2 mm. Carapace 3.7 mm long, 2.9 wide. First femur 4.4 mm, patella and tibia 4.7, metatarsus 4:6, tarsus 1.3. Sec- ond patella and tibia 4.1 mm, third 2.4, fourth 3.6. Male from Huitzilac, Morelos, Mexico. Carapace, dorsum, venter, sternum as in female (Figs. 71, 72). Distal portions of leg articles reddish black, elsewhere yellowish. Femur I with row of four to six macrosetae on anterior side; five to nine on anterov- entral side. Ratio of eye diameters: poste- rior medians and anterior medians 1.1, an- terior laterals 1.2, posterior laterals 1.1. Anterior median eyes separated by 1.8 di- ameters, posterior median eyes by 1.1, an- terior median eyes separated from anterior laterals by 3.2 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their di- ameters. Total length 7.5 mm. Carapace 3.8 mm long, 2.8 wide. First femur 6.2 mm, patella and tibia 6.3, metatarsus 7.1, tarsus 1.8. Second patella and tibia 5.3 mm, third 2.5, fourth 4. Diagnosis. Inside each epigynal depres- sion of M. spinipes and M. lacandon is a membrane that forms a slanted, oval- shaped opening (Figs. 69, 78). Within each oval-shaped opening is a dark mark, which in M. spinipes takes up a small part of that opening, resulting in a cross-eyed appear- ance (Fig. 69). Also, in M. spinipes (Fig. 69), the edges of the epigynal depressions are less distinct than in M. lacandon (Fig. 78). The distal embolic apophysis of M. spinipes is arrow-shaped (Fig. 79), in con- trast to a wider shovel-shape in M. lacan- don (Fig. 79). Variation. Average body length of 54 fe- males examined 9.1 mm, range 5.5 to 12.4 mm. Average body length of 55 males ex- amined 6.6 mm, range 3 to 10.8 mm. Natural History. Mature specimens have been collected between August and early November (Fig. 335). Elevations range from near sea level in California to 2.600 m in central Mexico. Variation in sexual dimorphism appears to correlate with habitat and social structure (Piel, 1996). Webs are found in dry regions among mesquite, Opuntia, Agave (ma- guey), cultivated Yucca, and Cactus. Spi- ders live in medium to small social colo- nies, which vary in size in accordance with local habitat quality (Uetz, 1988a,b). This behavioral and ecological relationship is METEPEIRA ® Piel 35 lacandon (10) Figures 60-66. Metepeira rectangula (Nicolet) (sp. 8; 37°48’S, 72°43’W). 60, male palpus, mesal. 61, epigynum, posterior. 62, epigynum, ventral. 63, male, dorsal. 64, male, ventral. 65, female, dorsal. 66, female, ventral. Figures 67-75. Metepeira spinipes F. O. P.-Cambridge (sp. 9 [67—72,74,75] 19°0’29’N, 99°15'50"W; [73] 39°18’N, 123°48’W). 67, male palpus, mesal. 68, epigynum, posterior. 69, epigynum, ventral. 70, male embolic division, Mental: 71, male, dorsal. 72, male, ventral. 73, female, ventral. 74, female, dorsal. 75, female, ventral. Figures 76-83. Metepeira lacandon new species (sp. 10; 16°45'N, 92°38’W). 76, male palpus, ribeak 77, epigynum, posterior. 78, epigynum, ventral. 79, male embolic division, ventral. 80, male, dorsal. 81, male, ventral. 82, female, dorsal. 83, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 36 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 thought to occur as a result of the spiders pursuing a risk-sensitive foraging strategy (Uetz, 1996). Distribution. Oregon to central Mexico (Levi, 1977, map 1; Map 6). Records Examined. MEXICO Aguascalientes: Hwy 45, 5.3 mi N Aguascalientes, 21°57'N, 102°17’W, 7.ix.1967 (R. E. Leech, REL). Baja California Sur: Sierra Laguna, 17 air mi ENE Todos Santos, 93°34'N, 110°0'W, 15.xii.1979 (C. E. Griswold, CAS). Chihuahua: 22.4 mi S Mifiaca, 28°24'N, 107°26'W, 23.viii.1950 (R. Smith, AMNH). Durango: 10 mi E El Salto, 23°12'’N, 105°52'’W, 8.viii.1947 (W. le Gertsch, AMNH):; 11 km W Suchil, 23°35'N, 104°5'W, 5.ix.1984 (W. Ile Pulawski, CAS): 6 mi NE El Salto, 23°19’N, 105°50’W, 11.viii.1947 (W. iE Gertsch, AMNH); Las Puentes [?], 26°49'N, 106°2'W, 23.vii.1947 (W. J. Gertsch, AMNH): Oti- napa, Q4°11'N, 105°2'W, 12.viii.1947 (W. Je Gertsch, AMNH): Palos Colorados, 24°2'N, 104°54’W, 5.viii.1947 (W. J. Gertsch, AMNH); Providencia, 26°44'N, 105°56’W, 24.viii.1947 (A. M. Davis, AMNH); SW Durango, 923°59'17"N, 104°45'47°W, 22.x.1994 (W. H. Piel, MCZ). Guanajuato: 30 mi SE Leon, 6 mi SE Silao, 20°52’N, 101°21'W, 6.ix.1964 (Jean & Wilton Ivie, AMNH); 6.3 mi NW Leén, 91°13'N, 101°43’W, 6.ix.1967 (R. E. Leech, REL): between Moroleon & Cuitzeo, 20°5’36’N, 101°9’28"W, 20.x.1994 (W. H. Piel, MCZ): near San Miguel de Allende, 20°55'N, 100°45’W, 16.ix.1976 (C. E. Griswold, CAS); S San Miguel de Allende, 20°46'36"N, 100°47'28"W, 20.x.1994 (W. H. Piel, MCZ); San Miguel de Allende. Road SW _ town, 20°52'N, 100°56’'W, 25.x.1982 (George Uetz, MCZ). Hidalgo: 18 mi E Huichapan, off Hwy 45, 20°23’N, 99°22'W, 25.viii.1984 (W. D. Sissom, C. Myers & L. Born, MCZ); 4 mi N Tizayuca, 19°54'N, 98°59'W, 20.xi.1946 (E. S. Ross, CAS); 41 km N Zimapan, 20°54'N, 99°13’W, 10.viii.1991 (W. H. Piel & G. S. Bodner, MCZ); Apulco, 20°19’N, 98°20'W, 6.x.1947 (H. Wagner, AMNH); Ozumbilla, 20°9’N, 101°16’W, 2.x.1957 (R. Dreisbach, MCZ); Pachuca, 20°7’N, 98°44'W, 30.viii.1957 (R. Dreisbach, MCZ); Tenango de Doria, 20°19'N, 98°13’W, 5.x.1947 (H. Wagner, AMNH). Jalisco: 12 mi S Mazamitla, 19°47'N, 103°8’W, 5.xii.1948 (H. B. Leech, CAS); Charco Ondo, 30 km W Ojuelos, 21°47’N, 101°53’W, 25.ix.1945 (H. Wagner, AMNH). México: Ixtapan de la Sal, 18°50’N, 99°41'W, 24.viii.1946 (H. Wagner, AMNH): Nevado de Toluca, 19°18’N, 99°44’W, 8.iv.1979 (George Uetz, MCZ); San Juan Teotihu- acan, 19°41'N, 98°52'W, 4.xi.1939 (C. M. Bogert & H. E. Vokes, AMNH); Tenancingo, 18°58'N, 99°36'W, 6.ix.1946 (H. Wagner, AMNH), 1.x.1946 (H. Wagner, AMNH), 15.x.1946 (H. Wagner, AMNH); Tenango del Valle, 19°7'N, 99°33’W, 25.vili.1946 (H. Wagner, AMNH), 27.viii.1946 (H. Wagner, AMNH); Teotihuacdn, 19°41'N, 98°52'W, 31.viiil1959 (A. F. Archer, AMNH); Tepotzotlan, 19°43'N, 99°13’W, 26.x.1982 (George Uetz, MCZ), 21.11.1983 (George Uetz, MCZ), 5.x.1983 (George Uetz, MCZ): Toluca, 19°18'N, 99°44’W, 10.viii.1978 (George Uetz, MCZ), 1.viii.1986 (George Uetz, MCZ); Toluca, at bottom of mountain near Parque Cierra Morelos, 19°18’N, 99°44'W, 10.viii.1978 (George Uetz, MCZ); Toluca, E of town on Paseo Tollocan [P], 19°18’N, 99°42’W, 23.x.1982 (George Uetz, MCZ). México D. F.: 19°25'N, 99°10'W, 12.x.1940 (H. Wagner, AMNBH), 28.xii.1940 (R. H. Crandall, AMNH), 15.ix.1943 (H. & D. Frizzell, AMNH): Contreras, 19°18’N, 99°17'W, 4.xii.1944 (H. Wagner, AMNH), 15.ix.1965 (N. L. H. Krauss, AMNH); Delegacion Tlalpan, Colonia Santa Ursula Xitla, 19°16’0"N, 99°10'25’"W, 12.x.1994 (W. H. Piel, MCZ): Desierto de los Leones, 19°22'’N, 99°16’W, 15.ix.1941 (H. Wagner, AMNH): EI Xitle, 18°61'N, 99°17’W [P], 12.viii.1942 (C. Tellez, AMNH): Haci- enda Cérdoba, 19°26'N, 99°10’/W [?], 29.x.1944 (H. Wagner, AMNH); Ouieros, 18°62’N, 99°17’W [?], 5.vii.1943 (M. Cardenas, AMNH): Mexico City, 19°25'’N, 99°10’W, 1.xi.1941 (C. Velo, AMNH), 25.ix.1957 (R. Dreisbach, MCZ);: Mixcoac, 19°23’N, 99°11'W (AMNH), 13.x.1940 (A. F. Archer, AMNH); Mixenac, 19°25'N, 99°10'W, 13.x.1940 (H. Wagner, AMNH); Pedregales, 18°60'N, 99°17'W [?], 15.viii. 1909 (AMNH); Petregal [?], 18°60'N, 99°17'W, 1.xii.1943 (AMNH); Rancho Cérdoba, 19°27'N, 99°10'W, 29.x.1944 (H. Wagner, AMNH): Tlaplan, 19°17'N, 99°10'W, 7.viii.1991 (W. H. Piel & G. S. Bodner, MCZ). Michoacan: 25 mi W. La Barca nr. Lago de Chapala, 20°17’N, 102°34’W, 11.ix.1976 (C. E. Griswold & Jackson, CAS); between Patzcuaro & Uruapan, 19°29'19"N, 101°48'20’"W, 19.x.1994 (W. H. Piel, MCZ); Hills N of Patzcuaro, 19°45’N, 101°36'W, 24.viii.1959 (A. F. Archer, AMNH); Hwy 110, 4 mi W. Jiquilpan, 19°59’N, 102°47'W 2.viii.1967 (R. E. Leech, REL); Hwy 15, 9.5 mi W. Morelia, 19°42’N, 101°16’W, 18.viii.1967 (R. E. Leech, REL); Lake Chapala, NW of Cojumatlan, 20°10'N, 102°53’'W, 7.ix.1966 (Jean & Wilton Ivie, AMNH); Monte de Zacapu, 19°47’N, 101°50’W, 24 viii.1959 (A. F. Archer, AMNH). Morelos: Cuer- navaca, 18°55’N, 99°15’W (AMNH), 15.ix.1941 (H. Wagner, AMNH), 18.xi.1946 (M. G. Bradt, AMNH): Cuernavaca/Tepotzotlan, interchange between I-95 & 115, 18°55'N, 99°13/W, 7.viii.1978 (MCZ): Huitzilac, 19°2'2'"N, 99°16'13’"W, 13.x.1994 (W. H. Piel, MCZ): North of Cuernavaca, 18°58’11"N, 99°14'37’W, 1ll.x.1994 (W. H. Piel, MCZ): S of Huitzilac, 19°0'29"N, 99°15’'50’W, 16.x.1994 (W. H. Piel, MCZ). Puebla: 6 mi E Rio Frio, 19°20'N, 98°35’W, 22.viii.1964 (Jean & Wilton Ivie, AMNH); Puebla, 19°3’N, 98°12'’W, 21.x.1982 (George Uetz, MCZ). San Luis Potost: 3 km W. Pilares, 21°55'34’N, 100°48’6"W, 21.x.1994 (W. H. Piel, MCZ): Cuidad del Maiz, 22°24'N, 99°36’W, 25.viii.1954 (R. Dreisbach, MCZ). Sonora: 46 mi S Agua Prieta on Highway 10, 31°0'N, 109°16’W, 15.viii.1959 (B. A. Branson, AMNH): Hermosillo, 29°4'N, 110°55’W, 20.ix.1952 (B. Malkin & V. E. Thatcher, AMNH): Sierra Man- zanal, 30°50'N, 110°10’W, 14.ix.1976 (Roth & Schroepfer, MCZ). Tlaxcala: Huamantla, 19°19'N, 97°56'W [P], 15.vii.1981 (C. Gold, CAS). Veracruz: 15 mi W. Banderilla, 19°39'N, 97°8’W, 31.x.1973 (S. GC. Williams & C. L. Mullinex, CAS); 15 mi West of Jalapa, 19°32'’N, 97°9’W, 23.vi.1946 (A. M. & L. I. Davis, AMNH). Zacatecas: 13 mi N. Sombrerete, 23°44'32"N, 103°47'10"W, 22.x.1994 (W. H. Piel, MCZ): Canutillo, 24°47'N, 101°31'W, 14.viii.1947 (W. J. Gertsch, AMNH); S. Zacatecas, 22°45'7'"N, 102°29'37"W, 22.x.1994 (W. H. Piel, MCZ). USA. Ar- izona: Southwestern Research Station, Chiricahua Mtns., 31°35’N, 109°14'W, 20.viii.1976 (V. Roth, MCZ). California: 26 mi W. Santa Rosa on Hwy 116, 38°31'N, 123°4'W, 19.ix.1976 (M. E. Thompson, MCZ): Mendocino, 39°18’N, 123°48'W, 18.viii.1959 (W. J. Gertsch, MCZ); Monterey, 36°36'N, 121°54'W, l.ix.1949 (A. F. Archer, MCZ): Pacific Grove, 36°37'N, 121°56’W (R. V. Chamberlin, MCZ):; Palo Alto, 37°27'N, 122°9'W (Doane, MCZ); Salt marsh on N shore of San Pablo Bay, Vallejo, 38°8'N, 122°27'W (D. Spiller, MCZ). 10. Metepeira lacandon new species Figures 76-83, 332; Map 6 Holotype. Male from San Cristobal, Chiapas, Mexico. The specific name is a noun in apposition after the Indian people who live in Chiapas. Holotype de- posited in the AMNH. Description. Female paratype from San Cristobal, Chiapas, Mexico. Reddish cara- pace, slightly darker in anterior half, ligh- ter behind lateral eyes (Fig. 82). Leg ar- ticles yellowish, gradually turning reddish brown distally. Femur I with row of four or five macrosetae on anterior side; two to four setae on anteroventral side. Anterior shoulders of abdomen black. Branches of white fleur-de-lis shape in dorsal folium thinner than in most species (Fig. 82). Venter of abdomen black with wide white median line that extends about half the distance between epigynal groove and spinnerets (Fig. 83). Pair of very small white spots on either side of spiracle. Ster- num black, often with central white spot (Fig. 83). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separated by 1.7 diameters, posterior median eyes by 1.0, anterior me- dian eyes separated from anterior laterals by 4.5 diameters of anterior lateral eyes, lateral eyes separated by 0.1 their diame- METEPEIRBA ° Piel 37 ters. Total length 11.5 mm. Carapace 4.6 mm long, 3.6 wide. First femur 5.3 mm, patella and tibia 5.3, metatarsus 4.9, tarsus 1.6. Second patella and tibia 4.4 mm, third Oui tountin 4 Male holotype. Carapace, dorsum, ven- ter, sternum as in female (Figs. 80, 81). Distal halves of femora, tibia reddish brown, elsewhere yellowish. Patellae, metatarsi reddish. Femur I with row of four or five macrosetae on anterior side; six or seven on anteroventral side. Ratio of eye diameters: posterior medians and an- terior medians 1.1, anterior laterals 1.5, posterior laterals 1.3. Anterior median eyes separated by 1.7 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 3.3 di- ameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. To- tal length 6.8 mm. Carapace 3.5 mm long, 2.7 wide. First femur 5.3 mm, patella and tibia 3.9, metatarsus 5.6, tarsus 1.7. Sec- ond patella and tibia 4.4 mm, third 2.3, fourth 3.4. Diagnosis. Inside each epigynal depres- sion of M. lacandon and M. spinipes is a membrane that forms a slanted, oval- shaped opening (Figs. 69, 78). Within each oval-shaped opening is a dark mark, which in M. lacandon takes up only a large part of that opening, resulting in a less cross- eyed appearance (Fig. 78). Also, in M. la- candon (Fig. 78), the edges of the epigynal depressions are more distinct than in M. spinipes (Fig. 69). The distal embolic apophysis of M. lacandon is more shovel- shaped (Fig. 79) than the thinner, arrow- shaped one in M. spinipes (Fig. 70). Variation. Average body length of three females examined 8.5 mm, range 7.5 to 10 mm. Average body length of four males examined 5.8 mm, range 4.2 to 6.8 mm. Natural History. Mature specimens were collected between July and Septem- ber (Fig. 332) from oak—pine woodland. Elevations range from 1,700 to 2,300 m. Distribution. Mountainous regions of Chiapas, Mexico (Map 6). 38 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Records Examined. MEXICO Chiapas: 12 km NW Comitan, 16°23'N, 92°15’W, 30.viii.1976 (E. S. Ross, CAS): 4 mi SE San Cristobal, 16°42'N, 92°36’W, 23.viii. 1966 (Jean & Wilton Ivie, AMNH); 5 km W San Cristobal de Las Casas on HWY 190, 16°44'’N 92°41'W, 27.vii.1983 (W. Maddison & R. S. Anderson, MCZ): 5 mi W San Cristobal, 16°45'N, 92°41’'W, 24 viii.1966 (Jean & Wilton Ivie, AMNH); San Cris- tobal, 16°45’N, 92°38’'W, 13.ix.1947 (H. Wagner, AMNH); San Cristobal de las Casas, 16°45’'N, 92°38'W, 22.vii.1947 (C. 6 ME Goodnight, AMNH); Tenejapa, 16°49'N, 92°31'W, 22.vii.1950 (C J. & M. Goodnight, AMNH). Metepeira nigriventris Group There are five species in the Metepeira nigriventris group: Metepeira nigriventris, Metepeira tarapaca, Metepeira calamuchi- ta, Metepeira galatheae, and Metepeira karkii. These closely related species are of- ten hard to distinguish because their gen- italia are similar, yet highly variable within a species. This species group is easily rec- ognized by the distinctive shape of the scape and similarities in palp morphology. Typically the base of the scape originates anteriorly and projects ventrally before curving posteriorly. This projection creates an overhang and a noticeable gap between the scape and the genital openings (e.g., le i, eld, IMI, 702). The embolus is fick and Thee a large, prominent, distal apophysis that Pees under the terminal apophysis (Figs. 84, 92, 100, 110, 121). 11. Metepeira nigriventris (Taczanowski) Figures, 84-91, 310; Map 5 Epeira nigriventris Taczanowski, 1878: 151, fig. 6, °. Bovalen lectotype from Lake Junin, Peru, in PAN, type lost. Keyserling, 1893: 217, fig. 161, 2, d. Araneus nigriventris:—Chamberlin, ‘1916: 248. Bon- net, 1955: 550. Metepeira nigriventris:—Chamberlin and Ivie, 1942: TA, figs. 211-214, 2, 6. Platnick, 1993: 449. Note. Although the type is lost, the type locality and Taczanowski’s descriptions are sufficient to rec- ognize the species. Description. Female from 12 km west of Tarma, Junin, Peru. Carapace dark brown, light around eyes with lateral posterior ex- tensions (Fig. 90). Proximal halves of leg articles yellow, distal halves black. Femur I with row of four macrosetae on anterior side; five on anteroventral side. Dorsum darker and white fleur-de-lis pattern small- er than in most species (Fig. 90). Venter mostly black with reduced, short, thin, white median line (Fig. 91). Sternum en- tirely black. Ratio of eye diameters: pos- terior medians and anterior medians 1.0, anterior laterals 1.4, posterior laterals 1.4. Anterior median eyes separated De : 8 di- ameters, posterior median eyes by 1.2, an- terior iravetienn eyes separated from anterior laterals by 5 diameters of anterior lateral eyes, lateral eyes separated by 0.4 their di- ameters. Total length 11 mm. Carapace 5.1 mm long, 4.2 wide. First femur 5.5 mm, patella and tibia 5.8, metatarsus 5.5, tarsus 1.9. Second patella and tibia 5.3 mm, third 3.3, fourth 4.5. Male from same locality as female. An- terior margin of chelicerae with large, swollen tooth and several denticles. Cara- pace reddish brown with lighter eye re- gion, lateral posterior extensions, and long, thin median white line (Fig. 88). Femur I with row of about five macrosetae on an- terior side; nine on anteroventral side. Coloration of legs, dorsum, venter, and sternum as in female (Figs. 88, 89). Ratio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separated by 1.8 diameters, posterior median eyes by 1.2, anterior median eyes separated from anterior laterals by 3.7 di- ameters of anterior lateral eyes, lateral eyes separated by 0.4 their diameters. To- tal length 9 mm. Carapace 4 mm long, 3.2 wide. First femur 5.8 mm, patella and tibia 6.2, metatarsus 6.5, tarsus 1.7. Second pa- tella and tibia 4.5 mm, third 2.8, fourth 4. Diagnosis. Metepeira nigriventris is eas- ily distinguished from other species in the M. nigriventris group by its dark pigmen- tation. As its name implies, the sternum is black and the white ventral mark on the abdomen is reduced to a much shorter and thinner line (Figs. 88-91). While the ster- num of M. karkii is similarly dark, the dor- sal and ventral markings on the abdomen are much lighter (compare Figs. 88—91 METEPEIRA ° Piel 39 ae au °° nigriventris calamuchita (13) Figures 84-91. Metepeira nigriventris (Taczanowski) (sp. 11; 11°25'S, 75°48’W). 84, male palpus, mesal. 85, epigynum, pos- terior. 86, epigynum, ventral. 87, male embolic division, ventral. 88, male, dorsal. 89, male, ventral. 90, female, dorsal. 91, female, ventral. Figures 92-99. Metepeira tarapaca new species (sp. 12; 21°39’S, 69°33’W). 92, male palpus, mesal. 93, epigynum, posterior. 94, epigynum, ventral. 95, male embolic division, ventral. 96, male, dorsal. 97, male, ventral. 98, female, dorsal. 99, female, ventral. Figures 100-107. Metepeira calamuchita new species (sp. 13; 32°4'S, 64°33’W). 100, male palpus, mesal. 101, epigynum, posterior. 102, epigynum, ventral. 103, male embolic division, ventral. 104, male, dorsal. 105, male, ventral. 106, female, dorsal. 107, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 40 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 with Figs. 125-128). The epigynum of M. nigriventris has a wider membrane over the openings that is visible on either side of a very wide scape (Fig. 86). In contrast, this feature is hidden behind a thinner scape in its likely sister species, M. tara- paca (Fig. 94). The overall shape of the epigynum (Fig. 86) and the pair of small notches in the posterior lobes (Fig. 85) also make this species distinctive. The male palp of M. nigriventris has an em- bolus that is relatively slimmer and more graceful than those of other species in the M. nigriventris species group (compare Fig. 84 with Bigs O27 MOOI UO! IDI). The shape of the embolus and its distal apophysis differs from other species (com- pare Fig. 87 with Figs. 95, 103, 112, 124). Variation. Average body length of 25 fe- males examined 9.5 mm, range 7.2 to 11.5 mm. Average body length of 16 males ex- amined 7.4 mm, range 5.3 to 9.5 mm. Natural History. Spiders are commonly found around Lake Titicaca living in me- dium and large colonies among power lines, Bolivian pines, Cactus, Bacharis, rock outcroppings, and tall grasses (L. Rayor, personal communication, and vari- ous locality labels). Mature specimens have been collected throughout the year except September and October (Fig. 310). Median elevation, 3,900 m. Distribution. High altitude regions of southern Peru and western Bolivia (Map 5). Records Examined. ARGENTINA Jujuy: Puma- huasi, 22°17'S, 65°41'W, 8.xi.1970 (L. E. Pefia, MCZ). BOLIVIA La Paz: 45 mi S La Paz, 17°9'S, 67°36'W, 25.11.1951 (E. S. Ross & Michelbacher, CAS); 70 mi S La Paz, 17°30'S, 67°36’W, 25.ii1.1951 (E. S. Ross & Michelbacher, CAS): La Paz, Avenida Sport Club, 16°30’S, 68°9’W, 4.i1.1959 (A. M. Nadler, AMNH); La Paz, in garden of house, 16°30'S, 68°9'W, 15.iv.1959 (R. Walsh, AMNH); Lake Titicaca, Copacabana, Yampupata, & Isla del Sol, 16°10'S, 69°S’W, 17.v.1995 (L. Rayor, MCZ); near La Paz, 16°30'S, 68°9'W, 24.v.1958 (R. Walsh, AMNH): S end of Lake Titicaca, 100 km NW La Paz, 16°10'S, 69°5'W, 5.vii.1958 (R. Walsh, AMNH);: Tiahuanaco, Puma Puerto Ruins, 16°33'S, 68°42'W, 1.ii.1973 (Ann Moreton, MCZ). Oruro: 6 km N Challapata, 18°51’S, 66°47'W, 23.11.1951 (E. S. Ross & Michelbacher, CAS); Gorge Uhuschlucht, near Oruro, 17°59’S, 67°9'W, 7.ii.1954 (Forster & Schindler, ZSM). Potos?: Villazon, 22°6'S, 65°36'W, 30.xii.1984 (L. E. Pefia, AMNH). PERU Apurimac: Chincheros, 13°30'48’S, 73°42'47’"W, 12.xii.1980 (C. Gold, CAS); Puna near Abancay, 13°38'2"S, 72°52'52”/W, 15.xii.1947 (W. Wey- rauch, CAS). Ayacucho: Puquio, 14°42'S, 74°8'W, 15.iv.1950 (F. Blancas, MUSM): San Antonio (Pu- quio), 14°47'S, 74°7’W, 1.xi.1985 (D. Silva, MUSM). Cusco: Cheqquerec, 13°23'S, 72°8'W, 2.ix.1993 (J. Ochoa Camara, MCZ): Cusco, 13°31'6"S, 71°58’41"W, 8.viii.1965 (P. & B. Wygodzinsky, AMNH). Junin: 8 mi W Tarma, 11°25'S, 75°48’W, 6.11955 (E. I. Schlinger & E. S. Ross, CAS); Cochas Bajo, 11 km W Tarma, 11°25'21"S, 75°46'1L1”"W, 27.iii.1988 (J. Palmer & D. Smith, MCZ); Cochas Bajo, 11 km W Tarma, rock ledge in agricultural val- ley, 11°25'21"S, 75°46’11"W, 29.iii1.1988 (J. Palmer, MCZ); Huancayo, 12°4'S, 75°14'W, 15.vi.1947 (W. Weyrauch, AMNH); Oroya, 11°32'S, 75°54'W, 12.iv.1914 (M. P. Anderson, AMNH). Lima: Bosque de Zarate, 11°53'S, 76°27'W, 18.i.1981 (J. Francke, MUSM). Puno: 10 mi N Ayaviri, 14°45'S, 70°35’W, 1.iii1.1951 (E. S. Ross & Michelbacher, CAS); Cama- cane, 15°55'S, 69°50’W, 20.xi1.1955 (L. E. Pefia, IRSNB); Isla Taquili, Lago Titicaca, 15°46’S, 69°41'W, 23.xii.1980 (C. Gold, CAS); Juli (col. Chu- cuito), 16°13'S, 69°27'W, 7.xi.1952 (F. Blancas, MUSM); near Chucuito, Lago Titicaca, 15°50'S, 69°48'W, 10.iii.1953 (M. Koepcke, MUSM); Puna, Lake Titicaca, 15°50'S, 70°2'W, 15.vi.1947 (W. Wey- rauch, AMNH); Puno, 15°50’S, 70°2’'W (Soukup, AMNH); Yunguyo, downtown plaza, 16°15'S, 69°5’W, 31.41.1973 (Ann Moreton, MCZ). 12. Metepeira tarapaca new species Figures 92-99, 305; Map 7 Holotype. Male from Quillagua, Antofagasta, Chile, 4.ii.1965, L. E. Pefia, in MCZ. The specific name is a noun in apposition after a Chilean province where it is abundant. Description. Female paratype from Quillagua, Antofagasta, Chile. Light red- dish brown carapace, white in center and around eyes with lateral posterior exten- sions (Fig. 98). Legs yellowish white, ringed brown at distal ends of articles. Fe- mur I with row of four macrosetae on an- terior side; three on anteroventral side. Dorsal folium white with black speckles (Fig. 98). Venter of abdomen black with wide, white median line, sometimes flanked by thinner white lines that togeth- er form a U-shape posteriorly (Fig. 98). Sternum black with median white line, sometimes broken (Fig. 99). Ratio of eye Estimated elevation (m 0 20 40 60 80 100 Percent of median white line on sternum Figure 108. Elevation of collection localities of mature female M. tarapaca with differing amounts of white on the sternum. Spiders with a median white line covering 100% of the sternum length are found at a wide altitude range. Spiders with a me- dian white mark covering only a short length of the sternum are only found at high elevations. Elevations estimated from NOAA database of 5- by 5-minute geographic tiles. diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separat- ed by 1.4 diameters. posterior median eyes by 1.0, anterior median eyes separated from anterior laterals by 3.4 diameters of anterior lateral eyes, lateral eyes separated by 0.1 their diameters. Total length 7.8 mm. Carapace 3.2 mm long, 2.8 wide. First femur 4.3 mm, patella and tibia 4.4, metatarsus 4, tarsus 1.2. Second patella and tibia 3.8 mm, third 2.1, fourth 3.2. Male holotype. Light reddish brown carapace, lighter around eyes and white mark in center (Fig. 96). Legs yellowish white, gradually growing darker toward distal ends of articles. Femur I with row of four macrosetae on anterior side; five on anteroventral side. Dorsal folium, venter, and sternum as in female (Figs. 96, 97). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.2, posterior laterals 1.1. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 1.0, anterior median eyes separated from anterior laterals by 2.5 di- ameters of anterior lateral eyes, lateral eyes separated by 0.1 their diameters. To- tal length 4.4 mm. Carapace 2 mm long, 1.6 wide. First femur 3.3 mm, patella and tibia 3.5, metatarsus 3.3, tarsus 1. Second patella and tibia 2.8 mm, third 1.4, fourth Dye Diagnosis. Unlike M. nigriventris or M. METEPEIRBA ° Piel 4l karkii, M. tarapaca has a white mark on the sternum (Figs. 97, 99). The epigynum is close to that of M. nigriventris, except that it is less robust (compare Fig. 93 with Fig. 85), and the membrane just over the openings cannot be seen behind the thin- ner scape (compare Fig. 86 with Fig. 94). Unlike M. calamuchita, the epigynum of M. tarapaca does not widen posteriorly (compare Fig. 94 with Fig. 102); unlike M. karkii, the posterior epigynal lobes are not swollen (compare Fig. 94 with Fig. 123); unlike M. galatheae, the black sclerotized circles behind the epigynal openings are larger and not shifted posteriorly (compare Fig. 94 with Figs. 118-120). The segment of the embolus between the basal and dis- tal embolic apophysis is relatively thicker than in other members of the M. nigriven- tris species group (compare Fig. 92 with Figs. 84, 100, 109, 121). The terminal di- vision of the male palp in M. tarapaca shows a distal embolic apophysis that dif- fers in shape from that of other species (compare Fig. 95 with Figs. 87, 103, 117, 124). Variation. Average body length of 22 fe- males examined 7.4 mm, range 5 to 11.5 mm. Average body length of six males ex- amined 5.9 mm, range 4.4 to 7.3 mm. The dorsum and venter are often darker than the holotype. Many lack the flanking lines and the U-shape on the venter; some at higher elevation have a much reduced white line on the sternum. Natural History. Mature specimens have been collected throughout the year, especially between January and April (Fig. 305). V. Roth (vial label) notes that these spiders live in a social colony. M. Roy (per- sonal communication) reports that colo- nies can reach 200 individuals. Median el- evation, 2,800 m. Dicom Moderately high altitudes in northern Chile and southern Peru (Map 0: Records Examined. CHILE Antofagasta: 7 km S Toconao, 23°11'S, 68°1'W, 25.xii.1988 (V. & B. Roth, CAS); Aguas Blancas [?] (= 24°11’S 69°51’W), To- conao, 23°11'S, 68°L’W, 11.x.1955 (L. E. Pefia, 42, Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 IRSN); Antofagasta, 93°39'S, 70°24'W, 15.xi.1975 (L. E. Pena, AMNH): Quillagua, 21°39'S, 69°33'W, 431.1965 (L. E. Pefia, MCZ), 211.1973 (W. C. Sedg- wick, MCZ); Rio Loa, 25 km S Quillagua, 21°54’S, 69°33'’W, 20.viii.1966 (E. Schlinger & M. Irwin, CAS). Elloa: Thermo Puritania, 35 km N San Pedro de Atacama, 22°37'S, 68°13’W, 25.xii.1988 (V. & B. Roth, CAS). Tarapaca: Arica Timar, Alociado [?], 18°45'S, 69°42’W, 22.ii1.1973 (N. Hichins, AMNH); Ariquilda, 19°38'S, 69°32’W, 29.iv.1969 (L. E. Pena, MCZ): Canchones, 20°27'S, 69°37'W, 29.i1.1973 (W. C. Sedgwick, MCZ); Chapiquilta, 19°18’S, 69°25'W, 6.vi.1968 (L. E. Petia, MCZ): Chiapa, 19°32’S, 69°13’W, 24.iv.1969 (L. E. Pena, MCZ); Pica, 20°30'S, 69°21'W, 26.i1.1973 (W. C. Sedgwick, MCZ); Poroma, Quebrada de Tarapaca, 19°52’S, 69°11'W, 20.iv.1969 (L. E. Pefia, MCZ); Quisama, 19°19’S, 69°28'W, 5.vi.1968 (L. E. Pefia, MCZ). PERU Are- quipa: Arequipa, 16°23'56"S, 71°32’6"W, 15.vii.1955 (A. Meza, MZSP); Chiguata, near Arequipa, 16°24’S, 71°24'W, 15.ii.1948 (W. Weyrauch, MUSM). 13. Metepeira calamuchita new species Figures 100-107, 308; Map 5 Holotype. Male from Calamuchita, Cordoba, Argen- tina, 15.i1.1955, J. M. Viana, in MACN. The specific name is a noun in apposition after the locality. Description. Female paratype from Cal- amuchita, Cordoba, Argentina. Carapace reddish brown, light around eyes with lat- eral posterior extensions (Fig. 106). Leg articles annulated distally. Femur I with row of six or seven macrosetae on anterior side; five on anteroventral side. Anterior portion of dorsal folium lighter than in most species (Fig. 106). Venter black with wide white median line and pair of large white spots on either side of spiracle (Fig. 107). Sternum brownish black with wide, white median line (Fig. 107). Ratio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 1.4, posterior laterals 1.3. Anterior median eyes separat- ed by 1.6 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 3.2 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 8.9 mm. Carapace 3.4 mm long, 2.8 wide. First femur 4.5 mm, patella and tibia 4.7, metatarsus 4.7, tarsus 1.3. Second patella and tibia 4.1 mm, third 2.3, fourth 3.5. Male holotype. Carapace brown with light, triangular, median mark pointing posteriorly (Fig. 104). Leg articles white, darkening brown distally. Femur I with row of four macrosetae on anterior side; five or six on anteroventral side. Anterior dorsal folium mostly white, posterior most- ly black (Fig. 104). Venter as in female (Fig. 105). Broad white median mark on sternum (Fig. 105). Ratio of eye diame- ters: posterior medians and anterior me- dians 1.1, anterior laterals 1.4, posterior laterals 1.4. Anterior median eyes separat- ed by 1.4 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 2.2 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 4.4 mm. Carapace 2.3 mm long, 1.8 wide. First femur 3.8 mm, patella and tibia 3.7, metatarsus 4, tarsus 1.2. Second patella and tibia 2.9 mm, third 1.5, fourth 2.5. Diagnosis. Metepeira calamuchita dif- fers from others in the M. nigriventris spe- cies group by the large, scooped openings, and posteriorly widening epigynum (Fig. 102). The epigyna of M. tarapaca and M. calamuchita both have deep scooped-out openings, but these are angled posteriorly in the latter (Fig. 102) and laterally in the former (Fig. 94). The male palp of M. cal- amuchita is easily distinguished from that of other species by the widened (Fig. 100) and inwardly curved (Fig. 103) “dewlap” under the embolus. Variation. Average body length of 13 fe- males examined 7.7 mm, range 5 to 9.8 mm. Average body length of five males ex- amined 5.2 mm, range 4 to 6.4 mm. Two specimens collected near 2,700 m in Itur- be, Jujuy, Argentina, have entirely dark sterna. In contrast, most specimens are found at lower elevations with white me- dian lines on their sterna. Natural History. Mature specimens have been collected between October and July, although they are probably available throughout the year (Fig. 308). Most ele- vations range from 150 to 1,700 m, with one population at 2,700 m. Distribution. Northern Argentina, at mostly lower altitudes east of the Andes (Map. 5). Records Examined. ARGENTINA Buenos Aires: Las Espadufias [?], Sierra de la Ventana, 38°9’S, 61°48'W, 15.x.1973 (Maury & Cesari, MACN); Sierra de la Ventana, 38°9'S, 61°48'W, 15.vii.1972 (Amarril- la, MACN). Coérdoba: C. Paz, 31°24'S, 64°31'W, 15.v.1940 (C. Marti, MACN); Calamuchita, 32°4'S, 64°33’W, 15.xii.1941 (J. M. Viana, MACN), 15.i.1955 (J. M. Viana, MACN); Mina Clareo, 31°43’S, 65°0'W, 15.iv.1973 (Stiebel, MACN). Jujuy: Iturbe, 22°59’S, 65°21'W, 22.ii.1983 (L. E. Pefia, AMNH). La Rioja: Ilias [?], 29°5’S, 66°19’W (P. M. Gomez, MACN). San Luis: Chosmes and Desaguadero (Mendoza), 33°24'30"S, 67°0'0"W, 14.iv.1967 (L. E. Pefia, MCZ). Santiago del Estero: Santiago del Estero, 27°47’S, 64°16'W (AMNH), 3.iv.1965 (H. W. Levi, MCZ): 70 km W Santiago, 27°47'S, 65°25'W, 3.iv.1965 (H. W. Levi, MCZ); Quebrachos: Sumampa, Parayacu [?], 29°22'S, 63°28'W, 15.xi.1944 (Maldonado Bruzzone, MLBP). 14. Metepeira galatheae (Thorell) Figures 108-120, 304; Map 7 Epeira galatheae Thorell, 1891: 53. Female holotype from “Cobija, Bolivia,” now, Cobija, Antofagasta, Chile, in the UZMK, examined. Araneus galatheae:—Bonnet, 1955: 506. Metepeira galatheae:—Levi, 1991: 179. Platnick, 1993. Metepeira cereicola nomen nudum, female in AMNH but no description has been found. Female man- uscript type from Salamanca, Coquimbo, Chile, collected by Archer on 30.iv.1961. Note. Thorell (1891) listed the holotype’s locality as “Cobija, Bolivia.” While Cobija, Bolivia exists (11°02'S 68°44'W), it is an unlikely locality for the corvette Galathea to visit on its voyage to southern Asia because it would have meant climbing over the Andes. Instead, the port town of Cobija, Chile (22°33'S 70°16'W) is much more likely, especially since this region of Chile was under Bolivian ad- ministration throughout the period of Galathea’s voyage, 1845-1847 (Paynter et al., 1975). Description. Female from Chile Chico, Aisén province, Chile. Reddish brown car- apace with white setae, light around eyes, lateral posterior extensions, central arrow- shaped mark (Fig. 113). Legs light yellow, articles annulated distally. Femur I with row of four or five macrosetae on anterior side; two to four on anteroventral side. Dorsum with black and white setae. Foli- um speckled brown with a white fleur-de- lis that reduces posteriorly (Fig. 113). Ven- ter brownish gray with wide median white METEPEIRA ° Piel 43 line and pair of large white spots on either side of spiracle. A pair of very faint white stitching parallel to and on either side of median line (Fig. 114). Sternum reddish brown, sometimes with small posterior and anterior white marks (Fig. 114). Ratio of eye diameters: posterior medians and an- terior medians 1.0, anterior laterals 1.2, posterior laterals 1.2. Anterior median eyes separated by 1.7 diameters, posterior median eyes by 1.2, anterior median eyes separated from anterior laterals by 3.6 di- ameters of anterior lateral eyes, lateral eyes separated by 0.1 their diameters. To- tal length 7.8 mm. Carapace 3.5 mm long, 2.7 wide. First femur 3.7 mm, patella and tibia 3.9, metatarsus 3.5, tarsus 1.2. Sec- ond patella and tibia 3.4 mm, third 2.1, fourth 3. Male from Chile Chico, Aisén province, Chile. Leg coloration as in female, except femur I mostly reddish brown with row of four macrosetae on anterior side; five on anteroventral side. Carapace, dorsum, ven- ter and sternum as in female (Figs. 111, 112). Ratio of eye diameters: posterior me- dians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separated by 1.6 diameters, posterior median eyes by 1.2, anterior me- dian eyes separated from anterior laterals by 3.3 diameters of anterior lateral eyes, lateral eyes separated by 0.4 their diame- ters. Total length 5.3 mm. Carapace 2.7 mm long, 2.1 wide. First femur 3.9 mm, patella and tibia 3.8, metatarsus 3.6, tarsus 1.1. Second patella and tibia 3.2 mm, third 1.8, fourth 2.6. Diagnosis. Although the sternum of M. galatheae varies from solid brown (Fig. 114) to brown with median white marks, the venter of the abdomen is surprisingly consistent. The venter has a wide brownish gray area and a short median white line flanked by very faint indications of parallel lines (Fig. 114). In contrast, the flanking lines in M. karkii thicken anteriorly (Fig. 128) and the venter of M. nigriventris is almost completely black (Fig. 91). Despite enormous variation in the hood of the 44 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Collection Day (since Jan 1) Latitude (Degrees North) Figure 109. Days on which mature M. galatheae were col- lected between 1937 and 1989 with latitude of the collection locality. Seasonality appears to be more restricted in southern regions than in northern regions. Scale of abscissa: —150 = August 3; 1 = January 1; 150 = May 30. scape with a corresponding variation in the position of the epigynal openings, it is nonetheless possible to distinguish the fe- males of M. galatheae from those of other Species in the M. nigriventris group. Whether ventrally or posteriorly posi- tioned, the epigynal openings and the darkened shadows of the sclerotized re- ceptacles beneath them are relatively smaller than those of M. calamuchita, M. tarapaca, and M. nigriventris (compare Figs. 118-120 with Figs. 86, 94, 102). Also, M. galatheae lacks the swollen posterior lobes present in M. karkii (compare Figs. 118-120 with Fig. 123). The shape of the embolus of M. galatheae varies significant- ly (compare Fig. 109 with Fig. 110). How- ever, unlike other species, the embolus of M. galatheae has a distinct, round and swollen protrusion (Fig. 117). Variation. Average body length of 80 fe- males examined 7.9 mm, range 4.8 to 12.5 mm. Average body length of 16 males ex- amined 5.9 mm, range 3.5 to 8.1 mm. Epi- gyna vary considerably. Many, similar to the holotype, open ventrally and resemble a posteriorly widened version of M. tara- paca (Figs. 119, 120); cleared epigyna show relatively straight ducts connecting the epigynal openings with the seminal re- ceptacles. These have a short distance be- tween the openings and the hood of the scape. Others, usually in southern Chile, have posterior openings and look surpris- ingly different (Fig. 118); cleared epigyna show S-shaped ducts connecting the open- ings with the seminal receptacles. These have an extended wrinkled area between the openings and the hood of the scape. However, several females [e.g., CHILE Bto-Bio: Chillan, 8.xi.1976 (G. Moreno, AMNH); Las Lajuelas, 11.i.1976 (G. Mo- reno, AMNH)] have epigyna that appear to be intermediate between the two forms. Furthermore, no somatic features were found to be sufficiently different, and little corresponding variation was found among sympatric males. It is possible that further collecting efforts will discover correspond- ing males, and future molecular studies may show that speciation has, in fact, oc- curred. But in the meantime, I am opting to treat both varieties as belonging to the same species. Natural History. Although mature spec- imens have been collected throughout the year (Fig. 304), the seasonality of this spe- cies appears to depend on its latitude. At the 45th southern parallel, spiders are usu- ally found in late November and Decem- ber; at the 40th parallel, spiders occur be- tween October and February; and at the 30th parallel, they are collected year round (Fig. 108). Median elevation, 550 m. Spi- ders are found on Patagonian scrub, dunes, and wire fences. Distribution. Chile and Argentina (Map 1s Records Examined. ARGENTINA Buenos Aires: Felipe Sola, 38°1'S, 62°50’W, 15.i.1944 (Prosen, MLP); Patagones, 40°48'S, 62°59’W, 15.ii.1937 (J. M. Viana, MACN): Sierra de la Ventana, 38°9’S, 61°48'W, 15.iii.1939 (J. C. Gario, MACN). Catamar- ca: Mutquin, 28°19'S, 66°10’W, 15.i1.1963 (O. de Fer- rariis, AMNH). Chubut: 15 km S Epuyen, 42°22'S, 71°21'W, 15.i.1986 (P. A. Goloboff, N. I. Platnick, & R. T. Schuh, AMNH): 19.5 km E Shaman, 44°27'S, 70°30'W, 19.xi.1966 (E. I. Schlinger & M. E. Irwin, CAS); 3 km N Puerto Lobos, 41°59'S, 65°6'W, 14.xii.1966 (E. I. Schlinger & M. E. Irwin, CAS); 35 km E Esquel, 42°54'S, 70°53’W, 18.xi.1966 (E. I. Schlinger & M. E. Irwin, CAS); El Hoyo [?], 42°4’S, 71°30'W (A. Kovacs, AMNH), 10.i.1962 (Andor Ko- vacs, AMNH); Epuyen, 42°15'S, 71°23'W, 18.xi.1962 (Andor Kovacs, AMNH); Leleque, 42°28’S, 71°6’W, 12.11.1965 (Andor Kovacs, AMNH); Los Manantiales, METEPEIRA ° Piel 45 A 4 ee 2 BS galatheae (14) karkil (is) Figures 110-120. Metepeira galatheae (Thorell) (sp. 14 [110-114,116,117,120] 46°33’S, 71°57’W; [115,119] 29°50’S, 70°2’W; [118] 33°30’'S, 71°25’W). 110, male palpus, mesal. 111, male, dorsal. 112, male, ventral. 113, female, dorsal. 114, female, ventral. 115, epigynum, posterior. 116, epigynum, posterior. 117, male embolic division, ventral. 118-120, epigynum, ventral. Figures 121-128. Metepeira karkii (Tullgren) (sp. 15; 51°38’S, 69°13’W). 121, male palpus, mesal. 122, epigynum, posterior. 123, epigynum, ventral. 124, male, dorsal. 125, male, ventral. 126, female, dorsal. 127, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. N of Comodoro-Rivadavia, 45°28'S, 69°29’'W, 19.xi.1985 (L. E. Pefia, AMNH): N of Camarones, Cantera, Namuncura, 44°46'S, 65°42/W, 17.xi.1985 (L. E. Pefia, AMNH): Rio Turbio, 42°13'S, 71°41’W (Andor Kovacs, AMNH), 12.i.1962 (Andor Kovacs, AMNH). Cordoba: 12 mi W Sampacho, 33°23’S, 64°43'W, 7.ii.1951 (E. S. Ross & Michelbacher, CAS): Arguello, 31°21'S, 64°15'W, 15.xii.1943 (J. A. De Car- \ lo, MACN); Calamuchita, 32°4'S, 64°33’'W, 15.xii.1940 (J. M. Viana, MACN); Sampacho, 33°23’S, 64°43'W, 7.i1.1951 (E. S. Ross & Michelbacher, CAS). Mendoza: Between Beazley and San Rafael, 34°10'S, 67°29'W, 4.iii.1983 (L. E. Petia, AMNH); Mendoza, 32°53'S, 68°49'W, 30.iii.1965 (H. W. Levi, MCZ); Us- pallata, 32°35'S, 69°20'W, 7.iii1.1983 (L. E. Pefia, AMNH). Neuquén: Catan Lil, Charahuilla, 39°45’S, 46 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 70°37'W, 15.ii.1971 (O. de Ferrariis, AMNH):; Cuba del Leon [?], 39°9’S, 70°53’W, 15.1.1975 (Maury, MACN); Lago Alumine, 38°55’S, 71°9’W, 15.i.1976 (O. de Ferrariis, AMNH); Zapala, 38°54’S, 70°4’W, 15.i.1958 (J. R. Navas, MACN). Rio Negro: Cerro Alto [?], 41°8’S, 70°40’W (MACN): Co. Leones, source of Rio Limay, 40°33’S, 70°26’W, 28.ii.1959 (J. R. Navas, MACN); El Bolson, 41°58’S, 71°31'W, 1.ii.1961 (A. Kovacs, AMNH), 17.x.1961 (Andor Ko- vacs, AMNH); El Bolson, 41°58’S, 71°35’'W, 2.ii1.1965 (Andor Kovacs, AMNH): General Roca, 39°3'S, 67°32'W, 15.ix.1964 (Bachmann, MEG). Salta: Maury [P], 24°40'S, 65°45'W, 15.1.1975 (MACN). San Juan: 10 km N Matagusanos, 31°10’S, 68°38’W, 13.i.1983 (L. E. Petia, AMNH). San Luis: INTA Experimental Station, E of Villa Mercedes, 33°40'S, 65°27'W, 8.xii. 1967 (C. R. Ward, CAS). Santa Cruz: 2.4 km S$ Fitz Roy, 47°2'S, 67°15'W, 12.xii.1966 (E. I. Schlinger & M. E. Irwin, CAS). Tucumdn: San Miguel de Tu- cuman, IML gardens, 26°49'S, 65°13'W, 19.xii.1979 (L. A. Stange, FSCA). BRAZIL Mato Grosso: Campo Grande, 20°27'S, 54°37'W, 7.ii.1952 (M. Alvarenga, MZSP). CHILE Aconcagua: W end tunnel, 85 km S Hlapel, 32°49'S, 71°7'W, 29.xi.1950 (E. S. Ross & Michelbacher, CAS). Aisén: 8 km W Chile Chico, 46°33'S, 71°57'W, 22.xi.1966 (E. I. Schlinger & M. E. Irwin, CAS); Chile Chico, near lake, 46°33’S, 71°43'W, 21.xi.1966 (E. I. Schlinger & M. E. Irwin, CAS). Antofagasta: 6 km N Muelle de Piedra, N Tal- tal, 25°21'S, 70°30’W, 4.ii.1942 (Junius Bird, AMNH); Caleta Hueso Parado, Taltal, 25°22’S, 70°2S’W, 1.ii.1941 (Junius Bird, AMNH); Cobija, 22°33’S, 70°16'W (ZMUC); Quebrada Paposo, 25°2'S, 70°27'W, 3.ii.1989 (L. Stange, FSCA). Araucania: Pe- mehue [?], 38°3'S, 71°43’W (L. E. Pefia, IRSNB): Villarrica, 36°16'S, 72°13'W, 25.xi.1963 (G. F. Ed- munds, AMNH). Atacama: 50-60 km S Copiapo, 2751'S, 70°20'W, 24.viii.1966 (E. I. Schlinger & M. E. Irwin, CAS); Copiapo, 27°22'S, 70°20’W (Cartis, MNRJ); Rio Copiapo, by the sea, 27°19’S, 70°56’W, 13.vi.1968 (L. E. Pefia, MCZ). Bio-Bio: 4 km E road to Pinto, 36°42’S, 71°53’W, 4.1.1976 (B. Moreno, AMNH); Chillén, 36°36’S, 72°7’W, 2.i.1976 (G. Mo- reno, AMNH), 21.ii.1978 (G. Moren, MCZ); Chillan, in cemetery, 36°36’S, 72°7'W, 8.xi.1976 (G. Moreno, AMNH); Cuesta de Quilmo, Chillin, 36°38'S, 72°12'W, 13.xi.1976 (G. Moreno, AMNH); El Aban- ico, 37°20'S, 71°31’W, 30.xii.1950 (E. S. Ross & Michelbacher, CAS); Las Lajuelas, 36°39'S, 72°8’W, 111.1976 (G. Moreno, AMNH). Concepcion, 36°50'S, 73°3'W (L. E. Pefia, IRSNB); Concepcion: Salta de Rio Laja, 37°13’S, 72°23’W, 30.i.1951 (E. S. Ross & Michelbacher, CAS); Nuble: 50 km E San Carlos, 36°25'S, 71°6’W, 26.xii.1950 (E. S. Ross & Michelbacher, CAS); Nuble: Cordillera de Chillén [?], 36°51'S, 71°24’W, 1.ii.1947 (L. E. Pefia, IRSNB): Rio Andalien, 36°44'S, 73°1’W, 25.iii.1979 (S. Gu- tierrez, MCZ). Coquimbo: 20 mi E La Serena, 29°54'S, 70°56’W, 3.vii.1950 (E. S. Ross & Michel- bacher, CAS); 5 mi N Ovalle, 30°31’S, 71°12’'W, 1.xii.1950 (E. S. Ross & Michelbacher, CAS); Banos del Toro, 29°50'S, 70°2’W, 15.ii.1947 (L. E. Pefia, IRSNB); Cerro Talinay, 30°50'29"S, 71°37'14”"W, 29.xi.1961 (A. F. Archer, AMNH); Cuesta las Cardas, Ovalle Rd., 30°17'S, 71°16'W, 13.xi.1961 (R. Wagen- knecht, AMNH); Hacienda Illapel, 31°36’S, 71°7’W, 3.x1.1954 (L. E. Pefia, IRSNB), 19.x.1966 (E. I. Schlinger, M. E. Irwin, & L. E. Pefia, CAS); Illapel: Salamanca: Fundo Quelén, 31°52’S,— 70°52’W, 30.iv.1961 (A. F. Archer, AMNH): La Serena, 29°54'28"S, 71°15'15"W, 15.ii.1947 (L. E. Pefia, IRSNB); Loma de Pefiuelas, 6 km S La Serena, 29°57'S, 71°18'W, 28.xi.1961 (A. F. Archer, AMNH): Quilacén, 16 km E La Serena, 29°54’S, 71°5’W, 2.x.1961 (R. Wagenknecht, AMNH). Los Lagos: Purr- anque, 40°55'S, 73°10'W, 15.ii.1955 (Edwin Reed, AMNH): Rio Bueno, 40°19’S, 72°58’W (L. E. Pejia, IRSNB): Valdivia: Neltume, 39°48'S, 71°57’'W, 23.x1.1988 (V. & B. Roth, CAS). Malleco: Angol, 37°48'S, 72°43'W, 29.i1.1951 (E. S. Ross & Michel- bacher, CAS). Maule: Linares, 35°51'S, 71°36’W (L. E. Pena, IRSNB), 15.1.1947 (L. E. Pena, IRSNB): Miraflores, Pedag. [?], 35°55’S, 71°39'W (Toro, AMNH). O’Higgins: Chépica, 34°44'S, 71°17'W, 15.xii.1947 (L. E. Pefia, IRSNB). Regidn Metropoli- tana: 34 km W Santiago, 33°30'S, 71°25'W, 19.xii.1950 (E. S. Ross & Michelbacher, CAS); Ba- tuco, nr. Santiago, 33°13'S, 70°47’W (Guil. Mann, AMNH); Lampa, 33°17'S, 70°54’W, 1.v.1979 (L. E. Pefia, AMNH). Santiago: El Golf [?], 33°30'S, 71°25'W, 9.iv.1961 (A. F. Archer & J. Aros, AMNH); Santiago, 33°30'S, 71°25'W (L. E. Pefia, IRSNB), 1.ii.1973 (W. C. Sedgwick, MCZ). Talca: 22 mi N Tal- ca, 35°7'S, 71°40'W, 22.xii.1950 (CAS). Valparaiso: Concon, in cow farm, 32°55'S, 71°31'/W, 4. iii. 1962 (H. Morales, AMNH); La Cruz, 32°53’'S, 71°16'W, 18.i.1973 (W. C. Sedgwick, MCZ); Llay-Llay, 32°51’S, 70°58’W, 20.i1.1973 (W. C. Sedgwick, MCZ): Los Mai- tenes [P], 32°59’S, 71°15’W, 14.x.1954 (L. E. Pefial, IRSNB); Quintay, 33°11'S, 71°42'W, 19.ii.1967 (E. I. Schlinger, CAS); Valparaiso, 33°2'S, 71°38’W (Edwin Reed, AMNH). 15. Metepeira karkii (Tullgren) Figures 121-128, 303; Map 5 Araneus karkii Tullgren, 1901: 219, 259. Female ho- lotype from Kark, Chile in the SMNH, examined. Metepeira labyrinthea:—Roewer, 1942: 868. Bonnet, 1957: 2821. Erroneous synonymy. Description. Female from Rio Gallegos, Santa Cruz Province, Argentina. Carapace reddish brown with white setae, light around eyes with lateral posterior exten- sions (Fig. 127). Legs light yellow, articles annulated distally. Femur I with row of three to four macrosetae on anterior side: one to four on anteroventral side. Dorsum covered with denser, longer, black and white setae than in most species. Folium mostly white with brown speckles (Fig. 127). Venter brownish gray with wide white median line; pair of large white spots on either side of spiracle. A pair of thin white lines, parallel to and on either side of median line, sometimes connect to a pair of thinner transverse white lines: one just posterior to the epigynal groove, one just anterior to the spinnerets (Fig. 128). Sternum dark reddish brown (Fig. 128). Ratio of eye diameters: posterior medians and anterior medians 1.2, anterior laterals 1.4, posterior laterals 1.5. Anterior median eyes separated by 1.8 diameters, posterior median eyes by 1.0, anterior median eyes separated from anterior laterals by 4.2 di- ameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. To- tal length 9.5 mm. Carapace 4 mm long, 3.4 wide. First femur 4 mm, patella and tibia 4.4, metatarsus 3.7, tarsus 1.2. Sec- ond patella and tibia 3.9 mm, third 2.5, fourth 3.8. Male from Rio Gallegos, Santa Cruz Province, Argentina. Carapace reddish brown with lighter eye region, lateral pos- terior extensions, and median arrowhead mark (Fig. 125). Legs light yellow, articles distally annulated reddish brown. Femur I with row of three to four macrosetae on anterior side; five to six on anteroventral side. Dorsal folium mostly white with brown speckles (Fig. 125). Venter and sternum as in female (Fig. 126). Ratio of eye diameters: posterior medians and an- terior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separated by 1.7 diameters, posterior median eyes by 1.2, anterior median eyes separated from anterior laterals by 2.8 di- ameters of anterior lateral eyes, lateral eyes separated by 0.4 their diameters. To- tal length 5.7 mm. Carapace 2.8 mm long, 2 wide. First femur 4.3 mm, patella and tibia 4.3, metatarsus 4.1, tarsus 1.2. Sec- ond patella and tibia 3.5 mm, third 1.9, fourth 2.9. Diagnosis. Females are easily separated from other species in the M. nigriventris METEPEIRBA °® Piel AT group by the thick posterior epigynal lobes (compare Fig. 123 with Fig. 119). The dis- tal embolic apophysis does not protrude out from under the terminal apophysis (Fig. 124) as it does in M. galatheae, M. tarapaca, and M. nigriventris (Figs. 87, 95, 119). The embolus of M. karkii differs from M. calamuchita by lacking the in- wardly curved “dewlap” under the embo- lus (compare Fig. 121 with Fig. 103). Variation. Average body length of 13 fe- males examined 6.8 mm, range 5 to 8.2 mm. Average body length of five males ex- amined 4.4 mm, range 2 to 5.3 mm. Dor- sal folia vary from white with little contrast and indistinct fleur-de-lis to darker with more contrast and distinct fleur-de-lis. Natural History. This species appears to be strongly seasonal: mature specimens have been collected between November and March (Fig. 303). Median elevation, 300 m. Spiders are found in pampas (tree- less grassland). Distribution. Lower altitudes in south- ern Argentina and Chile (Map 5). Records Examined. ARGENTINA Chubut: Puerto Piramides, Peninsula Valdes, 42°34'S, 64°17’W, 12.xi.1988 (V. & B. Roth, CAS). Neuquén: Laguna Blanca, 39°3’S, 70°23'W, 15.iii.1959 (J. Nara, MACN); Zapala, 38°54'S, 70°4'W, 15.i1.1958 (fe R. Na- vas, MACN); Zapala, Laguna Blanca, 38°54'S, 70°4'W, 15.1.1959 (J. R. Navas, MACN). Rio Negro: Cerro Alto [?], 41°8’S, 70°40’W (MACN): Coronel Juan José Gomez, 39°2'S, 67°39'W, 15.xi.1945 (Ibarra Grasso, MLP); Ne-Luan, 41°25'S, 68°45’W (MACN). Santa Cruz: Laguna Calafate, Precordilfera [?], 50°55’'S, 70°9'W, 22.1.1967 (P. San Martin, MCZ): Rio Gallegos, 51°38’S, 69°13’W, 20.i.1967 (P. San Martin, MCZ). BOLIVIA Santa Cruz: Patagonia: Estancia Monte, cerca Rio Coyby [?], 50°14’S, 68°55'W (B. Brown, AMNH). CHILE Magallanes: 4 km W La- guna Amarga, 50°59’S, 72°49'W, 8.xii.1966 (E. I. Schlinger & M. E. Irwin, CAS); Kark, 51°17'30"S, 72°32'30"W, 13.iii.1899 (FE. Nordenskidld, NRMS). Metepeira compsa Group: The three species in the M. compsa group include Metepeira compsa, Metepei- ra roraima, and Metepeira gressa. Along with the Metepeira nigriventris group, this group has a median apophysis with teeth on the face of the keel. Unlike the Mete- 48 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 peira nigriventris group, this group has smaller, slimmer distal embolic apophyses that do not arch up over the embolus tip (compare Fig. 149 with Fig. 84). The openings to the epigynum have distinct, sclerotized edges and are clearly visible ei- ther as circles (Figs. 131, 134) or eye- shaped ovals (Figs. 143, 151). 16. Metepeira compsa (Chamberlin) Figures 129-140, 311; Map 8 Aranea compsa Chamberlin, 1916: 252, fig. 6, 2. Fe- male holotype from Ollantaitambo, Cusco, Peru, in the MCZ, examined. Bonnet, 1955: 462. Araneus labyrintheus:—Petrunkevitch, 1926: 27. Er- roneous synonymy. Metepeira virginensis Chamberlin and Ivie, 1942: 70, figs. 188-190, 2, d. Female holotype from St. Thomas, U.S. Virgin Islands, in AMNH, examined. NEW SYNONYMY. Metepeira latigyna Chamberlin and Ivie, 1942: 70, figs. 191, 192, 2. Female holotype from Porto Ale- gre [Bahia], Brazil, in AMNH, examined. NEW SYNONYMY. Metepeira compsa:—Chamberlin and Ivie, 1942: 71, figs. 193-195, @, 3. Metepeira labyrinthea:—Bryant, 1942: 346. Metepeira perezxi Archer, 1965: 132, figs. 14, 16, 3, 2. Male holotype from subexperiment station, Is- abela, Puerto Rico, in AMNH, examined. Brignoli, 1983: 276. NEW SYNONYMY. Metepeira vaurieorum Archer, 1965: 133, figs. 15, 19, 3, 2. Male holotype from Usine de Robert, Mar- tinique, in AMNH, lost. Brignoli, 1983: 276. Lo- pez, 1993: 10, 11, figs. 1-4, 11, 6. NEW SYN- ONYMY. Note. Although the type for M. vaurieorum is lost, all examined records from Martinique (includ- ing ones from “Usine de Robert”) belong to M. compsa. Description. Female from Savonet, Cu- ragao, Netherlands Antilles. Carapace light around eyes with lateral posterior exten- sions (Figs. 135, 137). Only tibia IV ringed. Femur I with row of four macro- setae on anterior side; two or three fine setae on anteroventral side. Dorsum of ab- domen with usual Metepeira folium, though whiter than usual in some speci- mens (Figs. 135, 137). Venter with wide white median line flanked by two thin (Fig. 136) or wide (Fig. 138) white lines; pair of white spots on either side of spi- racle. Sternum has wide median white line widening anteriorly, sometimes broken (Figs. 136, 138). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separated by 1.3 diameters, posterior median eyes by 0.7, anterior median eyes separated from an- terior laterals by 1.8 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 3.9 mm. Car- apace 1.8 mm long, 1.4 wide. First femur 1.9 mm, patella and tibia 1.9, metatarsus 1.5, tarsus 0.7. Second patella and tibia 1.7 mm, third 0.9, fourth 1.5. Male from Savonet, Curagao, Nether- lands Antilles. Carapace light around eyes with lateral posterior extensions. Slightly lighter median triangular mark anterior to thoracic furrow (Fig. 139). Legs lightly ringed. Femur I with row of four macro- setae on anterior side, four on anteroven- tral side. Dorsum, venter, and sternum as in female (Figs. 139, 140). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separat- ed by 1.3 diameters, posterior median eyes by 0.6, anterior median eyes separated from anterior laterals by 1.4 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 3.6 mm. Carapace 1.8 mm long, 1.4 wide. First femur 2.7 mm, patella and tibia 2.8, metatarsus 2.4, tarsus 0.9. Second patella and tibia 2.3 mm, third 1.2, fourth 1.8. Diagnosis. Unlike other species, the openings to the epigynum of M. compsa are small and almost perfectly round and sclerotized around the rim (Fig. 131). In Peruvian and Argentinean populations the scape can be extremely wide, often entire- ly covering a ventral view of the openings (Fig. 134). The distal embolic apophysis on the male palp extends straight from its base and parallel to the embolus tip (Figs. 129, 132), in contrast to those of other spe- cies in the M. compsa species group that lift away from the embolus tip (Figs. 141, 149). Variation. Average body length of 22 fe- METEPEIRA ° Piel 49 roraima (i) Figures 129-140. Metepeira compsa (Chamberlin) (sp. 16 [129-131,135, 136,139,140] 12°20’N, 69°7’W; [132] 4°30’S, 81°8’W; [133,134] 4°51’S, 80°46’W; [137,138] 17°5'N, 61°42’W). 129, male palpus, mesal. 130, epigynum, posterior. 131, epigynum, ventral. 132, male palpus, mesal. 133, epigynum, posterior. 134, epigynum, ventral. 135, female, dorsal. 136, female, ventral. 137, female, dorsal. 138, female, ventral. 139, male, dorsal. 140, male, ventral. Figures 141-148. Metepeira roraima new species (sp. 17 [141-147] 3°22’N, 60°19’W; [148] 3°21’'N, 76°33’W). 141, male palpus, mesal. 142, epigynum, posterior. 143, epigynum, ventral. 144, male, dorsal. 145, male, ventral. 146, female, dorsal. 147, female, ventral. 148, female, dorsal. Scale bars: dorsum and venter figures 1.0 mm. males examined 5 mm, range 3.5 to 7.3 mm. Average body length of 18 males ex- amined 3.6 mm, range 2.4 to 4.7 mm. Enormous variation in the shape of the scape can be seen in this species: popula- tions in the Caribbean and northeastern South America have a narrow scape (Fig. 131), whereas populations in Argentina and Peru have a thick scape (Fig. 134). Somewhat less consistently parallel differ- ences can be seen in the shape of the me- dian apophysis: flagella are centered in Ca- ribbean and northeastern South America (Fig. 129) but shifted to the left in Argen- tina and Peru (Fig. 132). Further subtle differences appear in the shape of the em- 50 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 bolus tip and embolus arm (compare Fig. 129 with Fig. 132). These male and female genital differences are not sufficiently con- sistent to provide solid evidence for spe- ciation; thus, these populations will be treated as one species. Natural History. Mature specimens have been collected throughout the year (Fig. 311). Altitudes range from just above sea level (for Caribbean and eastern South American localities) to 4,000 m (for Pe- ruvian localities). Spiders have been found in everything from second growth mixed exotics (e.g., mango, citrus, banana) and mangroves to grasses and semidesert chap- arral. Distribution. Puerto Rico to northern Argentina, but absent from the Amazon (Map 8). Records Examined. ARGENTINA Buenos Aires: Punta Lara, 34°49'S, 57°59’W, 15.11.1941 (F. Moneos, MACN). Chaco: Basail, 27°52'S, 59°18’'W, 18.iv.1942 (MACN). Corrientes: Paso de la Patria, 27°19’S, 58°35'W, 151.1966 (M. E. Galiano, MEG). Entre Rios: Parana, 31°44’'S, 60°32'W (Rosenzwaig, MLP); Salto Grande, 31°13’S, 57°56'W, 15.iii.1964 (M. E. Galiano, MEG). Misiones: Posadas, 27°23'S, 55°53’ W, 15.ix.1963 (M. E. Galiano, MEG). Neuwquén: Piedra del Aguila, 40°3’S, 70°5’'W, 18.vii.1978 (Mision Cien- tifica Danesa, ZMUC). Santa Fé: Tostado, 29°14'S, 61°46'W (A. Giai, MACN). BOLIVIA La Paz: Apolo, 14°43'S, 68°31'W, 10.viii. 1989 (L. E. Pefia, AMNH). BRAZIL Bahia: Arquipélago dos Abrolhos, 17°40'S, 38°50'W, 28.xii.1887 (U.S.F.C., Voy. of Albatross, USNM); Parque Ondina, Salvador, 12°59’S, 38°31'W, 25.vii.1962 (A. F. Archer, AMNH); Porto Alegre, 18°5'S, 39°34’'W (AMNH). Minas Gerais: Pedra Azul, 16°1'S, 41°16’W, 15.xii.1970 (F. M. Oliveira, AMNH). Pernambuco: Pernambuco [?], 8°3'S, 34°54’W, 12.ii1.1927 (SMF), 8.iii.1955 (SMF). Rio de Janeiro: Lagomar [?], Macae, near sea, 22°23'S, 41°47'W, 17.vii.1986 (R. L. C. Baptista, MZSP). Rio Grande do Sul: Montenegro, 29°42'S, 51°28'W, 3.xi.1977 (E. H. Buckup, MCN); Rambo [?], 30°4'0"S, 51°11’W (MNRJ); Sao Leopoldo, 29°46'S, 51°9'W, 14.x.1965 (Celia Valle, MZSP). Sdo Paulo: Rubiao, Jr. [?], Bo- tucatu, 22°52’S, 48°26’W, 25.iv.1988 (R. L. C. Bap- tista, MZSP); Sao Paulo, Guaruja, 24°0’S, 46°16’W, 24 vii.1983 (R. Sievers, AMNH). BRITISH WEST INDIES Anegada: 18°45'N, 64°20'W, 12.xi.1966 (Harry Beatty, AMNH); center, nr. salt pond, 18°45'N, 64°20'W, 4.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH): West end, 18°45’N, 64°22'W, 4.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH); Anegada Settlement: 18°45’N, 64°20'W, 5.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Antigua: Coolidge Airport, 17°6'N, 61°51’W, 15.xi.1967 (N. L. H. Krauss, AMNH): Devil’s Bridge, 17°5'N, 61°42'W, 30.vi.1963 (Rick & E. N. Kjellesvig-Waering, AMNH); Jolly Beach, 17°4’N, 61°53’W, 20.ix.1963 (E. N. Kjellesvig- Waering, AMNH); Lignum Vitae Bay: Jolly Beach [?], 17°4'N, 61°53'W, 19.vi.1968 (E. N. Kjellesvig-Waer- ing, AMNH); Redonda Island, from webs spun be- tween boulders on beach, 16°55'’N, 62°19'W, 10.iv.1956 (J. F. G. Clarke, USNM); Reeds Point, nr. Jolly Beach, 17°4'N, 61°53’W, 2.vii.1963 (E. N. Kjel- lesvig-Waering, AMNH); Saint John’s, 17°6'N, 6L°51'W, 15.viii. 1967 (N. L. H. Krauss, AMNH), 15.xii. 1967 (N. L. H. Krauss, AMNH). Dead Man’s Chest [?]: 18°22'N, 64°34'W, 26.v.1966 (Island Pro- ject Staff, Univ. of Puerto Rico, AMNH). East Seal Dog: 18°30'N, 64°25'W, 7.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). George Dog Is- land: 18°30'N, 64°27'W, 7.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH); 4 mi W Virgin Gorda Isl, high on dry cliffs, 18°30'N, 64°27'W, 25.ii1.1979 (K. Johnson, AMNH). Ginger Island: 18°24'N, 64°28’W, 25.v.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Great Dog: 18°29'N, 64°27'W, 7.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Green Cay, near Tortola: 18°27'N, 64°42'30"W, 14.viii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH). Island O. near Anegada Settlement: 18°45'N, 64°20’W, 5.vi.1966 (Is- land Project Staff, Univ. of Puerto Rico, AMNH). Is- land R. near Anegada Settlement: 18°45’'N, 64°20'W, 4.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Jost Van Dyke: 18°28'N, 64°45'W, 30.viii.1965 (H. Heatwole, R. Levins & F. Mac- Kenzie, AMNH). Large mangrove patch nr. Settle- ment Anegado: 18°45'N, 64°20'W [?], 5.vi.1966 (Is- land Project Staff, Univ. of Puerto Rico, AMNH). Lit- tle Camanoe: 18°28'N, 64°33’W, 25.v.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Little Jost Van Dyke: 18°27'N, 64°43'W, 27.vii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH). Little Tobago: 18°26'N, 64°51'W, 4.iv.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Montserrat: Gage’s Soufriére [?], 16°43’N, 62°10’W, 28.vii.1972 (N. L. H. Krauss, AMNH); Plymouth, 16°42’N, 62°13'W, 15.xi.1967 (N. L. H. Krauss, AMNH). Neck- er Island: 18°33'N, 64°21'W, 6.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). nr. Anegada Set- tlement: Byers Bache [P?], 18°45'N, 64°20’'W, 5.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Peter Island: 18°22'N, 64°35'W, 6.vii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH), 12.v.1966 (Percy Chubb, AMNH). Prickly Pear Is- land: 17°10'N, 61°48'W, 6.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Saint Christo- pher Island: Basseterre, 17°18'N, 62°43'W, 6.ii.1968 (B. Malkin, AMNH). Salt Island: 18°23'N, 64°31'W, 24.v.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Sandy Key: near Tortola, 18°13'N, 63°7'W, 31.viii.1965 (H. & A. Heatwole, AMNH). Tobago Is- land: 18°27'N, 64°48'W, 2.iv.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Tortola: 18°27'N, 64°36'W, 8.vii.1958 (A. F. Archer, C. Hel- sley, & M. Sanderson, AMNH); Beef Island, 18°27'N, 64°31'W, 21.vii.1965 (H. Heatwole, R. Levins & F. MacKenzie, AMNH); Greater Camanoe Isl, 18°29'N, 64°32’W, 1.vii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH); Little Thatch Island, 18°23'N, 64°42'W, 16.viii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH); Long Bay Estate, 18°24'N, 64°41'W, 24.vii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH); Marina Key [?], 18°27'N, 64°36'W, 4.vii.1965 (H. Heatwole & R. Levins, AMNH); Prospect Reef, S Roadtown, 18°25'N, 64°36'W, 23.iii. 1979 (K. Johnson, AMNH), 31.iii.1979 (K. Johnson, AMNH); Road to town, 18°27’N, 64°36'W, 15.vii.1972 (N. L. H. Krauss, AMNH): Sandy Spit [?], 18°27’N, 64°36’W, 14.viii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH). Tortola Island: 18°27'N, 64°36'W, 15.vii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH). Virgin Gorda: 18°30'N, 64°24’W, 15.xi. 1966 (Harry Beatty, AMNH): Baths & Devil’s Bay, 18°26'N, 64°27'W, 25.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH); Cooper Mine Trail, 18°26'N, 64°26’W, 25.vi.1966 (Is- land Project Staff, Univ. of Puerto Rico, AMNH); Sa- vana Bay & Pond Bay, 18°28’N, 64°25’W, 27.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH); Trellis Bay (W end of island), nr. harbor, 18°27'N, 64°26'W, 27.iii.1979 (K. Johnson, AMNH); Virgin Gorda Mountain, 18°30'N, 64°24'W, 26.vi.1966 (Is- land Project Staff, Univ. of Puerto Rico, AMNH). Virgin Islands Area: Island Q [?], 18°45'N, 64°20'W, 5.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). West Seal Dog Island: 18°29'N, 64°28'W, 7.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). CHILE Tarapaca: Arica, 18°29’S, 70°20'W, 28.i.1973 (W. C. Sedgwick, MCZ); Azapa Arica, 18°31'S, 7O°1L1’W, 9.xi.1955 (L. E. Petia, IRSNB):; Lluta, 18°24'S, 70°19’W, 12.xi.1955 (L. E. Pefia, IRSNB); Rio Lluta, 18°24'S, 70°19’W, 12.xi.1955 (L. E. Pena, IRSNB); Sobraya, 18°32'S, 70°9'W, 10.xi.1955 (L. E. Petia, IRSNB); Taltape, Camarones Valley, 18°59'S, 69°47'W, 29.i1.1973 (MCZ). COLOM- BIA Magdalena: Cabaiia “Villa Culebra,” 10 km E Station Marta, 11°12’N, 74°7’W, 15.x.1985 (H.-G. Miiller, SMF); Casajera [P], 11°O0’N, 74°15’W, 1.ii.1974 (J. A. Kochalka, JAK); Cienaga, 11°1’N, 74°15'’W, 30.i1.1974 GE A. Kochalka, JAK); Gaira, 11°11'N, 74°13'W, 15.xii.1975 (W. Eberhard, MCZ). DOMINICAN REPUBLIC Barahona: Patos, 10°38'N, 61°52'’W, 24.ix.1944 (R. H. Montgomery, AMNH). FRENCH WEST INDIES Guadeloupe: Deshaies, 16°18’N, 61°48’W, 28.vi.1960 (C. & P. Vau- rie, AMNH): Domaine Duclos [?], 16°16’N, 61°31'W, 25.vi.1960 (C. & P. Vaurie, AMNH), 15.vii.1960 (C. & P. Vaurie, AMNH); Marie-Galante, in citrus, 15°56'N, 61°16’W, 15.iii.1977 (W. H. Whitcomb, FSCA): Pointe-a-Pitre, [let A Boissard, 16°14'N, 61°34'W, 26.vi.1960 (C. & P. Vaurie, AMNH): Terre- de-Haut, Les Saintes, 15°58’N, 61°35/W, 2.vii.1960 (C. & P. Vaurie, AMNH). Martinique: Ansemitian [?], METEPEIRA ® Piel Sail Trois-Ilets, 14°33'N, 61°2'W, 10.vi.1960 (C. & P. Vau- rie, AMNH):; Diamant, 14°29'N, 61°2’W, 17.vi.1960 (C. & P. Vaurie, AMNH), 18.vi.1966 (C. & P. Vaurie, AMNH); Fort de France, 14°36’N, 61°5’W, 15.xii.1950 (N. L. H. Krauss, MCZ); Pointe Ferret, La Caravelle, 14°45'N, 60°54’W, 19.vi.1960 (C. & P. Vaurie, AMNH); Sainte-Anne, 14°26’N, 60°53’W, 20.vi. 1966 (C. & P. Vaurie, AMNH): Usine de Robert [P], 14°41'’N, 60°57'W, 16.vi.1960 (C. & PB. Vaurie, AMNH). GRENADA nr. Saint Georges: 12°3'N, 61°45'W, 3.vi.1950 (Leo Isaacs, AMNH). NETH- ERLAND ANTILLES Bonaire: Red Pond [?], 12°12'N, 68°15'W, 3.i1.1968 (B. Malkin, AMNH). Cu- racgao: 12°11'N, 68°58'W, 13.i1.1968 (B. Malkin, AMNH); Fuik (Oostpunt), muddauber nests, 12°4'N, 68°49'W, 20.xii.1962 (H. W. Levi & B. de Jong, MCZ); Groot Sint Joris, plantation, 12°14’N, 69°3'W, 22,.xii.1962 (H. W. Levi & B. de Jong, MCZ); Groote Berg, 12°11'N, 69°0'W, 19.xii. 1962 (H. W. & L. Levi, MCZ): Hato, 12°11'N, 68°58’W, 28.xii.1967 (B. Mal- kin, AMNH), 7.i.1968 (B. Malkin, AMNH): Piscadera Baai, 12°8'N, 68°59’W, 18.xii.1962 (H. W. Levi, MCZ), 20.xii.1962 (H. W. Levi, MCZ): Savonet; shady ravine, 12°20'N, 69°7’W, 28.xii.1962 (H. W. Levi, MCZ):; SE of airport, 12°10’N, 68°54’W, 20.xii.1962 (H. W. Levi & B. de Jong, MCZ): Siberié, 12°14'/N, 69°3’W, 25.xii. 1962 (H. W. Levi, MCZ): Sint Jan, 12°15'N, 69°6’W, 25.xii. 1962 (H. W. Levi & B. de Jong, MCZ); Willemstad, Jewish Cemetary, 12°7'N, 68°57'W, 24.xii. 1962 (H. W. Levi, MCZ). Sint Eustatius: Oranjestad, 17°29'N, 62°59'W, 18.i.1968 (B. Malkin, AMNH). Sint Maarten: nr. Juliana Air- port [P], 18°4’N, 63°4'W, 24.ii.1965 (H. Heatwole & F. MacKenzie, AMNH). PARAGUAY Alto Paranda: Taguarazaya [?], 25°30'S, 54°50’W (AMNH). PERU Ancash: Huaraz, 9°32'S, 77°32'W, 6.xii. 1980 (C. Gold, CAS). Apurimac: 35 mi E Abancay, 13°38'S, 72°22'W, 5.i1.1951 (E. S. Ross & Michelbacher, CAS); 40 mi E Abancay, 13°38’S, 72°20'W, 4.iii.1951 (E. S. Ross & Michelbacher, CAS); Abancay, 13°38'2"S, 72°52'53"W, 6.iii.1951 (E. S. Ross & Michelbacher, CAS). Cajamarca: Jaen, 5°42'34"S, 78°48'32"W, 17.v.1967 (A. F. Archer, AMNH). Cusco: 40 mi W Cusco, 13°32'S, 72°33'W, 5.iii.1951 (E. S. Ross & Michelbacher, CAS); Hacienda Urco, near Calea, 13°22’S, 71°54'W, 19.ix.1939 (Karl P. Schmidt, AMNH); Huacerpay [?], 13°37'S, 72°13'W, 10.ix.1993 (J. Ochoa Camara, MCZ); Ollantaitambo, 13°15'17'S, 72°15'48"W, 15.vii.1911 (Yale Peruvian Expedition, AMNH), 15.xii.1980 (C. Gold, CAS): Pisac, 13°25'21"S, 71°50'48’W, 13.xii.1980 (C. Gold, CAS); Urubamba, 13°18’28"S, 72°6'55’"W, 15.i.1965 (Carras- co, MCZ), 18.11.1965 (H. W. Levi, MCZ), 6.viii. 1987 (D. Silva, MUSM). Huanuco: Huanuco, 9°55’S, 76°14'W, 6.x.1946 (J. C. Pallister, AMNH). Ica: km 367 between Ica and Nazca, sandy semidesert, 14°24'S, 75°23'W, 22.41.1952 (W. Weyrauch, CAS): Nazca, 14°50'S, 74°57'W, 15.iv.1951 (P. Aguilar, CAS). Piura: Cerro Prieto, La Brea, 4°41'S, 81°6’W (CAS); Mallares, 4°51'S, 80°46’'W, 8.vi.1941 (H. E. F. & D. E. F, CAS), 13.vii.1941 (WH. E.F & DL ELF 52 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 CAS); N of Negritos, Parifias Valley, 4°41’S, 81°18’W, 9.x.1938 (D. L. & H. E. Frizzell, CAS); nr. Negritos, 4°42'S, 81°18'W, 5.iii.1939 (H. E. F, CAS): nr. Se- chura, 5°33'S, 80°51'W, 4.xi.1941 (D. L. F, CAS); Ne- gritos, 4°49'S, 81°18’W, 15.xi.1934 (H. E. F, CAS), 15.iii.1941 (H. E. F, CAS): Parifias, 4°30’S, 81°8’W, 7.v.1939 (D. L. & H. E. Frizzell, CAS), 31.v.1939 (D. L. & H. E. Frizzell, CAS), 15.x.1939 (D. L. & H. E. Frizzell, CAS); Parifias Valley, 4°30'S, 81°8'’W, 6.iii.1939 (D. L. & H. E. Frizzell, CAS), 8.iv.1939 (D. L. & H. E. Frizzell, CAS); Portachuelo, 4°59’S, 79°54'W, 29.iv.1939 (M. H. & H. E. Frizzell, CAS): Quebrada Mogollon, 4°32’S, 81°4’W, 30.iv.1939 (D. L. & H. E. Frizzell, CAS), 11.vi.1939 (D. L. & H. E. Frizzell, CAS); Rio Chira Valley, N of Amotape, 4°53'S, 81°1L'W, 13.xi.1938 (D. L. & H. E. Frizzell, CAS): S of Sechura, 5°33'S, 80°51'W, 25.x.1941 (D. L. F, CAS): San Lorenzo [?], Zona Alta Este Herrera, 5°4'S, 79°47'W, 12.iv.1969 (P. Aguilar, MCZ); Sullaiia, 4°53'S, 80°41'W, 8.x.1939 (D. L. & H. E. Frizzell, CAS), 5.x.1941 (D. L. & H. E. Frizzell, CAS). Tum- bes: 34 km E, 25 km N Punta Parifias, 4°18’S, 80°51’W, 1.1.1939 (D. L. & H. E. Frizzell, CAS). SAINT LUCIA Castries: 14°1'N, 61°0'W, 28.vii.1963 (E. N. Kjellesvig-Waering, AMNH). TRINIDAD & TOBAGO Tobago Island: Buccoo Bay, 11°10'N, 60°48'W, 15.viii.1965 (E. N. Kjellesvig-Waering, AMNH); Guayaguayare. Point, 10°8'’N, 61°2'W, 14.ix.1963 (E. N. Kjellesvig-Waering, AMNH); Pi- geon Point, 11°10'N, 60°50’W, 18.viii.1937 (E. D., MCZ); Salybia Bay, by shore, 10°42'N, 61°2’W, 15.ii.1972 (J. A. L. Cooke, MCZ); Toco, 10°50'N, 60°57'W, 19.iv.1964 (Erik N. Kjellesvig-Waering, AMNH). Trinidad: Gasparee, 10°46'N, 61°19'W, 3.xi.1944 (R. Montgomery, AMNH). URUGUAY Co- lonia: Punta Gorda [?], 34°28'S, 57°51'W, 25.ii.1968 (R. Capocasale & L. Bruno, CAS). USA Puerto Rico: Algodones Key, 18°12’N, 65°41'W, 15.x.1964 (H. Heatwole, R. Levins & F. MacKenzie, AMNH); Bal- neario [?] Guajataca, 18°21'N, 66°55'W, 4.vii.1958 (A. F. Archer, AMNH): Bafios de Coamo, 17°59’'N, 67°3'W, 2.iv.1990 (H. W. & L. Levi, MCZ), 3.iv.1990 (H. W. & L. Levi, MCZ); Barranquitas, 18°11'N, 66°18'W, 28.xii.1977 (J. Coddington, USNM); below Quebradillas along old RR track, 18°28'N, 66°56'W, 30.iii1.1989 (H. W. & L. Levi, MCZ): Cabeza de Perro Island, 18°15’N, 65°35’W, 16.i1.1965 (H. Heatwole & F. MacKenzie, AMNH); Cayo Don Luis, 17°57’'N, 66°58'W, 12.i.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH); Cayo Norte, off Culebra, 18°20'N, 65°15’W, 14.iv.1965 (H. Heatwole & F. MacKenzie, AMNH); Cayo Pifierito, 18°15'N, 65°36'W, 24.ix.1964 (H. Heatwole & F. MacKenzie, AMNH):; Channel at Culebra, Isla del Diablo, 18°23'N, 65°40'W, 12.viii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH); Cuevas de los Alfaros, Barrio Mora, 18°29'N, 67°1'W, 20.vii.1958 (A. F. Ar- cher & Rolle, AMNH):; Culebra Island, 18°19’N, 65°17'W, 19.vii.1965 (F. MacKenzie, AMNH); Cule- bra, near Dewey, 18°18'N, 65°18’W, 10.viii.1965 (Is- land Project Staff, Univ. of Puerto Rico, AMNH); Cu- lebrita Island, 18°19'N, 65°14'W, 15.iv.1965 (H. Hea- twole & F. MacKenzie, AMNH), 11.viii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH); Dese- cheo Is, 18°23’N, 67°29'W, 19.11.1914 (AMNH); De- secheo Island, 18°23'N, 67°29'W, 28.v.1965 (H. Hea- twole, R. Levins & F. MacKenzie, AMNH); Faro de Cabo Rojo, 18°5'N, 67°9’W, 13.iii.1961 (F. Rolle, AMNH); Frank Key, Isl. #13 nr. La Parguera area, 17°58'N, 67°3'W, 14.i.1966 (Island Project Staff, Univ. of Puerto Rico, MCZ); Heatwole Island, off Culebrita, 18°19’N, 65°13’W, 14.iv.1965 (H. Heatwo- le & F. MacKenzie, AMNH); Hormiqueros, 18°9'N, 67°S8'’W, 16.11.1962 (Aida Velez, AMNH), 11.iii.1962 (Aida Velez, AMNH), 19.iii.1962 (Aida Velez, AMNH); Isabela, subexperiment station, 18°30'N, 67°1'W, 17.viii.1957 (A. F. Archer, AMNH); Jayuya, coffee plantation, 18°13'N, 66°37'W, 23.iii.1986 (H. W. & L. Levi, MCZ); Juana Diaz, 18°3'N, 66°31’W, (xi LOW (jay Eai€axico,, JEC)LeviniseRockaaial: 18°12’N, 65°41'W, 15.x.1964 (H. Heatwole, R. Levins & F. MacKenzie, AMNH); Loma Tinaja [?], S of La- guna Cartagena, 17°59'N, 67°6'W, 5.vii.1958 (A. F. Archer & M. Sanderson, AMNH); Luquillo, near beach, 18°23'N, 65°44'W, 24.i1.1932 (A. S. Mills, AMNH), 24.xii.1985 (V. & B. Roth, CAS), 27.iii.1988 (H. W. & L. Levi, MCZ); Mayagiiez, 18°12'N, 67°9'W, 15.vii.1958 (A. F. Archer, MCZ); Mayagiiez, 5 km N university campus, 18°12'N, 67°9'W, 5.i.1964 (MCZ); Mayagiiez: Las Mesas, 18°11'N, 67°6'W, 20.xi.1960 (F. Rolle, AMNH); Mayagiiez, university farm N university campus, 18°12’N, 67°9'W, 7.ii.1964 (MCZ); McKenzie Key, Key #3 [P?], 18°12'N, 65°41'W, 29.x.1964 (H. Heatwole, R. Levins & F. MacKenzie, AMNH); Mona Island, Serrallés, 18°5'N, 67°54'W [P], 7.iv.1944 (Beatty, MCZ); Muertos Is- land, beating bushes, 17°54'N, 66°32'W, 28.v.1959 (Jordan & Martorell, AMNH); Muertos Island, mud nests of Sceliphron caementarium, 17°54'N, 66°32'W, 27.v.1959 (Medina & Martorell, AMNH); N slope Ti- naja, nr. Cartagena Lagoon, beating and sweeping, 18°23'N, 67°10’W, 5.vii.1958 (M. W. Sanderson, AMNH);: Palominitos, 18°20'N, 65°34'W, 16.vi.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH): Palomino Island, 18°21'N, 65°34'W, 7.xi.1964 (H. Heatwole & F. MacKenzie, AMNH); Parguera, 17°59'N, 67°3'W, 25.iii.1990 (H. W. & L. Levi, MCZ); Patillas, 18°0’N, 66°1’W, 3.iv.1931 (Mills & Leonard, AMNH); Pico Atalaya, Rte. 2, nr. Afiasco, 18°18'N, 67°11'W, 3.vii.1958 (M. W. Sanderson, AMNH); Pi- fieros Island, 18°15’N, 65°35’'W, 24.ix.1964 (H. Hea- twole & F. MacKenzie, AMNH); Playa de Humacao, 18°10'N, 65°45'W, 23.v.1964 (MCZ); Quebradillas, around hotel, 18°29'N, 66°56’W, 30.iii.1989 (H. W. & L. Levi, MCZ); Ratones Island, 17°56’N, 66°17'W, 19.xi.1964 (H. Heatwole & R. Levins, AMNH); Rom- ero III [?], 17°57'N, 67°0’W, 13.i.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH); Rubianes, Caimito Bajo [?], Cord. Jaicoa, 18°26'N, 67°8'W, 19.vii.1958 (A. F. Archer & Rolle, AMNH); Santurce, 18°27'N, 66°4'W, 28.iii.1931 (A. S. Mills, AMNH); South of Corozo, Cabo Rojo salt flats, edge of salt ponds, 17°56'N, 67°11'W, 23.11.1990 (H. W. & L. Levi, MCZ); Valle de Lajas, 18°1'N, 67°8'W, 3.vi.1958 (A. F. Archer, AMNH): xeric hills N of Gudnica, 18°0'N, 66°55’/W, 28.viii.1957 (A. F. Archer, AMNH); Zancudo Island (Isleta Marina), 18°20’N, 65°37'W, 2.xi.1964 (H. Heatwole & F. MacKenzie, AMNH). U.S. VIRGIN ISLANDS Big Cockroach: 18°24'N, 65°4'W, 7.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Congo Cay: 18°22'N, 64°48’W, 12.xi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Grass Cay: 18°22'N, 64°50'W, 12.xi.1966 (Univ. of Puerto Rico, AMNH). Great Saint James Island: 18°19'N, 64°50'W, 13.xi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Hans Lollik Is- land: 18°24'N, 64°55'W, 6.iv.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Little Cock- roaches: 18°25'N, 65°3’W, 7.vi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Little Hans Lol- lick: 18°25'N, 64°54'W, 5.iv.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Little Saint James Is- land: 18°18'N, 64°50'W, 13.xi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Mingo Cay: 18°22'’N, 64°49'W, 12.xi.1966 (Island Project Staff, Univ. of Puerto Rico, AMNH). Saint John: 18°20'N, 64°45'W, 9.iii. 1925 (F. E. Lutz, AMNH), 11.vii.1958 (A. F. Archer & M. Sanderson, AMNH), 15.xii.1967 (N. L. H. Krauss, AMNH); Cruz Bay, 18°20'N, 64°48'W, 27.11.1964 (A. M. Chickering, MCZ); nr. Cinnamon & Hart Bays on W half of island, 18°20’N, 64°46'W, 2.viii.1976 (D. E. & D. N. Rosen, AMNH). Saint Thomas: 18°24'N, 64°55'W, 24.11.1925 (AMNH), 24.vii.1925 (A. Petrunkevitch, CAS), 24.xi1.1925 (AMNH); Crown Mountain, 18°21'N, 64°58'W, 30.viii.1957 (A. F. Archer, AMNH), 7.vii.1958 (A. F. Archer, AMNH); Denmark Hill [?], 18°24'N, 66°55'W, 1.ix.1957 (A. F. Archer, AMNH): Flagstok Hill, Stumpy Bay, 18°22’N, 65°0'W, 9.vii.1958 (A. F. Archer, AMNH); Harman’s, Char- lotte Amalia, 18°21'N, 64°56’W, 2.ix.1957 (A. F. Ar cher & family, AMNH); Hassell Island, 18°20'N, 64°56'W, 1.ix.1957 (A. F. Archer, AMNH), 15.ii.1964 (A. M. Chickering, MCZ), 10.viii.1966 (M. L. Pres- sick, AMNH). St. Croix: Christiansted, 17°45'N, 64°42'W, 3.vi.191]1 (AMNBH), 15.i1.1955 (A. M. Nad- ler, AMNH); East End, 17°45'N, 64°40’W, 15.xii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH); Green Key, 17°46'N, 64°40’W, 17.iv.1964 (H. Hea- twole, MCZ); Mount Eagle, dryish forest, 17°46'N, 64°49'W, 15.xii.1965 (Island Project Staff, Univ. of Puerto Rico, AMNH); Protestant Cay, 17°45’N, 64°42’W, 18.iv.1964 (H. Heatwole, MCZ): St. Croix, 17°45'N, 64°54'W, 1.ix.1966 (Chickering, MCZ), 6.ix.1966 (Chickering, MCZ). VENEZUELA Depen- dencias Federales: Patos, 10°38'N, 61°52'W, 23.ix.1944 (R. H. Montgomery, AMNH). Districto Federal: Punta Tanaguarena, in coastal bldg. & gar- den, 10°37'N, 66°48'W, 26.xii.1970 (W. B. Peck, CAS). METEPEIRA ® Piel 53 17. Metepeira roraima new species Figures 141-148, 316; Map 8 Holotype. Female from Rio Surumu, Roraima, Brazil, x.1966, M. Abrorenga, in MZSP. The specific name is a noun in apposition after the locality. Description. Female holotype. Brown carapace; lighter around eyes (Fig. 146). Coxae, femora, tibiae, and patellae tan, lighter ventrally. Metatarsi, tarsi white. Fe- mur I with row of three or four macrose- tae on anterior side; none on anteroventral side. Dorsal folium with typical Metepeira fleur-de-lis pattern (Fig. 146). Venter of abdomen brown with wide white median line, flanked by thinner white lines that to- gether form a T-shape posteriorly (Fig. 147). Pair of small white spots on either side of T-shape mark. Sternum brown with median white line (Fig. 147). Ratio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 1.5, posterior laterals 1.3. Anterior median eyes separat- ed by 1.2 diameters, posterior median eyes by 0.6, anterior median eyes separated from anterior laterals by 1.9 diameters of anterior lateral eyes, lateral eyes separated by 0.1 their diameters. Total length 4 mm. Carapace 1.8 mm long, 1.4 wide. First fe- mur 1.9 mm, patella and tibia 2.1, meta- tarsus 1.8, tarsus 0.7. Second patella and tibia 1.7 mm, third 1.0, fourth 1.6. Male paratype from Rio Surumu, Ro- raima, Brazil. Carapace brown; lighter around eyes with median white mark (Fig. 144). Legs brown, white at base of femora. Femur I with row of four to five macro- setae on anterior side; four to five on an- teroventral side. Dorsal folium, venter, and sternum as in female (Figs. 144, 145). Ra- tio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 1.6, posterior laterals 1.4. Anterior median eyes separated by 1.4 diameters, posterior median eyes by 0.6, anterior median eyes separated from anterior laterals by 1.6 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 2.9 mm. Carapace 1.3 mm long, 1.1 wide. First femur 1.9 mm, patella and 54 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 tibia 2, metatarsus 1.6, tarsus 0.6. Second patella and tibia 1.5 mm, third 0.8, fourth eS, Diagnosis. The epigynum of M. roraima differs from that of M. compsa by having oval openings (Fig. 143) instead of round openings (Figs. 131, 134); it differs from M. gressa by being more translucent and by having a narrower scape (compare Fig. 143 with Fig. 151). Unlike M. gressa, the embolus tip of M. roraima is more bent and the distal apophysis is as wide as the embolus tip (compare Fig. 141 with Fig. 149). Unlike M. compsa, the distal embolic apophysis projects away from the embolus tip (Fig. 141) instead of extending straight from the base and parallel to the embolus tip (Figs. 129, 132). Variation. Average body length of eight females examined 5 mm, range 4 to 7 mm. Average body length of five males exam- ined 3.1 mm, range 2.8 to 4 mm. Colom- bian specimens are much darker than those from northern Brazil and southern Guyana, suggesting the possibility that these populations represent separate cryp- tic species. In fact, the carapace on Col- ombian specimens is often jet black with the white marks on the dorsal folium much reduced (Fig. 148). However, there is little corresponding genitalic difference, especially among males; consequently, these populations are deemed to be con- specific. Natural History. Mature M. roraima specimens from eastern South America have been collected in November and De- cember; specimens from western South America have been collected between March and July (Fig. 316). Median eleva- tion, 1,000 m. Distribution. Western Colombia, north- er Brazil, and southern Guyana (Map 8). Records Examined. BRAZIL Roraima: Rio Suru- mu, Jerrit Rio Branco [?], 3°22'’N, 60°19’W, x.1966 (M. Abrorenga, MZSP). COLOMBIA. Valle del Cau- ca: Atuncela, 3°46’N, 76°42'W, 19.vii.1970 (W. Eber- hard, MCZ): Cali, around house, 3°27'N, 76°31’W (W. Eberhard, MCZ); Palmira, 3°32’N, 76°17'W, 1.iii. 1964 (Ballo, CAS); Rio Pance, below Buenos Ai- res, 3°55'N, 76°8’W, 5.iv.1970 (W. Eberhard, MCZ); Rio Pance, near Cali., 3°21’N, 76°33’'W, 8.v.1970 (W. Eberhard, MCZ), 8.vi.1970 (W. Eberhard, MCZ), 15.vi.1970 (W. Eberhard, MCZ), 23.vi.1970 (W. Eberhard, MCZ), 25.vi.1973 (W. Eberhard, MCZ). GUYANA Upper Takutu: Isherton, on lat. 2, 10 mi E Rupununi River, 2°19'N, 59°22’W, 15.xi.1937 (W. G. Hassler, AMNH). 18. Metepeira gressa (Keyserling) Figures 149-156, 300; Map 8 Epeira gressa Keyserling, 1892: 166, fig. 123, 2. Five female syntypes from Taquara, Rio Grande do Sul, Brazil, in BMNH, examined. One specimen des- ignated lectotype. Epeira seditiosa Keyserling, 1893: 212, fig. 157, d. Male holotype from Rio Grande do Sul, Brazil, in BMNH, examined. NEW SYNONYMY. Araneus gressus:—Petrunkevitch, 1911: 314. Roewer, 1942: 844. Bonnet, 1955: 511. Araneus seditiosus:—Petrunkevitch, 1911: 314. Roewer, 1942: 852. Bonnet (1955: 592) erroneously suggests that Petrunkevitch (1911: 314) synony- mized Araneus scitulus (Blackwall, 1863) with Ar- aneus seditiosus. Eustala_ seditiosa:—Mello-Leitaio, 1943: 179. Erro- neous transfer. Metazygia gressa:—Mello-Leitao, 1943: 187. Erro- neous transfer. Metepeira gressa:—Levi, 1991: 179. Platnick, 1993: 449. Metepeira seditiosa:—Levi, 1991: 180. Platnick, 1993: 449. Note. Examination of voucher specimens of Vi- era suggest that in the behavioral studies of Viera (1986, 1989) and Viera and Costa (1988), the name “Metepeira sp. A” is, in fact, M. gressa. Description. Female from Punta del Es- pinillo, Montevideo, Uruguay. Chelicerae brown, lighter on distal inside margins. Carapace brown; yellowish white across and just behind posterior eye row; median white line reaching thoracic furrow, some- times thickened into arrow shape (Fig. 155). Proximal halves of femora, white; re- mainder black. Patellae black ventrally, white dorsally. Distal halves of tibiae, black; remainder white with black annu- lation. Femur I with row of four macro- setae on anterior side; none on antero- ventral side. Dorsal folium white fleur-de- lis pattern on black, margined with wavy white lines (Fig. 155). Venter black with short, wide, white median line; pair of small white spots on either side of spiracle (Fig. 156). Sternum black with median white line, often broken (Fig. 156). Ratio of eye diameters: posterior medians and anterior medians 1.2, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separated by 1.8 diameters, posterior median eyes by 1.0, anterior median eyes separated from anterior laterals by 2 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 3.3 mm. Carapace 1.8 mm long, 1.4 wide. First femur 1.9 mm, patella and tibia 2, metatarsus 1.6, tarsus 0.7. Second patella and tibia 1.8 mm, third 1.1, fourth 1R6: Male from Punta del Espinillo, Monte- video, Uruguay. Coloration of chelicerae and carapace as in female except median line brighter, thickened into arrow shape (Fig. 153). Leg coloration as in female. Fe- mur I with row of four macrosetae on an- terior side; five to six on anteroventral side. Dorsal folium, venter, and sternum as in female (Figs. 153, 154). Ratio of eye di- ameters: posterior medians and anterior medians 1.0, anterior laterals 1.5, posterior laterals 1.4. Anterior median eyes separat- ed by 1.3 diameters, posterior median eyes by 0.8, anterior median eyes separated from anterior laterals by 1.9 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 3 mm. Carapace 1.8 mm long, 1.3 wide. First fe- mur 2.5 mm, patella and tibia 2.3, meta- tarsus 1.9, tarsus 0.9. Second patella and tibia 1.8 mm, third 1.1, fourth 1.6. Diagnosis. The epigynum of M. gressa differs from that of M. compsa by having oval openings (Fig. 151) instead of round openings (Figs. 131, 134); it differs from M. roraima by being less translucent and by having a wider scape (compare Fig. 151 with Fig. 143). Unlike M. roraima, the em- bolus tip in M. gressa is not as bent, and the distal apophysis is thinner than the point on the embolus where it is attached (compare Fig. 149 with Fig. 141). Unlike M. compsa, the distal embolic apophysis projects away from the embolus tip (Fig. 149) instead of extending straight from the METEPEIRBA ® Piel 55 base and parallel to the embolus tip (Figs. 12932) Variation. Average body length of 14 fe- males examined 4.8 mm, range 3.3 to 6.8 mm. Average body length of five males ex- amined 3.4 mm, range 2.8 to 4.3 mm. Natural History. In Uruguay, M. gressa live in small colonies of up to five individ- uals surrounding the inflorescences of Eryngium sp. (Viera and Costa, 1988). The number of sticky spirals (14 below the hub, 22 above the hub) and radii (c. 40) are the same irrespective of age; however the length and width of the web differ be- tween juveniles (c. 6 and 5 cm, respective- ly) and adults (c. 9 and 7 cm, respectively) (Viera, 1992). Mature specimens are most often collected between September and March at low elevation (Fig. 300). Distribution. Northern Argentina, Par- aguay, and Uruguay (Map 8). Records Examined. ARGENTINA Buenos Aires: Arrecifes, 34°3'S, 60°7’W, 17.i.1939 (Biraben, MLP): Boulogne, 34°30'S, 58°34’W, 15.x.1938 (Prosen, MLP); Colon [?], 33°53’29"S, 61°6'35’W, 20.ix.1944 (Torres, MLP): Moreno, 34°39'S, 58°48’W, 15.ii. 1966 (Rossi and Maury, MACN); San Isidro, 34°27'S, 58°30'W, 15.xii.1937 (Peregra, MACN); San Mi- guel—San Fernando, 34°29'S, 58°39'W, 15.vii.1940 (F. Morrios, MACN); Sierra de la Ventana, 38°9'S, 61°48'W, 15.iii.1939 (S. H. Bavio, MACN), 15.xi.1954 (Fritz, MACN), 31.x.1969 (Carlos Grisolia, MCZ), 15.vii.1972 (Amarrilla, MACN); Tigre, 34°25'S, 58°34'W (J. M. Viana, MACN). Entre Rios: Concep- cién del Uruguay, 32°29'0"S, 58°13'42"W, 4.11941 (Prosen, MLP); Salto Grande, 31°13'S, 57°56’W, 15.iii.1964 (M. E. Galiano, MEG). Santa Fé: Floren- cia Varelo [?], 28°2’S, 59°15’W, 15.xii.1939 (F. Morris, MACN). Tucumdén: 30 km S Concepcion, 27°36'S, 65°35'W, 16.1.1983 (L. E. Pehla, AMNH). PARA- GUAY Itapua: San Luis, 27°6'S, 56°36’W, 15.x.1908 (AMNH). URUGUAY Canelones: Montevideo: De- tras del Cerro, Camino de las tropas [?], 34°45’S, 56°10'W, 6.ii.1963 (R. Capocasale and L. Bruno, CAS). Colonia: Punta Gorda. [?], 34°28'S, 57°51'W, 25.ii.1968 (R. Capocasale and L. Bruno, CAS), 26.ii.1965 (R. Capocasale and L. Bruno, CAS). Mal- donado: Colonia, Cerro de las Animas. Small stones in native forest, 34°46’S, 55°19'W, 30.x.1967 (P. San Martin, MCZ). Montevideo: Punta del Espinillo, 34°50'S, 56°26'W, 12.x.1983 (Carmen Viera, MCZ), 15.xii. 1983 (Carmen Viera, CV). Treinta y Tres: Que- brada de los Cuervos, 33°10'S, 54°27'W, 27.x.1990 (Lopez, Perez, Viera, MCZ). 56 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Metepeira incrassata Group Species in the M. incrassata group all lack a keel on the median apophysis. Males have a distal embolic apophysis that either projects toward the embolus tip (e.g., Fig. 157), forms a sharp corner but does not actually project forward (e.g., Fig. 185), or forms a smoother, rounded, and gradual comer (e.g., Figs. 192, 206). The M. in- crassata species group includes Metepeira maya, Metepeira inca, Metepeira gosoga, Metepeira olmec, Metepeira comanche, Metepeira pimungan, Metepeira triangu- laris, Metepeira arizonica, Metepeira atas- cadero, and Metepeira incrassata. 19. Metepeira maya new species Figures 157-163, 317; Map 14 Holotype. Male from North Bay, Twin Cays, Stann Creek District, Belize, 14.iii.1986, P. Sierwald, in USNM. The specific name is a noun in apposition after the Indian people of southern Mexico and Central America. Description. Female paratype from Twin Cays, Stann Creek District, Belize. Carapace brown with large white eye re- gion, white lateral posterior extensions, and short median posterior extension (Fig. 162). Legs white, ringed black on distal ends of articles. Femur I with row of four macrosetae on anterior side; none on an- teroventral side. Dorsum of abdomen with usual Metepeira folium, though frequently with deep red pigmentation on anterior half (Fig. 162). Venter wide, white median line with pair of large white spots on either side of spiracle (Fig. 163). Sternum with wide median white line widening anteri- orly with constriction in center (Fig. 163). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separated by 1.2 diameters, posterior median eyes by 0.8, anterior median eyes separated from anterior laterals by 2 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 3.9 mm. Carapace 1.8 mm long, 1.4 wide. First femur 2.2 mm, patella and tibia 2.1, metatarsus 1.6, tarsus 0.8. Sec- ond patella and tibia 1.8 mm, third 1.0, fourth 1.6. Male holotype. Carapace dark with light region around eyes and light triangular mark anterior to thoracic furrow (Fig. 160). Legs darker than in female. Femur I with row of four macrosetae on anterior side; four on anteroventral side. Dorsum, venter, sternum as in female except me- dian white line on sternum usually broken (Figs. 160, 161). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.4, posterior laterals 1.2. Anterior median eyes separated by 1.2 diameters, posterior median eyes by 0.9, anterior median eyes separated from an- terior laterals by 1.7 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 3 mm. Car- apace 1.5 mm long, 1.2 wide. First femur 2.2 mm, patella and tibia 2.3, metatarsus 1.8, tarsus 0.9. Second patella and tibia 1.8 mm, third 1.0, fourth 1.4. Diagnosis. Unlike others in the M. in- crassata species group, M. maya and M. inca both have distal embolic apophyses that project forward, forming rounded bumps (Figs. 157, 164). The male palps of these two species differ by the thickness of the flagellae and base on the median apophysis: thicker in M. maya than in M. inca (compare Fig. 157 with Fig. 164). The epigynum of M. maya resembles those of M. inca and M. comanche because the oval epigynal openings are formed out of mem- branous surfaces that are distinctly sepa- rate from the base of the scape (Figs. 159, 166, 187). The epigynal openings of M. maya (Fig. 159) are much wider than those of M. comanche (Fig. 187); and the posterior lobes on M. maya are thickened (Fig. 159) but not pointed as in M. inca (Fig. 166). Variation. Average body length of five females examined 5.2 mm, range 3.9 to 6.3 mm. Average body length of four males examined 2.8 mm, range 2.3 to 3.1 mm. Natural History. Mature specimens have been collected in March through August METEPEIRA ® Piel 597 Figures 149-156. Metepeira gressa (Keyserling) (sp. 18; 34°50’S, 56°26’W). 149, male palpus, mesal. 150, epigynum, posterior. 151, epigynum, ventral. 152, male embolic division, ventral. 153, male, dorsal. 154, male, ventral. 155, female, dorsal. 156, female, ventral. Figures 157-163. Metepeira maya new species (sp. 19; 16°50'N, 88°5’W). 157, male palpus, mesal. 158, epigynum, posterior. 159, epigynum, ventral. 160, male, dorsal. 161, male, ventral. 162, female, dorsal. 163, female, ventral. Figures 164-170. Metepeira inca new species (sp. 20; 4°30’S, 81°8’W). 164, male palpus, mesal. 165, epigynum, posterior. 166, epigynum, ventral. 167, male, dorsal. 168, male, ventral. 169, female, dorsal. 170, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 58 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 (Fig. 317). Habitats range from mangroves at sea level to pine forests at 1,600 m. Distribution. Southern Mexico to Costa Rica (Map 14). Records Examined. BELIZE Stann Creek: Twin Cays, Andera Flats, 16°50'N, 88°5'W, 20.iii.1986 (P. Sierwald, USNM); Twin Cays, North part of West Pond, West of Swamp Doc, 16°50'N, 88°5'W, 28.iii. 1986 (P. Sierwald, USNM); Twin Cays, North- west Point, North Bay, 16°50'N, 88°5’W, 14.iii.1986 (P. Sierwald, USNM); Twin Cays, Northwest Point, North Bay, ground not flooded, 16°50'N, 88°5'W, 14.iii.1986 (P. Sierwald, USNM); Twin Cays, red mangrove, 16°50'N, 88°5'W, 17.v.1985 (Feller, USNM); Twin Cays, white mangrove, 16°50'N, 88°5'W, 15.vi.1984 (Feller, USNM), 5.vi.1985 (Erwin, Mathis, Sims, USNM). COSTA RICA San José: San José, 9°56’N, 84°5’W (Tristan & Banks, MCZ). GUA- TEMALA Chiquimula: Chiquimula, 14°48’N, 89°33'W, 22.vii.1947 (C. & P. Vaurie, AMNH). Sa- catepequez: Antigua, 14°34'N, 90°44’W, 16.vili. 1947 (C. & P. Vaurie, AMNH). MEXICO Chiapas: Com- itan de Dominguez, 16°15'N, 92°8’W, 19.vii.1950 (C. J. & M. Goodnight, AMNH); near Rio San Gregorio, between Comitan and Ocotal, 15°45’N, 92°0'W, 18.vii.1950 (C. J. & M. Goodnight, AMNH); pine for- est, 15 mi NW Arriaga, 16°25'N, 94°1'W, 27.viii.1966 (Jean & Wilton Ivie, AMNH); Tuxtla Gutierrez, 16°45'N, 93°7'W, 10.vi.1964 (Pallister, AMNH). NIC- ARAGUA Matagalpa: Matagaslpa, 12°53'N, 85°57'W, 15.vii. 1989 (R. Reinbold, JMM). 20. Metepeira inca new species Figures 164-170, 318; Map 11 Holotype. Male from Parifias Valley, Piura, Peru, 21.v.1939, D. L. & H. E. Frizzell, in CAS. The specific name is a noun in apposition after the Que- chuan people who once ruled Peru. Description. F emale paratype from Par- ifias Valley, Piura, Peru. Light reddish brown carapace, white around eyes, very faint light mark in center (Fig. 169). Legs white; slightly darker on dorsal surfaces. Femur I with row of four to five macro- setae on anterior side; none on anterov- entral side. Dorsal folium white; darker on shoulders and posteriorly (Fig. 169). Ven- ter grayish brown with wide white median mark inside U-shaped pattern; pair of white spots on either side of spiracle (Fig. 170). Sternum white (Fig. 170). Ratio of eye diameters: posterior medians and an- terior medians 0.9, anterior laterals 1.0, posterior laterals 1.1. Anterior median eyes separated by 1.6 diameters, posterior median eyes by 1.3, anterior median eyes separated from anterior laterals by 2.5 di- ameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. To- tal length 6.8 mm. Carapace 2.9 mm long, 2.3 wide. First femur 3 mm, patella and tibia 3.3, metatarsus 2.6, tarsus 1. Second patella and tibia 2.8 mm, third 1.7, fourth 2-5. Male holotype. Light reddish brown carapace, white around eyes, faint light tri- angular mark in center (Fig. 167). Legs white, save for light reddish brown on dor- sal surfaces of femora and patellae. Femur I with row of three macrosetae on anterior side; two to three on anteroventral side. Dorsal folium white with wavy black lines thickening posteriorly (Fig. 167). Venter and sternum as in female (Fig. 168). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.1. Anterior median eyes separated by 1.3 diameters, posterior median eyes by 0.8, anterior median eyes separated from anterior laterals by 1.9 di- ameters of anterior lateral eyes, lateral eyes almost touching. Total length 3.7 mm. Carapace 1.9 mm long, 1.4 wide. First fe- mur 2.9 mm, patella and tibia 2.9, meta- tarsus 2.7, tarsus 1. Second patella and tib- ia 2.4 mm, third 1.3, fourth 1.9. Diagnosis. Unlike others in the M. in- crassata species group, M. inca and M. maya both have distal embolic apophyses that project forward, forming rounded bumps (Figs. 157, 164). The male palps of these two species are separated by the thinner flagellae and base on the median apophysis of M. inca (compare Fig. 164 with Fig. 157). The epigynum of M. inca resembles those of M. maya and M. com- anche because the oval epigynal openings are formed out of membranous surfaces that are distinctly separate from the base of the scape (Figs. 159, 166, 187). The epi- gynal openings of M. inca (Fig. 166) are much wider than those of M. comanche (Fig. 187); the posterior lobes on M. inca are pointed (Fig. 166), not just thickened, as in M. maya (Fig. 159). Variation. Average body length of six fe- males examined 5.7 mm, range 5.2 to 6.8 mm. Average body length of four males examined 3.7 mm, range 2.8 to 4.4 mm. Coloration varies significantly among lo- calities, especially in the males, where some are frequently much darker than others. Natural History. Mature specimens have been collected in April through Oc- tober (Fig. 318) at elevations between 300 and 600 m. Distribution. Most northern tip of Peru (Map 11). Records Examined. PERU Piura: 12 mi N Man- cora, 4°0'S, 80°54'W, 11.xii.1938; Parifias Valley, 4°30'S, 81°8'’W, 3.iv.1939, 8.iv.1939, 16.iv.1939, 7.V.1939, 21.v.1939, 25.vi.1939, 3.vii.1939, 6.viii. 1939, 15.viii. 1939, 15.x.1939; Quebrada de Parifias, 4°32'S, 81°17'W, 14.iv.1939, 7.v.1939, 21.v.1939; Quebrada Mogollon, 4°32'S, 81°4’W, 30.iv.1939, 11.vi.1939, 18.vi.1939, 21.vi.1939, 11.vii.1939, 16.vii.1939, 24.ix.1939 (all records: D. L. & H. E. Frizzell, CAS). 21. Metepeira gosoga Chamberlin and Ivie Figures 171-177, 322; Map 9 Metepeira gosoga Chamberlin and Ivie, 1935: 21 figs 82-83, 2. Female holotype from Pilot Knob Valley, Mohave Desert, California, in the AMNH, exam- ined. Roewer, 1942: 868. Bonnet, 1957: 2820. Levi, 1977: 200, figs. 28-36. Description. Female from Baja Califor- nia Norte, Mexico. Carapace dirty yellow- ish brown, lighter anterior half, darker di- amond-shaped mark behind eyes (Fig. 176). Legs same color as carapace; dark rings on distal ends of articles. Femur I with row of three to five macrosetae on anterior side; two or three macrosetae on anteroventral side. Folium lighter than in other species, darkening posteriorly (Fig. 176). Venter black surrounded by yellow. Wide white median line with pair of large white spots on either side of spiracle. Sometimes with parallel pair of lateral white lines that join transverse white line posteriorly (Fig. 177). Sternum black with wide, white line widening anteriorly, often METEPEIRA ° Piel 59 broken in center (Fig. 177). Ratio of eye diameters: posterior medians and anterior medians 0.9, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separat- ed by 1.2 diameters, posterior median eyes by 0.6, anterior median eyes separated from anterior laterals by 3.4 diameters of anterior lateral eyes, lateral eyes separated by 0.4 their diameters. Total length 8.3 mm. Carapace 4 mm long, 3.3 wide. First femur 4.5 mm, patella and tibia 4.8, meta- tarsus 4.5, tarsus 1.3. Second patella and tibia 4.3 mm, third 2.5, fourth 3.8. Male from Baja California Norte, Mex- ico. Male carapace, dorsum, venter, ster- num as in female (Figs. 174, 175). Femur I with row of five macrosetae on anterior side: row of nine macrosetae on anterov- entral side. Ratio of eye diameters: poste- rior medians and anterior medians 0.8, an- terior laterals 1.1, posterior laterals 1.3. Anterior median eyes separated by 1 di- ameter, posterior median eyes by 0.8, an- terior median eyes separated from anterior laterals by 2.1 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their di- ameters. Total length 5.5 mm. Carapace 2.8 mm long, 2.3 wide. First femur 4.5 mm, patella and tibia 4.5, metatarsus 4.5, tarsus 1.4. Second patella and tibia 3.8 mm, third 1.9, fourth 3. Diagnosis. Within the M. incrassata spe- cies group, only M. gosoga, M. inca, and M. maya have projecting distal embolic apophyses (Figs. 157, 164, 171). Of these, only M. gosoga has a distinctly pointed one (Fig. 171). The epigynum of M. gosoga dif- fers from those of other species in the M. incrassata group by the small, oval shape of the openings (Fig. 173). Variation. Average body length of two females examined 6.8 mm, range 5 to 8.5 mm. Average body length of two males ex- amined 4.9 mm, range 4.1 to 5.7 mm. Natural History. Mature specimens have been collected in June through Au- gust (Fig. 322) between 500 and 2,000 m. Levi (1977) notes that these have been collected on Opuntia and desert vegeta- tion. 60 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Distribution. Southwestern U.S. and Baja California Norte, Mexico (Levi, 1977, map 1; Map 9). Records Examined. MEXICO Baja Calif. Norte: 23 mi S Catavina, 29°32'N, 114°57'W, 20.vi.1983 (L. Strange & R. Miller, FSCA). USA Arizona: Show Low Lake, 34°12'’N, 110°O’W, 15.viii.1966 (MCZ). 22. Metepeira olmec new species Figures 178-184, 324; Map 11 Holotype. Female from Fortin de las Flores, Vera- cruz, Mexico, 26.vi.1944, L. I. Davis, in AMNH. The specific name is a noun in apposition after the ancient Indian people of southern Veracruz and Ta- basco. Description. Female holotype. Carapace reddish brown with white region around eyes that extends posteriorly along lateral margins (Fig. 183). Legs ringed on distal ends of articles. Femur I with row of four macrosetae on anterior side; none on an- teroventral side. Dorsum of abdomen with usual Metepeira folium (Fig. 183). Venter of abdomen with wide, long white median line set within surrounding U-shaped markings. Pair of white spots on either side of spiracle (Fig. 184). Sternum has wide median white line widening anteri- orly with broken constriction in center (Fig. 184). Ratio of eye diameters: poste- rior medians and anterior medians 1.0, an- terior laterals 1.1, posterior laterals 1.5. Anterior median eyes separated by 1.5 di- ameters, posterior median eyes by 0.8, an- terior median eyes separated from anterior laterals by 2.1 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their di- ameters. Total length 5.9 mm. Carapace 2.8 mm long, 2 wide. First femur 2.8 mm, patella and tibia 2.8, metatarsus 2.5, tarsus 1. Second patella and tibia 2.5 mm, third eS tourtiar 23. Male paratype from Fortin de las Flo- res, Veracruz, Mexico. Carapace dark with light region around eyes and a light mark anterior to thoracic furrow (Fig. 181). Legs darker than in female. Macrosetae on legs variable with spider’s size—femur I with row of three or four macrosetae on anterior side; one to three on anteroven- tral side. Dorsum, venter, sternum as in female, except median white line on ven- ter shortened to spot and median white line on sternum often broken (Figs. 181, 182). Ratio of eye diameters: posterior me- dians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separated by 1.3 diameters, posterior median eyes by 0.8, anterior me- dian eyes separated from anterior laterals by 1.3 diameters of anterior lateral eyes, lateral eyes separated by 0.1 their diame- ters. Total length 2.8 mm. Carapace 1.4 mm long, 1.2 wide. First femur 1.7 mm, patella and tibia 1.6, metatarsus 1.1, tarsus 0.7. Second patella and tibia 1.3 mm, third 0.8, fourth 1.1. Diagnosis. Female M. olmec are diag- nosed by their almost perfectly round epi- gynal depressions with uniform thickness around the posterior edges (Fig. 180). The male embolus is relatively short and has a vestigial distal embolic apophysis that does not project forward, but instead drops off, forming a sharp curve (Fig. 178). Among other species in the M. incrassata species group, this embolus shape is also seen in M. comanche and M. triangularis: unlike M. comanche (Fig. 185), the darker, scler- otized portion of the embolus does not ex- tend over the hump of the distal embolic apophysis; unlike M. triangularis (Fig. 199), the flagellae are thicker than the base of the median apophysis. Variation. Average body length of four females examined 6.2 mm, range 5.6 to 6.7 mm. Average body length of three males examined 2.3 mm, range 2.2 to 2.6 mm. Natural History. Mature specimens have been collected perennially (Fig. 324) at elevations between 500 and 1,400 m. Distribution. Montane rain forests from Veracruz to Panama (Map 11). Records Examined. COSTA RICA San José: San Antonio de Escanza, 9°59'N, 84°11'W [?], 4.iii.1984 (W. Eberhard, MCZ). MEXICO Veracruz: Fortin de las Flores, 18°54'N, 97°0’W, 26.vi.1944 (L. I. Davis, AMNH)); Papantla, 20°27'N, 97°19’W, 12.x.1947 (H. Wagner, AMNH). PANAMA Chiriqut: Boquete, 8°47'N, 82°26’W, 15.viii.1950 (A. M. Chickering, METEPEIRA ® Piel 61 7 gosoga comanche (23) Figures 171-177. Metepeira gosoga Chamberlin and Ivie (sp. 21; 29°32'N, 114°57’W). 171, male palpus, mesal. 172, epigynum, posterior. 173, epigynum, ventral. 174, male, dorsal. 175, male, ventral. 176, female, dorsal. 177, female, ventral. Figures 178-184. Metepeira olmec new species (sp. 22; 18°54'N, 97°0’W). 178, male palpus, mesal. 179, epigynum, posterior. 180, epigynum, ventral. 181, male, dorsal. 182, male, ventral. 183, female, dorsal. 184, female, ventral. Figures 185-191. Metepeira comanche Levi (sp. 23 [185,188,189] 33°22’N, 99°56’W; [186,187] 25°45’/N, 101°55’W; [190, 191] 26°41'N, 101°23’W). 185, male palpus, mesal. 186, epigynum, posterior. 187, epigynum, ventral. 188, male, dorsal. 189, male, ventral. 190, female, dorsal. 191, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 62 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 MCZ), 7.viii.1954 (A. M. Chickering, MCZ). Panama: Cerro Galera, 8°55'N, 79°38'W, 7.i.1977 (H. W. Levi & Y. Lubin, MCZ). 23. Metepeira comanche Levi Figures 185-191, 331; Map 11 Metepeira comanche Levi, 1977: 204, figs. 61-69, ¢, 2. Male holotype from 9.7 km west of O’Brien, Haskell Co., Texas, in the MCZ, examined. Brig- noli, 1983: 275. Description. Female from Gloria, Coa- huila, Mexico. Carapace dark with light re- gion around eyes, extending posteriorly behind posterior lateral eyes (Fig. 190). Legs ringed. Femur I with row of four ma- crosetae on anterior side; one or two on anteroventral side. Dorsal folium high in contrast; black comma-shaped markings shadow white fleur-de-lis pattern (Fig. 190). Venter of abdomen with wide white median line posteriorly set within sur- rounding U-shaped marking. Pair of white spots on either side of spiracle (Fig. 191). Sternum has wide median white line wid- ening anteriorly (Fig. 191). Ratio of eye diameters: posterior medians and anterior medians 0.9, anterior laterals 1.2, posterior laterals 1.1. Anterior median eyes separat- ed by 1.2 diameters, posterior median eyes by 1.0, anterior median eyes separated from anterior laterals by 2.7 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 7.8 mm. Carapace 3.3 mm long, 2.5 wide. First femur 3.6 mm, patella and tibia 3.9, metatarsus 3.4, tarsus 1.2. Second patella and tibia 3.3 mm, third 2, fourth 2.9. Male from 9.7 km west of O’Brien, Has- kell Co., Texas. Carapace light around eyes, darker posteriorly (Fig. 188). Legs ringed. Macrosetae on femur I variable; usually row of four macrosetae on anterior side, six on anteroventral side. Dorsum with usual Metepeira folium (Fig. 188). Venter with median oval white mark; pair of white spots on either side of spiracle (Fig. 189). Median white line on sternum usually broken (Fig. 189). Ratio of eye di- ameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separat- ed by 1.4 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 1.6 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 3.1 mm. Carapace 1.7 mm long, 1.3 wide. First femur 2.4 mm, patella and tibia 2.2, metatarsus 2, tarsus 0.9. Second patella and tibia 1.9 mm, third 1.0, fourth 1.5. Diagnosis. The epigynum of M. com- anche resembles those of M. maya and M. inca because the oval epigynal openings are formed out of membranous surfaces that are distinctly separate from the base of the scape (Figs. 159, 166, 187). The epi- gynum of M. comanche differs from M. maya and M. inca by having much narrow- er epigynal openings (compare Fig. 187 with Figs. 159, 166). The male embolus is relatively short and has a vestigial distal embolic apophysis that does not project forward, but instead drops off, forming a sharp curve (Fig. 185). Among other spe- cies in the M. incrassata species group, this embolus shape is also seen in M. olmec and M. triangularis: unlike M. olmec (Fig. 178), the darker, sclerotized portion of the embolus extends over the hump of the dis- tal embolic apophysis; unlike M. triangu- laris (Fig. 199), the flagellae and base of the median apophysis are wide (Fig. 185). Natural History. Mature specimens have been collected from May to Novem- ber (Levi, 1977, Fig. 331). Distribution. New Mexico, Texas, and northern Mexico (Levi, 1977, map 1; Map LIEN, Records Examined. MEXICO Coahuila: Gloria, 26°41'N, 101°23'W, 24.viii.1947 (W. J. Gertsch, AMNH); Paila, 25°45’N, 101°55’W, 21.viii.1947 (W. J. Gertsch, AMNH). 24. Metepeira pimungan new species Figures 192-198, 330; Map 9 Holotype. Male from San Miguel Island, Santa Bar- bara Co., Califoria, USA, 20.vii.1968, M. E. Thompson, in the MCZ. The specific name is a noun in apposition after the Indian people who once lived on the Channel Islands. Description. Female paratype from San Miguel Island, California USA. Carapace with large white eye region and lateral posterior extensions. White median arrow- shaped mark extends to thoracic furrow (Fig. 197). Legs mostly white with rings on legs III and IV. Femur I with row of four macrosetae on anterior side; one on anter- oventral side. Dorsum of abdomen with usual Metepeira folium (Fig. 197); venter wide, with long white median line, flanked by shorter parallel thin white lines on ei- ther side. Pair of white spots on either side of spiracle. Sternum black with posterior drop-shaped white mark (Fig. 198). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.4, posterior laterals 1.3. Anterior median eyes separated by 1.6 diameters, posterior median eyes by 1.0, anterior median eyes separated from anterior laterals by 3.4 di- ameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. To- tal length 6.9 mm. Carapace 3.3 mm long, 2.6 wide. First femur 3.5 mm, patella and tibia 3.5, metatarsus 2.8, tarsus 1.3. Sec- ond patella and tibia 3 mm, third 1.9, fourth 2.8. Male holotype. Carapace, legs, abdo- men, venter, sternum as in female, though legs darker and not ringed (Figs. 195, 196). Femur I with row of four to five ma- crosetae on anterior side; four to five on anteroventral side: both rows concentrated distally on femur. Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separated by 1.6 diameters, posterior median eyes by 0.8, anterior median eyes separated from an- terior laterals by 1.9 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 4.5 mm. Car- apace 2.3 mm long, 1.8 wide. First femur 3.1 mm, patella and tibia 3, metatarsus 2.8, tarsus 1. Second patella and tibia 2.5 mm, third 1.3, fourth 1.9. Diagnosis. Like M. atascadero, but un- like other species in the M. incrassata group, the posterior end of the sternum of METEPEIRBA ® Piel 63 M. pimungan has a white water drop mark (Figs. 196, 198). Unlike M. atascadero, M. pimungan lacks U-shaped white markings on the venter, just anterior to the spiracle (compare Fig. 198 with Fig. 221). In con- trast to other species in the M. incrassata group, the embolus on male M. pimungan specimens is curved to form a gentle S- shape (Fig. 192). The female has a squar- ish-shaped black mark inside each epigyn- al depression (Fig. 198) which is not seen in other M. incrassata group species. Variation. Average body length of four females examined 7.7 mm, range 6.9 to 8.5 mm. Average body length of five males ex- amined 5.3 mm, range 4.6 to 6.4 mm. Natural History. Mature specimens have been collected in August (Fig. 330). Webs are found near the ground, shel- tered from the perennial strong winds by Coryopsis and lupines. Distribution. Endemic to San Miguel Island (Map 9). Records Examined. USA California: San Miguel Is- land, 34°2’28.6"N, 120°21'13.6"W, 11.viii.1995 (W. H. Piel, MCZ); 34°2'6.1"N, 120°21'9.1"W, 12.viii.1995 (W. H. Piel, MCZ); 34°2’N, 120°22’W, 20.viii.1968 (M. E. Thompson, MCZ); 34°3'3.1"N, 120°21'53.3"W, 13.viii.1995 (W. H. Piel, MCZ): behind dunes on beach, 34°2'43.8"N, 120°21'0.8"W, 11.viii.1995 (W. H. Piel, MCZ). 25. Metepeira triangularis (Franganillo) Figures 199-205, 337; Map 14 Mangora triangularis Franganillo, 1930: 21-22. Fe- male holotype from Sierra Maestra, lost. Platnick, 1993: 449. Metepeira labyrinthea:—Franganillo, 1936: 75. Bry- ant, 1940: 341. Platnick, 1993: 449. Metepeira triangularis:—Archer, 1958: 15, fig. 37, @. Metepeira acostai Archer, 1958: 15, fig. 36, 2. Female holotype from the savannas, Agramont, Camagiiey Province, Cuba, in the AMNH, examined. NEW SYNONYMY. Metepeira bani Archer, 1965: 132, figs. 11, 17 2, 13 3. Male and female syntypes from Bani, Domini- can Republic, in the AMNH, examined. NEW SYNONYMY. Brignoli, 1983: 275. Note. Franganillo’s collection vials are not la- beled, making it impossible to find the holotype. Based on the descriptions by Franganillo (1930), Archer (1958), the illustrations of Archer (1958, 1965), and material determined by Archer, I con- clude that there is no evidence to support three 64 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 separate species. For the name M. triangularis, I designate a female neotype from Camagiiey Prov- ince, Cuba. 21°30'N, 78°10'W, x.1954 (J. T. Acosta, AMNBH). Description. Female from La Descu- bierta, Lago Enriquillo, Isla Cabritos, Do- minican Republic. Light reddish brown carapace, white around eyes (Fig. 204). Legs yellowish white, sometimes ringed brown at distal ends of articles. Femur I with row of three, or sometimes four, ma- crosetae on anterior side; none on anter- oventral side. Dorsal folium white with slightly darker markings posteriorly (Fig. 204). In some cases, dark anterior marks on shoulders of dorsum. Venter of abdo- men black with wide white median line, sometimes flanked by thinner white lines that together form an anchor shape (Fig. 205). Pair of small white spots on either side of spiracle. Sternum black with me- dian white line, wider anteriorly (Fig. 205). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.4, posterior laterals 1.4. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 1.1, anterior median eyes separated from anterior laterals by 3.3 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 5.9 mm. Carapace 2.7 mm long, 2 wide. First femur 2.6 mm, patella and tibia 2.9, metatarsus 2.5, tarsus 1. Second patella and tibia 2.4 mm, third 1.4, fourth DAD Male from La Descubierta, Lago Enri- quillo, Isla Cabritos, Dominican Republic. Carapace as in female, though lighter eye region often extends to cover entire ante- rior half (Fig. 202). Legs yellowish white. Distal half of femora often dark. Femur I with row of four macrosetae on anterior side; three to five on anteroventral side. Dorsal folium white with slightly darker markings posteriorly (Fig. 202). Venter of abdomen as in female; sternum black with median white line (Fig. 203). Ratio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 0.7, posterior laterals 1.1. Anterior median eyes separat- ed by 1.6 diameters, posterior median eyes by 0.8, anterior median eyes separated from anterior laterals by 1.0 diameters of anterior lateral eyes, lateral eyes separated by 0.1 their diameters. Total length 4.2 mm. Carapace 2 mm long, 1.5 wide. First femur 2.9 mm, patella and tibia 3, meta- tarsus 2.8, tarsus 1.1. Second patella and tibia 2.4 mm, third 0.8, fourth 1.3. Diagnosis. The epigynum of M. trian- gularis differs from other species by the thin, sclerotized openings in the shape of Ray-Ban sunglasses (Fig. 201). The male embolus has a vestigial distal embolic apophysis that does not project forward, but instead drops off, forming a sharp curve (Fig. 199). Among other species in the M. incrassata species group, shorter versions of this embolus shape is also seen in M. comanche and M. olmec: unlike M. comanche (Fig. 185), the darker, sclero- tized portion of the embolus does not ex- tend over the hump of the distal embolic apophysis; unlike M. olmec (Fig. 178), the flagellae and base of the median apophysis are much thinner (Fig. 199). Variation. Average body length of seven females examined 5.5 mm, range 4.6 to 6.3 mm. Average body length of five males ex- amined 3.8 mm, range 3.2 to 4.3 mm. Natural History. Mature specimens have been collected from June to Febru- ary, although I suspect that like other Ca- ribbean species, M. triangularis is actually entirely perennial (Fig. 337). Distribution. Dominican Republic and Cuba (Map 14). Records Examined. CUBA Camagtiey Prov.: 21°30'N, 78°L0’W, x.1954 (J. T. Acosta, AMNH). Ma- tanzas: Matanzas, Parque Watkins, 23°3'N, 81°35’W, 9.viii.1955 (A. F. Archer, AMNH); Pan de Palenque [?], 23°1'N, 81°43’W, 11.viii.1955 (A. F. Archer, AMNH). Oriente: Banes, 20°58'’N, 75°43'W, 2.viii.1955 (A. F. Archer, AMNH); Cuabitas, Santiago, 20°4'N, 75°48'W, 15.xii.1955 (P. Alayo, AMNH): Puerto Boniato, 20°7'N, 75°47'W, 4.xi.1945 (P. Alayo, AMNH). Sierra las Casas: Isla de Pinos, 21°53'N, 82°48’W, 17.viii. 1955 (A. F. Archer, AMNH). DO- MINICAN REPUBLIC Azua: Hatillo, 18°24’'N, 70°32'W, 30.i1.1991 (Felix E. Del Monte, CAS). Ban- ahona: Sierra Martin Garcia [?], 18°25’N, 70°30'W, 8.viii.1958 (A. F. Archer & E. de Boyrie Moya, METEPEIRA ° Piel 65 =. (= 5%) a 30 4 203 triangularis (25) Dios 209 mj arizonica (26) Figures 192-198. Metepeira pimungan new species (sp. 24; 34°2’N, 120°22’W). 192, male palpus, mesal. 193, epigynum, posterior. 194, epigynum, ventral. 195, male, dorsal. 196, male, ventral. 197, female, dorsal. 198, female, ventral. Figures 199-205. Metepeira triangularis (Franganillo) (sp. 25; 18°34'N, 71°44’W). 199, male palpus, mesal. 200, epigynum, posterior. 201, epigynum, ventral. 202, male, dorsal. 203, male, ventral. 204, female, dorsal. 205, female, ventral. Figures 206-212. Metepeira arizonica Chamberlin and Ivie (sp. 26; 28°53'N, 113°4’W). 206, male palpus, mesal. 207, epigynum, posterior. 208, epigynum, ventral. 209, male, dorsal. 210, male, ventral. 211, female, dorsal. 212, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 206 66 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 AMNRH). La Independencia: 2 km SE EI Limon, Ji- mani. S side of road, 18°25'’N, 71°45'’W, 30.i1.1991 (Felix E. Del Monte, MNSD); Isla Cabritos: La Des- cubierta, Lago Enriquillo, 18°34’N, 71°44’W, 18.vi.1981 (E. Marcano, MNSD). Montecristi: Car- retera Montecristi, El Morro [?], 19°54'N, 71°39'W, 2.xii.1991 (Felix E. Del Monte, MNSD), 21.xii.1991 (Felix E. Del Monte, MNSD). Peravia: Las Dunas, Prov. Bani [?], 18°25'N, 71°25'W, 23.i.1992 (Felix E. Del Monte, MNSD). Prov. Trujillo Valdez: W_ of Bani, 18°17'N, 70°22’W, 8.viii.1958 (A. F. Archer, AMNH). San Juan: 1 km S Las Matas de Farfan, 18°51'N, 71°31'W, 25.viii.1970 (B. Patterson, MCZ). 26. Metepeira arizonica Chamberlin and Ivie Figures 206-213, 319; Map 9 Metepeira arizonica Chamberlin and Ivie, 1942: 69, figs. 182-187, 2, 3d. Female holotype from Canyon Lake, Arizona, in the AMNH, examined. Levi, 1977: 200, figs. 12, 13, 39-46, 2, 3. Brignoli, 1983: OS Description. Female from Isla Partida, Baja Calif. Norte, Mexico. General color- ation high in contrast. Carapace dark brown; large white eye region, white lat- eral posterior extensions, and short median posterior extension (Fig. 211). Distinct black rings on all legs. Femur I with row of four macrosetae on anterior side; four on anteroventral side. Dorsum of abdo- men with usual Metepeira folium, though generally slightly lighter (Fig. 211); venter wide, with long white median line set within surrounding U-shaped markings. Pair of white spots on either side of spi- racle (Fig. 212). Wide median white line widening anteriorly with constriction in center, set on black sternum (Fig. 212). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.4, posterior laterals 1.2. Anterior median eyes separated by 1.4 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 2.7 di- ameters of anterior lateral eyes, lateral eyes separated by 0.1 their diameters. To- tal length 6.2 mm. Carapace 2.8 mm long, 2.2 wide. First femur 3.4 mm, patella and tibia 3.5, metatarsus 3, tarsus 1.2. Second patella and tibia 3 mm, third 1.9, fourth 2.8. Male from Isla Partida, Baja Calif. Nor- te, Mexico. Carapace, legs, abdomen, ven- ter, sternum as in female, though legs darker with less distinct rings, and white line on sternum often broken (Figs. 209, 210). Femur I with row of four macrosetae on anterior side; five to eight on anterov- entral side. Ratio of eye diameters: poste- rior medians and anterior medians 1.1, an- terior laterals 1.4, posterior laterals 1.2. Anterior median eyes separated by 1.5 di- ameters, posterior median eyes by 0.7, an- terior median eyes separated from anterior laterals by 1.6 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their di- ameters. Total length 4.4 mm. Carapace 2.3 mm long, 1.8 wide. First femur 3.3 mm, patella and tibia 3.3, metatarsus 3.2, tarsus 1.2. Second patella and tibia 2.8 mm, third 1.5, fourth 2.2. Diagnosis. Unlike all other species in the M. incrassata group, the epigynum of M. arizonica is puffy and swollen, causing the openings to reduce (from a ventral view) to crescent-shaped slits (Fig. 187). The embolus differs from others in the M. incrassata group because the distal em- bolic apophysis is reduced to a gentle arch- ing curve (Fig. 206). Variation. Average body length of nine females examined 5.5 mm, range 4.7 to 6.3 mm. Average body length of 13 males ex- amined 3.9 mm, range 3.1 to 5.1 mm. In some males a small ventral keel is visible extending beyond the flagella of the me- dian apophysis (Levi, 1977, fig. 46). Natural History. Mature specimens are most commonly collected between April and August (Fig. 319), from dry oak—pine— juniper woodland, alfalfa, cactus, and de- sert shrub (Levi, 1977). Localities have a large range in altitude, from sea level to 2,300 m; however, spiders from southern regions live at higher elevations than spi- ders from northern regions (Fig. 213). Distribution. American southwest to San Luis Potosi (Levi, 1977, map 1; Map 9). Records Examined. MEXICO Baja Calif. Norte: 15 Estimated Elevation (meters) 22 24 26 28 30 32 34 Latitude (degrees north) Figure 213. The elevation of collection localities for M. arizon- ica at their corresponding latitudes. Species-specific altitudes appear to decrease with distance from the equator. Elevations estimated from NOAA database of 5- by 5-minute geographic tiles. mi S Punta Prieta, 28°58'N, 114°17'W, 7.vii.1973 (S. C. Williams & K. B. Blair, CAS); 24 mi S Santa Inés, 29°20'N, 114°20'W, 7.vii.1973 (S. C. Williams & K. B. Blair, CAS); 26 mi S San Felipe, 30°38'N, 114°50’W, 15.iv.1965 (D. Q. Cavagnaro, C. E. & E. S. Ross, V. L. Vesterby, CAS); 6 km NW Racho Santa Inés, 29°43'N, 114°43’W, 28.iii.1981 (Paul E. Blom, WHC); 6 mi S San Felipe, 30°55'N, 114°52’W, 14.iv.1965 (D. Q. Cavagnaro, C. E. & E. S. Ross, V. L. Vesterby, CAS); 9 km NW Racho Santa Inés, 29°46'N, 114°46’W, 3.vii.1981 (David Crowe, WHC): Arroyo de Calamajué at carbonated spring, 29°38’N, 114°25'W, 26.vi.1973 (S. C. Williams & K. B. Blair, CAS); Cafion de Guadalupe, off Laguna Salada, 32°10'N, 115°48’'W, 11.1.1958 (V. Roth, AMNH); El Mayor, 32°5'N, 115°13’W, 15.vi.1952 (M. Cazier, W. Gertsch, & R. Schrammel, AMNH); Isla Mejia, 29°34'N, 113°35’W, 30.iv.1921 (J. C. Chamberlin, MCZ); Isla Partida, 28°53’N, 113°4’W, 2.vii.1921 GE C. Chamberlin, MCZ); Isla San Lorenzo, N end, 28°40'N, 112°52'W, 24.vi.1921 (J. C. Chamberlin, CAS); Isla San Lorenzo, south Tobart, 28°36'N, 112°46’W [P], 9.iv.1921 (J. C. Chamberlin, CAS); Puerto Refugio, N end Angel de la Guarda Isl. Webs in cracks in cliff overhanging beach, 29°34'N, 113°32'W, 18.iv.1962 (Howard W. Campbell, CAS); San Pedro Martir, 30°45’N, 115°13'W, 18.iv.1921 (J. C. Chamberlin, CAS). Coahuila: 25 mi SE San Pedro, 25°35'N, 102°50'W, 21.viii.1947 (W. J. Gertsch, AMNH): Cabos, 25°35'N, 101°43’W, 21.viii.1947 (W. J. Gertsch, AMNH); Gloria, 26°41’N, 101°23’W, 24.viii.1947 (W. J. Gertsch, AMNH); Saltillo, 25°25'N, 101°0'W, 23.v.1952 (M. Cazier, W. Gertsch, & R. Schrammel, AMNH). Durango: La Loma, 25°27'N, 103°40'W, 20.viii.1947 (W. J. Gertsch, AMNH); Providencia, 26°44'N, 105°56'W, 24 viii.1947 (A. M. Davis, AMNH); San Isidro, 60 mi NW Durango, 25°1'’N, 105°6'W, 19.viii.1947 (W. J. Gertsch, AMNH). San Luis Potost: 3 km W. Pilares, 21°55'34"N, 100°48’6"W, 21.x.1994 (W. H. Piel, MCZ). Sinaloa: Las Saleras Is. (= Isla Saliaca [?]), 25°11'N, 108°20’W [?], 13.vi.1921 (J. C. Chamberlin, METEPEIRA ® Piel 67 CAS). Sonora: 20 mi N Hermosillo, 29°8’N, 110°58'W, 13.ix.1966 (Jean & Wilton Ivie, AMNH): 20 mi SW Sonoyta, 31°38'N, 113°4'W, 13.vi.1952 (W. J. Gertsch, AMNH); 22 mi E Hermosillo on the banks of Rio Sonora, 29°4'’N, 110°55’W, 17.viii.1959 (B. A. Branson, AMNH); Los Angeles, 29°27'N, 110°46'W, 4.x.1966 (V. Roth, AMNH): Puerto Pef- asco at seashore, 31°20'N, 113°33’W, 3.iv.1968 (D. E. Bixler, MCZ). Zacatecas: 14 mi S Fresnillo, 23°5’'N, 102°45'W, 4.vili.1954 (W. J. Gertsch, AMNH);: 4 mi NE Concepcién del Oro, 24°41’N, 101°23'W, 4.vii.1984 (J. B. Woolley, AD). USA Arizona: Santa Catalina Mtns., mile 7.9 of Catalina Highway from Tuscon to Mt. Lemmon., 32°17’N, 110°48’W, 19.vi.1985 (W. Maddison, MCZ): Tucson, 32°15’'N, 110°57'W, 5.vii.1991 (W. H. Piel & G. S. Bodner, MCZ). California: Corn Springs, Chuckawalla Mtns., 10 mi SE Desert Center, 33°33’N, 115°24’W, 29.xi.1963 (D. C. Lorn, MCZ); Manzanita chaparral, San Gabriel Canyon, Coldbrook Ranger Station, B34°11'N, 117°53'W, 19.viii.1964 (L. Pinter, MCZ). Texas: Big Bend Nat'l Park, Old Ranch House on St. Elena Rd., 29°10’N, 103°30’W, 25.v.1967 (E. Sabath, MCZ). 27. Metepeira atascadero new species Figures 214-221, 336; Map 14 Holotype. Male from San Miguel de Allende, Guana- juato, Mexico, 25.x.1982, George Uetz, in MCZ. The specific name is a noun in apposition after the Rancho Hotel Atascadero in San Miguel de Allen- de. This name was coined by George Uetz when he and his colleagues stayed in Rancho Hotel Atas- cadero while studying the behavioral ecology of this species. This name has been in informal use in the literature (e.g., Uetz and Hodge, 1990). Description. Female paratype from San Miguel de Allende, Guanajuato, Mexico. Reddish brown carapace, white around eyes, faint light marks extend posteriorly behind lateral eyes (Fig. 220). Legs yel- lowish, ringed brown at distal ends of ar- ticles. Femur I with row of four macro- setae on anterior side; three on anterov- entral side. Largest branches of fleur-de- lis pattern on dorsal folium form large paired white spots (Fig. 220). Venter of ab- domen black with wide white median line, flanked by white U-shape mark (Fig. 221). Pair of small white spots on either side of spiracle. Sternum black with white poste- rior mark (Fig. 221). Ratio of eye diame- ters: posterior medians and anterior me- dians 1.0, anterior laterals 1.6, posterior laterals 1.3. Anterior median eyes separat- 68 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 ed by 1.4 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 3.7 diameters of anterior lateral eyes, lateral eyes separated by 0.4 their diameters. Total length 7.5 mm. Carapace 3.1 mm long, 2.5 wide. First femur 4 mm, patella and tibia 4, metatarsus 3.9, tarsus 1.3. Second patella and tibia 3.3 mm, third 1.8, fourth 2.8. Male holotype. Reddish brown cara- pace, lighter around eyes, faint light marks extend posteriorly behind lateral eyes and median eyes (Fig. 218). Legs yellowish white, each article gradually turning red- dish brown distally. Femur I with row of four or five macrosetae on anterior side; eight or nine on anteroventral side. Larg- est branches of fleur-de-lis pattern on dor- sal folium form large paired white spots (Fig. 218). Venter of abdomen black with wide white median line, flanked by faint white U-shape mark (Fig. 219). Pair of small white spots on either side of spiracle. Sternum black with white posterior mark (Fig. 219). Ratio of eye diameters: poste- rior medians and anterior medians 1.0, an- terior laterals 1.7, posterior laterals 1.4. Anterior median eyes separated by 1.8 di- ameters, posterior median eyes by 1.0, an- terior median eyes separated from anterior laterals by 3.6 diameters of anterior lateral eyes, lateral eyes separated by 0.5 their di- ameters. Total length 5 mm. Carapace 2.5 mm long, 1.8 wide. First femur 4 mm, pa- tella and tibia 4.2, metatarsus 4.5, tarsus 1.4. Second patella and tibia 3.3 mm, third les founth: 25: Diagnosis. The male embolus of M. atascadero has a protruding bump _ be- tween the embolus tip and the basal em- bolic apophysis (Figs. 214, 217) which sep- arates it from all other Metepeira species. As with M. incrasssata (Fig. 224) and, to a lesser degree, M. triangularis (Fig. 201), the epigynal depressions on either side of the scape of M. atascadero (Fig. 216) are sclerotized to create a scooped and slick quality. The darker epigynal openings in- side the depressions are hidden anteriorly under the scape’s hood. Metepeira atas- cadero’s epigynum differs from that of M. incrassata by its much smaller depressions (compare Fig. 216 with 224). Variation. Average body length of 12 fe- males examined 7.9 mm, range 6 to 9.5 mm. Average body length of 12 males ex- amined 5.7 mm, range 3.5 to 6.9 mm. Natural History. Mature specimens have been collected from the end of Au- gust to the end of October (Fig. 336). This species lives solitarily: only 20% of spiders live in aggregations of two or more indi- viduals (Uetz and Hodge, 1990). It is thought that the low level of social behav- ior in M. atascadero occurs because the species pursues a risk-prone foraging strat- egy (Uetz, 1988a,b). Spiders are found be- tween 1,500 and 2,500 m elevation. Distribution. Mexican highlands from Durango to Guerrero (Map 14). Records Examined. MEXICO Coahuila: Saltillo, 14 mi E in Larrea Desert, 25°25’N, 100°55’W, 28.vii.1944 (AMNH). Durango: El Tascate, 26°12'N, 105°7'W, 27.vii.1947 (W. J. Gertsch, AMNH); Yer- banfs, 80 mi NW Durango, 24°45'N, 103°50'W, 19.viii. 1947 (W. J. Gertsch, AMNH). Guanajuato: 20 mi E. Guanajuato, 21°1’N, 100°57'W, 15.ix.1976 (C. E. Griswold & Jackson, CAS); Guanajuato, 21°1’N, 101°15'W (N. Banks, MCZ); San Miguel de Allende, 20°55'N, 100°45’W, 25.x.1982 (George Uetz, MCZ), 14.x.1983 (MCZ), 14.x.1985 (MCZ), 17.x.1985 (MCZ), 26.x.1985 (MCZ). Guerrero: Tecalpulco, 25 km N Iguala, 18°29'N, 99°38’W, 1.1.1948 (AMNH). Hidalgo: Ozumbilla, 20°9’N, 101°16’W, 2.x.1957 (R. Dreisbach, MCZ). Jalisco: Cyarco Onda, 30 km W Ojuelos, 21°47'N, 101°53’W (H. Wagner, AMNBH). Michoacan: Hills N of Patzcuaro, 19°45'N, 101°36’W, 24 viii.1959 (A. F. Archer, AMNH); Hwy 110, 4 mi W Jiquilpan, 19°59'N, 102°47'W, 2.viii.1967 (R. E. Leech, REL). Zacatecas: East of Guadalupe, 22°46'N, 102°31'W, 21.viii.1959 (A. F. Archer, AMNH): Guadalupe, 92°45'N, 102°31'W, 16.viii. 1947 (W. J. Gertsch, AMNH). 28. Metepeira incrassata F. O. P.-Cambridge Figures 222-228, 323; Map 9 Epeira Saléi Keyserling, 1864: 93, fig., ¢, 2. Female holotype from Oaxaca, Mexico, in BMNH, exam- ined. Keyserling, 1892: 196, fig. 145, 2. Roewer, 1942: 851. NEW SYNONYM Y. Metepeira incrassata F. O. P.-Cambridge, 1904: 460, fig. 11, 2. Female holotype from Jalapa, Mexico, in BMNH. Roewer, 1942: 868. Bonnet, 1957: 2821. METEPEIRA ® Piel 69 ventura (29) Figures 214-221. Metepeira atascadero new species (sp. 27; 20°55'N, 100°45’W). 214, male palpus, mesal. 215, epigynum, posterior. 216, epigynum, ventral. 217, male embolic division, ventral. 218, male, dorsal. 219, male, ventral. 220, female, dorsal. 221, female, ventral. Figures 222-228. Metepeira incrassata F. O. P.-Cambridge (sp. 28 [222,225—228] 19°4’N, 97°2’W; [223,224] 18°54’N, 97°0’W). 222, male palpus, mesal. 223, epigynum, posterior. 224, epigynum, ventral. 225, male, dorsal. 226, male, ventral. 227, female, dorsal. 228, female, ventral. Figures 229-235. Metepeira ventura Chamberlin and lIvie (sp. 29; 28°5’N, 114°8’W). 229, male palpus, mesal. 230, epigynum, posterior. 231, epigynum, ventral. 232, male, dorsal. 233, male, ventral. 234, female, dorsal. 235, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 70 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Aranea sallei:—F.O.P.-Cambridge, 1904: 519. Roew- er, 1942: 851. Araneus incrassata:—FPetrunkevitch, 1911: 289. Araneus sallei:—Petrunkevitch, 1911: 314. Metepeira salei:—Levi, 1991: 180. Platnick, 1993: 449. Note. Although M. incrassata is not the oldest name, it has been cited more than 20 times in the general literature (e.g., Caraco et al. 1995; Hieber and Uetz, 1990; Hodge and Uetz, 1992, 1995, 1996; Jakob et al., 1996; Rayor, 1996; Rayor and Uetz, 1990, 1993; Uetz, 1988a,b, 1989, 1991, 1992, 1996: Uetz and Hieber, 1994, 1997; Uetz and Hod- ge, 1990; Uetz et al., 1994). In contrast, the senior synonym has not been cited outside of taxonomic catalogues. In compliance with Article 79 of the ICZN (1985), I choose to assert the priority of the more popular junior name. Description. Female from Rancho Chu- la-Vista, Veracruz, Mexico. Brown cara- pace, darker around margins, lighter around eyes and behind lateral eyes (Fig. 227). Legs brown, yellow at base of articles and on articles distal to the femur for legs I and II. Femur I with row of three to four macrosetae on anterior side; one on anter- oventral side. Dorsal folium darker than most Metepeira species. White fleur-de-lis pattern with thin branches (Fig. 227). Ven- ter dark brown to black with white median mark that is shorter than it is in most other species (Fig. 228). Sternum dark brown to black with one or two small white spots, usually in the center or anteriorly (Fig. 228). Ratio of eye diameters: posterior me- dians and anterior medians 0.9, anterior laterals 1.3, posterior laterals 1.1. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 1.1, anterior me- dian eyes separated from anterior laterals by 4.1 diameters of anterior lateral eyes, lateral eyes separated by 0.5 their diame- ters. Total length 7.6 mm. Carapace 3.6 mm long, 2.9 wide. First femur 3.9 mm, patella and tibia 4.1, metatarsus 3.5, tarsus 1.4. Second patella and tibia 3.7 mm, third 2:3. fount ois. Male from Rancho Chula-Vista, Vera- cruz, Mexico. Carapace, dorsum, venter, sternum a darker version of female (Figs. 225, 226). Base of femora yellow, remain- der dark brown; other articles gradually turning lighter distally. Femur I with row of four to six macrosetae on anterior side; six to nine on anteroventral side. Ratio of eye diameters: posterior medians and an- terior medians 1.0, anterior laterals 1.4, posterior laterals 1.2. Anterior median eyes separated by 1.4 diameters, posterior median eyes by 1.0, anterior median eyes separated from anterior laterals by 3.7 di- ameters of anterior lateral eyes, lateral eyes separated by 0.4 their diameters. To- tal length 7 mm. Carapace 3.3 mm long, 2.7 wide. First femur 4.9 mm, patella and tibia 5.3, metatarsus 4.8, tarsus 1.7. Sec- ond patella and tibia 4.3 mm, third 2.2, fourth 3.4. Diagnosis. Overall pigmentation of M. incrassata is darker than all others in its species group. The thin branches on the dorsal folium (Figs. 225, 227), the almost entirely dark sternum, and the shortened median line on the venter (Figs. 226, 228) are distinctive. As with M. olmec (Fig. 178), M. comanche (Fig. 185), and M. tri- angularis (Fig. 199), the embolus of M. in- crassata (Fig. 222) curves off sharply from a distal embolic apophysis that does not project forward. Compared to M. olmec (Fig. 178) and M. comanche (Fig. 185), M. incrassata’s embolus tip beyond the distal apophysis is relatively longer and not as curved (Fig. 222). Male M. incrassata are easily separated from M. triangularis by the shape of the median apophysis (com- pare Fig. 222 with Fig. 199). As with M. atascadero (Fig. 216) and, to a lesser de- gree, M. triangularis (Fig. 201), the epi- gynal depressions on either side of the scape of M. incrassata (Fig. 224) are scler- otized to create a scooped and slick qual- ity. The darker epigynal openings inside the depressions are hidden anteriorly un- der the scape’s hood. Metepeira incrassa- ta’s epigynum differs from those of M. atascadero and M. triangularis by its much larger and disk-shaped depressions (com- pare Fig. 224 with Figs. 201, 216). Variation. Average body length of 31 fe- males examined 7.6 mm, range 6.4 to 9.1 mm. Average body length of 22 males ex- amined 6.2 mm, range 4.3 to 8 mm. Natural History. Mature specimens have been collected in February through October but are most frequently found in July (Fig. 323). This species lives socially: half of all spiders live in aggregations of 1,000 or more individuals (Uetz and Hod- ge, 1990). Colonies are frequently found spanning telephone lines, houses, fences, and other man-made structures. It is thought that this high level of social be- havior occurs because the species pursues a risk-averse foraging strategy in which the variance in survival attenuates with in- creasing colony size (Uetz, 1988a,b). Ele- vations center around 800 m and range from about 500 to 1,500 m. Distribution. Tropical Mexico from San Luis Potosi to southern Veracruz and Oa- xaca (Map 9). Records Examined. MEXICO Hidalgo: 20 mi S of Jacala, 20°50'N, 99°16’W, 18.iv.1946 (L. I. Davis & M. Johnston, AMNH); Chapulhuacan, 21°10’N, 98°54'’W, 20.v.1952 (M. Cazier, W. Gertsch, & R. Schrammel, AMNH), 27.vii.1966 (Jean & Wilton Ivie, AMNH), 16.vii.1969 (S. & J. Peck, MCZ). Méx- ico: 1 mi S Palomas, 19°50’N, 99°5’W, 28.vii.1950 (AMNH). Oaxaca: Oaxaca, 17°3'N, 96°43’W (Nathan Banks, MCZ). Puebla: Huauchinango, 20°11'N, 98°3'W, 7.x.1947 (H. Wagner, AMNH); north of Xic- otepec de Judrez, 20°18’N, 97°57'W, 19.iv.1967 (W. B. Peck, MCZ). San Luis Potost: 10km W Xilitla on r120, 21°22'N, 99°4'W, 10.viii.1991 (W. H. Piel & G. S. Bodner, MCZ). Veracruz: 1 mi SW Tlapacoyan, 19°57'N, 97°14’W, 16.vii.1973 (A. Newton, MCZ): 2 mi N Fortin de las Flores, 18°56'N, 97°1'W, 5.viii. 1966 (Jean & Wilton Ivie, AMNH): 5 mi E Ori- zaba, 18°51'N, 97°4’W, 25.vii.1956 (W. Gertsch & V. Roth, AMNH); Coatepec, 19°27’N, 96°58’W, 28.vii.1955 (C. & P. Vaurie, AMNH), 19.vii.1991 (W. H. Piel & G. S. Bodner, MCZ): Fortin de las Flores, 18°54’N, 97°O'W, 7.vii.1947 (G. & M. Goodnight, AMNH), 25.iv.1963 (W. J. Gertsch & W. Ivie, AMNH), 10.vii.1976 (A. Newton, MCZ), 21.x.1982 (George Uetz, MCZ), 17.vii.1991 (W. H. Piel & G. S. Bodner, MCZ); Jalapa, 19°32'N, 96°55'W, 14.ii.1948 (H. Wagner, AMNH), 15.iii.1948 (H. Wagner, AMNH), 15.x.1962 (N. L. H. Krauss, AMNH): Los Naranjos, 18°21'N, 96°10'W, 4.iii.1948 (H. Wagner, AMNH): near Monte Blanco, 18°58'’N, 97°1'W, 3.vili.1973 (A. Newton, MCZ): Orizaba, 18°51'N, 97°4'W, 6.vii.1963 (D. Bixler, MCZ); Rancho Chula- Vista, N of Cérdoba, 19°4'’N, 97°2'W, 18.vii.1991 (W. H. Piel & G. S. Bodner, MCZ). METEPEIRA ® Piel ital Metepeira ventura Group Spiders in the M. ventura group (Me- tepeira ventura, Metepeira revillagigedo, Metepeira celestun, Metepeira uncata, Me- tepeira crassipes, Metepeira chilapae) are closely related to those in the M. minima group (M. petatlan, M. minima, M. paci- fica, M. jamaicensis). Females in the M. ventura group have epigynal openings that are wider than long and shaped as trian- gles (Fig. 231), ovals (Fig. 252), or squares (Figs. 259, 266), with their posterior edges open. The posterior edges are observed to be open because the posterior lobes at their distal edge are wider than the distal end of the scape. In many other species, the gap between the lobes is narrower than the scape and therefore hidden from a ventral view, giving the impression that the openings form closed shapes (e.g., Fig. 123). In addition, with the exception of M. celestun (Fig. 245), the scape is relatively thin throughout its entire length (Figs. 231, 238, 252, 259, 266). In males, the em- bolus can be slim and elongated with a gentle curve (e.g., Figs. 236, 243), or as with some species in the M. minima group, it can be tapering to a sharp bend right at the tip (e.g., Figs. 229, 264). In general, the larger flagellum is a simple tapering extension off the base (Figs. 229, 243, 250, 257, 264, and to a lesser extent, Fig. 236), as opposed to being a distinctly separate structure that abruptly curves off the base (e.g., Fig. 129). Although the flagellae can be thin, as in the M. minima species group, they are not set off on a distinctly narrower stalk (e.g., compare Fig. 264 with Fig. 278). 29. Metepeira ventura Chamberlin and Ivie k Figures 229-235, 320; Map 13 Metepeira ensenada Chamberlin and Ivie, 1942: 65, figs. 166-168, 3. Male holotype from beach near Ensenada, Mexico, in the AMNH, examined. Syn- onymized by Levi (1977). Metepeira ventura Chamberlin and Ivie, 1942: 67, figs. 175-179, 2. Female holotype, 1 male and 3 female paratypes from between Oxnard and Santa Monica, California, USA, in the AMNH, examined. 2. Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Levi, 1977: 204, figs. 53-60. Brignoli, 1983: 275. Note. As first reviser, Levi (1977) preferred to use the name M. ventura because a larger number of specimens were available from the type locality. Description. Female from 10 mi north of Colonia Guerrero, Baja California Nor- te, Mexico. Carapace brown with white eye region, lateral posterior extensions, and white median line (Fig. 234). Legs yel- lowish tan with darker rings on fourth pair of legs. Femur I with row of four macro- setae on anterior side; two on anteroven- tral side. Dorsum of abdomen with usual Metepeira folium, lighter in anterior third (Fig. 234). Venter of abdomen with wide white median line posteriorly set within hint of surrounding U-shaped marking. Pair of white spots on either side of spi- racle (Fig. 235). Sternum has wide median white line widening anteriorly, sometimes fragmented (Fig. 235). Ratio of eye di- ameters: posterior medians and anterior medians 1.0, anterior laterals 1.6, posterior laterals 1.4. Anterior median eyes separat- ed by 1.4 diameters, posterior median eyes by 0.7, anterior median eyes separated from anterior laterals by 2.9 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 5.5 mm. Carapace 2.8 mm long, 2.2 wide. First femur 3.3 mm, patella and tibia 3.3, metatarsus 3, tarsus 1.2. Second patella and tibia 2.9 mm, third 1.7, fourth 2.5. Male from 10 mi north of Colonia Guerrero, Baja California Norte, Mexico. Carapace, abdomen, venter as in female (Figs. 232, 233). Ringed legs darker than in female. Femur I with row of four ma- crosetae on anterior side; six to seven on anteroventral side. Median white line on sternum often broken (Fig. 233). Ratio of eye diameters: posterior medians and an- terior medians 0.9, anterior laterals 1.2, posterior laterals 1.1. Anterior median eyes separated by 1.6 diameters, posterior median eyes by 0.8, anterior median eyes separated from anterior laterals by 1.9 di- ameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. To- tal length 4.3 mm. Carapace 2.2 mm long, 1.7 wide. First femur 3.6 mm, patella and tibia 3.6, metatarsus 3.7, tarsus 1.3. Sec- ond patella and tibia 2.8 mm, third 1.5, fourth 2.3. Diagnosis. Unlike other members of the M. minima species group, the larger fla- gellum on the median apophysis of M. ventura is a tapering extension off the base (Fig. 229), as opposed to arising from a distinctly separate stalk (Fig. 286). Like M. uncata, the epigynal openings of M. ven- tura (Fig. 231) are somewhat triangular in shape, in contrast to oval (Fig. 288) or slit- shaped (Fig. 280) as in other members of the M. minima species group. The rela- tively shorter scape and the dark circles inside the epigynal openings of M. ventura (Fig. 231) contrast with M. uncata’s rela- tively longer scape and the dark sinuous shapes inside the epigynal openings (Fig. 252) Variation. Average body length of eight females examined 6.7 mm, range 5.6 to 9.7 mm. Average body length of seven males examined 5.1 mm, range 3.9 to 7 mm. Natural History. Mature specimens are mostly collected April through September (Fig. 320). Altitudes range from near sea level to 1,000 m. Distribution. California to northwestern Mexico (Levi, 1977, map 1; Map 13). Records Examined. MEXICO Baja Calif. Norte: 10 m E El Rosario, 30°1L1'N, 115°46’W, 8.vii.1973 (S. C. Williams & K. B. Blair, CAS); 10 mi N Col. Guerrero, 28°5'N, 114°8’W, 1.ix.1957 (V. Roth, AMNH); 2 mi SE Eréndira, 31°19’N, 116°19’W, 12.v.1973 (S. C. Williams & K. B. Blair, CAS); Isla Cedros, Gran Cafi- on, 28°12'N, 115°15’W, 10.iii.1945 (B. F. Osorio Taf- all, AMNH): Isla de Cedros, Cerro de Cedros, 28°12'N, 115°15’W, 1.vii.1983 (V. F. Lee, CAS); Isla de Cedros, Punta Norte [?], 28°22'N, 115°14’W, 3.vii.1983 (V. F. Lee, CAS); Isla de Cedros, trail to Cerro de Cedros, at spring, 28°12’N, 115°15’W, 27.ix.1984 (D. B. Weissman, V. F. Lee, CAS); Islas San Benito, Benitos del Oeste, 28°18’N, 115°35’W, 4.vii.1983 (D. C. Lightfoot & V. F. Lee, CAS); Islas San Benito, Middle Island, 28°19’N, 115°34'W, 9.iv.1981 (Stanley C. Williams, CAS); Islas San Be- nito, South Island, 28°18’N, 115°35'W, 9.iv.1981 (Stanley C. Williams, CAS); near Consuelo, 6 mi NW El Rosario, 30°11’N, 115°46’W, 18.iv.1965 (D. Q. Cavagnaro, C. E. & E. S. Ross, V. L. Vesterby, CAS); Rancho Las Parritas, 10 mi S San Quintin, 30°20'N, 115°57'W, 27.vi.1977 (C. E. Griswold, CAS); Santo Tomas, 31°33’N, 116°24’W, 8.vii.1953 (W. J. & J. W. Gertsch, AMNH). Baja Calif. Sur: Bahia de los Muertos, 23°58'’N, 109°50'W, 20.xii.1958 (H. B. Leech, CAS); Desierto del Vizcaino, Laguna Ojo de Liebre, sobre frutilla, 27°43’N, 114°15’W, 11.xi.1981 (A. Cota & M. Jiménez, MLJ); E edge of Sierra Pla- ceres, 27°35'N, 114°30'W, 25.iii.1984 (W. J. Pulawski, CAS). Sonora: 6 mi E Navojoa, 27°6'N, 109°23’W, 23.viii. 1965 (W. J. Gertsch & R. Hastings, AMNH). USA California: 1 mi NW Winchester (Double Butte), 33°43’N, 117°6’W, 7.xii.1976 (W. Icenogle, MCZ); Lompoc, by US 63, 34°38'N, 120°27'W, 17.viii.1966 (L. & P. Pinter, MCZ); Lucia, Monterey Co., 36°1'N, 121°33'W, 15.ix.1964 (L. Pinter, MCZ); Manzanita chaparral, San Gabriel Canyon, Coldbrook Ranger Station, 34°11'N, 117°53’W, 29.v.1965 (L. Pinter, MCZ); Santa Catalina Island Area near Hay- press Res, 33°23'’N, 118°25’'W, 6.ii.1993 (Martin G. Ramirez & Laura B. Fandino, MCZ):; Santa Catalina Island, Mt. Torquemada, 33°26’N, 118°33'W, 15.viii. 1965 (L. Pinter, MCZ): Winchester, 33°42'N, 117°5'W, 11.v.1970 (W. Icenogle, MCZ); Winchester, Double Butte, 33°42’N, 117°5’W, 19.v.1974 (W. Icen- ogle, MCZ). Nevada: Quin River Crossing, 41°35’N, 118°27'W, 21.vi.1975 (G. F. Knowlton, MCZ). 30. Metepeira revillagigedo new species Figures 236-242, 301; Map 12 Holotype. Female from south of Isla Socorro, Archi- piélago de Revillagigedo, State of Colima, Mexico, 15.xii.1988, M. Jiménez, in MCZ. The specific name is a noun in apposition after the locality. Description. Female holotype. Carapace dark brown with light region in median eye quadrangle and surrounding lateral eyes. Pair of darker, elliptic, walnut leaf shapes on lighter patch near center of car- apace (see Fig. 241). Darker rings on distal ends of femora, patellae, tibiae, and prox- imal dorsal portion of tibia I, Il. Femur I with three to four macrosetae on anterior side; none on the anteroventral side. Dor- sum of abdomen with white oak leaf foli- um, margined with dark markings, partic- ularly on trailing edges of lobes (Fig. 241). Coloration has a slightly golden hue. Dark lateral band wraps around sides of abdo- men and stretches up over anterior dorsal portion. Venter with two white spots on either side of spiracle. Anterior to spiracle, slight indication of a V-shaped mark with longitudinal extensions reaching halfway up abdomen. Wide, white medial longitu- METEPEIRA ® Piel 73 dinal line to epigynal groove (Fig. 242). Sternum with wide, white longitudinal mark widening anteriorly (Fig. 242). Ratio of eye diameters: posterior medians and anterior medians 0.8, anterior laterals 1.1, posterior laterals 1. Anterior median eyes separated by 1.3 diameters, posterior me- dian eyes by 1.0, anterior median eyes sep- arated from anterior laterals by 2.5 diam- eters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 7.5 mm. Carapace 3.2 mm long, 2.5 wide. First femur 3.5 mm, patella and tibia 3.8, metatarsus 3.2, tarsus 1.2. Second pa- tella and tibia 3.2 mm, third 1.8, fourth 2S: Male paratype from Isla Socorro, Colli- ma. Carapace dirty yellowish brown with lighter posteriorly pointing acute triangle in center (Fig. 239). Legs same color as carapace, except lighter on proximal half of femora. Femur I with row of four ma- crosetae on anterior side, row of two ma- crosetae on anteroventral side. Male ab- domen lighter and with less contrast than female (Fig. 239). Venter with a much re- duced and shorter longitudinal line than female. Sternum as in female, except lon- gitudinal line more often broken (Fig. 240). Ratio of eye diameters: posterior me- dians and anterior medians 1.0, anterior laterals 1.1, posterior laterals 1.1. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 0.8, anterior me- dian eyes separated from anterior laterals by 1.4 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diame- ters. Total length 3.2 mm. Carapace 1.7 mm long, 1.1 wide. First femur 2.5 mm, patella and tibia 2.5, metatarsus 2.3, tarsus 0.9. Second patella and tibia 2 mm, third 1.0, fourth 1.5. Diagnosis. The epigynum of M. revilla- gigedo looks very different from those of other species in the M. ventura group. In- stead of having small, sharply delineated depressions on either side of the scape (e.g., Fig. 259), M. revillagigedo has larger but more gradual depressions (Fig. 238). Despite the unique appearance of the epi- 74 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 gynum, the affinity between M. revillagi- gedo and the M. ventura species group can be seen in the black comma-shaped marks inside the epigynal depressions (Fig. 238) similar to those in, for example, M. ven- tura (Fig. 231). The male emboli of M. revillagigedo (Fig. 236) and M. celestun (Fig. 243) are very similar in shape and relatively longer than in other species in the M. ventura group. Males of M. revil- lagigedo differ from males of M. celestun by the thickness and shape of the median apophysis (compare Fig. 236 with Fig. 943). Natural History. Specimens have only twice been collected: once in December and once in April (Fig. 301). This species is known to live in fig trees. Distribution. This species is found on Isla Socorro in the Pacific Ocean (Map 12), the island being one of several that form the Archipiélago de Revillagigedo. It is well isolated, situated about 465 km south of Baja California Sur and 588 km west of the Jalisco coastline. Records Examined. MEXICO Colima: Archipiéla- go de Revillagigedo: Isla Socorro, 18°45'N, 110°57’W, 26.iv.1932 (Templeton, Crocker Exped., CAS); Ar- chipiélago de Revillagigedo: Sur de la Isla Socorro, 18°44'N, 110°57'W, 15.xii.1988 (M. Jiménez, MCZ). 31. Metepeira celestun new species Figures 243-249, 326; Map 13 Holotype. Male from Celestun, Yucatan, Mexico, 24 .vii.1991, W. H. Piel & G. S. Bodner, in MCZ. The specific name is a noun in apposition after the locality. Note. Since males of this species were never found with females, one cannot be sure absolutely that they are conspecific. Several facts argue for conspecificity: all specimens were collected in sim- ilar habitats and during the same season; dorsal fo- lium of males and females share similar patterns and both have hint of gold coloration; based on parallel correspondence with other species, the male palp is arguably compatible with the female's epigynum. Description. Female paratype from Edzna, Campeche, Mexico. Carapace yel- lowish tan with lighter area around eyes (Fig. 248). Legs yellowish tan with darker rings on fourth pair of legs. Femur I with row of four macrosetae on anterior side: sometimes four on anteroventral side. Dorsal folium as in other Metepeira, ex- cept speckled an unusual reddish gold col- or (Fig. 248). Venter wide white median line with pair of large white spots on either side of spiracle (Fig. 249). Sternum has wide median white line widening anteri- orly with constriction in center (Fig. 249). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 2.4 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 5.2 mm. Carapace 2.5 mm long, 1.9 wide. First femur 2.6 mm, patella and tibia 2.8, metatarsus 2.2, tarsus 0.9. Sec- ond patella and tibia 2.4 mm, third 1.4, fourth 2. Male holotype. Carapace dark with light region around eyes and a light triangular mark anterior to thoracic furrow (Fig. 246). Coxae and proximal third of femora white, remaining two-thirds black; other articles similarly ringed. Femur I with row of four macrosetae on anterior side; three on anteroventral side. Dorsal folium with slight gold iridescence. Fleur-de-lis pat- tern leafy and pinnate (Fig. 246). Venter, sternum as in female, except median white line on sternum often broken (Fig. 247). Ratio of eye diameters: posterior medians and anterior medians 1.1, anterior laterals 1.4, posterior laterals 1.3. Anterior median eyes separated by 1.6 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 1.4 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 2.8 mm. Carapace 1.4 mm long, 1.1 wide. First femur 1.8 mm, patella and tibia 1.8, metatarsus 1.4, tarsus 0.8. Sec- ond patella and tibia 1.5 mm, third 0.8, fourth 1.2. Diagnosis. The overall coloration of M. celestun in alcohol has an unusually golden quality that is rare among preserved Me- METEPEIRBA ° Piel 75 Para ry 3 je ij Pa revillagigedo 239 ¥ 240 (30) uncata (32) Figures 236-242. Metepeira revillagigedo new species (sp. 30; 18°44’N, 110°57’W). 236, male palpus, mesal. 237, epigynum, posterior. 238, epigynum, ventral. 239, male, dorsal. 240, male, ventral. 241, female, dorsal. 242, female, ventral. Figures 243-249. Metepeira celestun new species (sp. 31 [243,246,247] 20°56’N, 90°21’W; [244,245,248,249] 19°35’N, 90°15'W). 243, male palpus, mesal. 244, epigynum, posterior. 245, epigynum, ventral. 246, male, dorsal. 247, male, ventral. 248, female, dorsal. 249, female, ventral. Figures 250-256. Metepeira uncata F. O. P.-Cambridge (sp. 32 [250,251,253-256] 14°40’N, 92°9’W; [252] 14°49’N, 91°31’W), 250, male palpus, mesal. 251, epigynum, posterior. 252, epigynum, ventral. 253, male, dorsal. 254, male, ventral. 255, female, dorsal. 256, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 76 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 tepeira. Unlike all other members of the M. ventura species group, the scape of M. celestun is very wide at its base, thereby partly concealing the epigynal openings (Fig. 245). However, if the scape is dis- carded, the exposed openings have black comma-shaped marks similar to those of M. ventura (Fig. 231), thereby confirming the species’ affinities with other members in the M. ventura species group. Unlike other members, the posterior lobes nearly touch each other near their base (Fig. 244). The emboli of M. celestun and M. revillagigedo are very similar in shape and relatively longer than in other species in the M. ventura group (Fig. 243). Metepei- ra celestun differs from M. revillagigedo by the thickness and shape of the median apophysis (compare Fig. 243 with Fig. 236). Variation. Average body length of four females examined 5.6 mm, range 5.2 to 6.4 mm. Average body length of three males examined 2.7 mm, range 2.2 to 2.9 mm. Female scape sometimes wider than it ap- pears in Fig. 245, resembling Metepeira arizonica. Legs and carapace color vary from yellowish tan to orange-red. Natural History. Mature specimens have been collected in July (Fig. 326) in forested clearings, roadside bushes, palms, and swampy areas near the beach. Distribution. Yucatan peninsula (Map IB), Records Examined. MEXICO Campeche: Edzna, 19°35'N, 90°15’W, 22.vii.1991 (W. H. Piel & G. S. Bodner, MCZ). Quintana Roo: Kohunlich ruins, 9 km S Franciso Villa, 18°26’N, 88°48’W, 15.vii.1983 (R. S. Anderson, MCZ). Yucatan: 3km S San Felipe, 21°32'N, 88°14'’W, 25.vii.1991 (W. H. Piel & G. S. Bodner, MCZ); 4 km N Xocenpich, 12 km N Piste, on road to Dzitas, 20°47'N, 88°34'W, 20.vii.1983 (W. Maddison, MCZ); Balankanche Cave, 2 km E Chi- chen Itza, 20°40’N, 88°33’W, 19.vii.1983 (W. Mad- dison, MCZ): beach north of Celesttin, 20°56'N, 90°21'W, 24.vii.1991 (W. H. Piel & G. S. Bodner, MCZ): Chichen Itza ruins on HWY 180, seasonal for- est, 20°40'’N, 88°34'W, 19.vii.1983 (W. Maddison & R. S. Anderson, MCZ); Uxmal, 20°22’N, 89°46’W, 23.vii.1991 (W. H. Piel & G. S. Bodner, MCZ). 32. Metepeira uncata F. O. P.-Cambridge Figures 250-256, 329; Map 12 Metepeira uncata F. O. P.-Cambridge, 1903: 459, fig. 8, 6. Male holotype from Santa Ana, Guatemala, in BMNH. Roewer, 1942: 868. Bonnet, 1957: 2823. Araneus uncatus:—Petrunkevitch, 1911: 321. Description. Female from Ayutla, San Marcos, Guatemala. Carapace dark brown with light region surrounding the eyes and extending posteriorly behind the lateral eyes. Pair of darker feather-shaped patches anterior of thoracic furrow, touching pos- teriorly to create a U-shape (see Fig. 255). Dark rings on distal half of leg articles, an- terior side of patellae, and dark markings on dorsal, proximal end of tibia I and II. Femur I with four macrosetae on anterior side; one on anteroventral side. Dorsum of abdomen with white fleur-de-lis folium; this pattern thinner than in most other species. Fleur-de-lis pattern on a dark background, outlined on either side by thin white stripe (Fig. 255). Dark lateral band follows sides of abdomen and stretches up over anterior dorsal portion. Venter with two white spots on each side of spiracle; wide, white, medial longitudi- nal line starts anterior to the colulus and ends posterior to the epigynal groove. Thin, faint lines run parallel to wide me- dian one anterior to white spots. White patch separates epigynum and dark de- pression posterior to pedicel (Fig. 256). Sternum black with white dewdrop- shaped mark at posterior end (Fig. 256). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separated by 1.6 diameters, posterior median eyes by 1.1, anterior median eyes separated from anterior laterals by 4 di- ameters of anterior lateral eyes, lateral eyes separated by 0.4 their diameters. To- tal length 7.8 mm. Carapace 3.7 mm long, 2.9 wide. First femur 4 mm, patella and tibia 4.3, metatarsus 3.8, tarsus 1.4. Sec- ond patella and tibia 3.6 mm, third 2.3, fountineoror Male from Ayutla, San Marcos, Guate- mala. Carapace dirty yellowish brown with lighter mark in center (Fig. 253). Same color as carapace except lighter on proxi- mal half of femora. Femur I with row of nine macrosetae on anterior side; five on anteroventral side. Male abdomen similar to female, but leaves of fleur-de-lis pattern thinner (Fig. 253). Venter with a shorter longitudinal line, as compared with female (Fig. 254). Sternum with posterior white dewdrop as in female, and additional an- terior white mark near labium (Fig. 254). Ratio of eye diameters: posterior medians and anterior medians 0.9, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separated by 1.5 diameters, posterior median eyes by 1.0, anterior median eyes separated from anterior laterals by 2 di- ameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. To- tal length 4.5 mm. Carapace 2.3 mm long, 1.8 wide. First femur 3.4 mm, patella and tibia 3.4, metatarsus 3.2, tarsus 1.1. Sec- ond patella and tibia 2.7 mm, third 1.4, fourth 2. Diagnosis. The generally darker pig- mentation (Fig. 255) and the small white mark on the sternum (Fig. 256) separate M. uncata from all other members of the M. ventura species group. Like M. ventura (Fig. 231) but unlike other members of the M. ventura species group, the open- ings to the epigynum are more triangular to oval (Fig. 252), rather than square (Figs. 259, 266). Also distinctive are the sinuous black lines that form upside-down U- shapes inside each epigynal opening (Fig. 252), in contrast to the comma-shaped marks in M. ventura or M. chilapae (Figs. 231, 266). Like M. chilapae but unlike oth- er members of the M. ventura group, the median apophysis and flagellae of M. un- cata are slimmer and of uniform thickness (Figs. 250, 264). The palp of M. uncata differs from that of M. chilapae by the more basal position of the bend in the em- bolus tip (compare Fig. 250 with Fig. 264). Variation. Average body length of five females examined 8.2 mm, range 7.5 to 8.8 METEPEIRA ® Piel UH mm. Average body length of four males examined 5.6 mm, range 4.5 to 6.4 mm. Natural History. Mature specimens have been collected in August (Fig. 329). Distribution. Southwestern Guatemala to northern Costa Rica (Map 12) at alti- tudes ranging from 100 to 3,000 m. Records Examined. COSTA RICA Cordillera: 20 km N Siquires, 10°9'N, 84°17'W, 15.viii.1980 (W. Eberhard, MCZ). GUATEMALA El Quiche: Chichi- castenango, 14°56'N, 91°7’W, 6.viii.1947 (C. & P. Vaurie, AMNH). Huehuetenango: Todos Santos Cu- chumatan, 15°31’N, 91°37'W, 16.viii.1979 (C. E. Griswold, CAS). Quetzaltenango: E] Baul, 1 km S Quetzaltenango, 14°49'N, 91°31'W, 14.viii.1979 (T. C. Meikle & C. E. Griswold, CAS); Quetzaltenango, 14°50'N, 91°31'W, 16.viii.1950 (C. J. & M. Good- night, AMNH). San Marcos: Ayutla, 14°40'N, 92°9'W, 19.viii.1947 (AMNH). 33. Metepeira crassipes Chamberlin and lIvie Figures 257-263, 309; Map 12 Metepeira josepha Chamberlin and Ivie, 1942: 64, fig. 165, 2. Female holotype from Kings Mtn. near Palo Alto, California in the AMNH. Synonymized by Levi (1977). Metepeira crassipes Chamberlin and Ivie, 1942: 66, figs. 171-173, 2, 3. Male holotype, female, male paratypes from Laguna Beach, California in the AMNH. Levi, 1977: 202, figs. A= 52 ORO Note. As first reviser, Levi (1977) preferred to use name M. crassipes. Description. Female from Isla San Pe- dro Nolasco, Sonora, Mexico. Carapace with large white eye region and lateral posterior extensions (Fig. 262). Legs ringed at distal ends of articles—though sometimes only lightly. Femur I with four macrosetae on anterior side aligned in straight row; one to four light setae on an- teroventral side. Dorsum of abdomen with usual Metepeira folium (Fig. 262); venter wide, with long white median line set within U-shaped thinner white lines. Pair of white spots on either side of spiracle (Fig. 263). Sternum black with wide me- dian white line widening anteriorly (Fig. 263). Ratio of eye diameters: posterior me- dians and anterior medians 1.1, anterior laterals 1.4, posterior laterals 1.3. Anterior median eyes separated by 1.4 diameters, 78 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 posterior median eyes by 0.8, anterior me- dian eyes separated from anterior laterals by 3 diameters of anterior lateral eyes, lat- eral eyes separated by 0.1 their diameters. Total length 5.7 mm. Carapace 2.8 mm long, 2.1 wide. First femur 3.2 mm, patella and tibia 3.1, metatarsus 2.8, tarsus 1. Sec- ond patella and tibia 2.7 mm, third 1.6, fourth 2.5. Male from Winchester, California. Car- apace as in female except often with me- dian white mark. Femur I with row of four macrosetae on anterior side; about four to eight on anteroventral side. Dorsum, ven- ter as in female (Fig. 260); median white line on sternum more often broken (Fig. 261). Ratio of eye diameters: posterior me- dians and anterior medians 1.0, anterior laterals 1.6, posterior laterals 1.4. Anterior median eyes separated by 1.2 diameters, posterior median eyes by 0.6, anterior me- dian eyes separated from anterior laterals by 2 diameters of anterior lateral eyes, lat- eral eyes separated by 0.3 their diameters. Total length 4 mm. Carapace 2.1 mm long, 1.5 wide. First femur 3.2 mm, patella and tibia 3.2, metatarsus 3.2, tarsus 1.1. Sec- ond patella and tibia 2.5 mm, third 1.3, fOumtle 2) Diagnosis. Unlike other species in the M. ventura species group, the openings to the epigynum of M. crassipes resemble those of M. chilapae because they are shaped like squares with rounded edges (compare Fig. 259 with Fig. 266). How- ever, compared to M. chilapae, the epigyn- al openings of M. crassipes are relatively smaller and the scape is relatively longer. The embolus of M. crassipes is distin- guished by its gentle curvature (Fig. 257), in contrast to much more abrupt curvature seen in other species in the M. ventura group (e.g., Fig. 264), with the exception of M. revillagigedo. The median apophysis of M. crassipes (Fig. 257) is slimmer than that of M. revillagigedo (Fig. 236). Variation. Average body length of four females examined 6 mm, range 5 to 6.5 mm. Average body length of two males ex- amined 4.4 mm, range 4.1 to 4.7 mm. Sometimes the scape is greatly swollen, adding to its relative width. In such cases the scape can cover the epigynal openings, which may mislead the investigator to con- fuse it for Metepeira arizonica. Natural History. Levi (1977) reports that males in the U.S. have been collected from April to October, primarily in Cali- fornia buckwheat and sage. Mexican re- cords expand this seasonality to include the entire year (Fig. 309). Distribution. From northern California in the U.S. to Baja California Sur, Mexico (Levi, 1977, map 1; Map 12). Records Examined. MEXICO Baja Calif. Norte: Santo Tomas, 31°33'N, 116°24’W, 8.vii.1953 (W. J.& J. W. Gertsch, AMNH), 12.xi.1976 (S. C. Williams & K. B. Blair, CAS). Baja Calif. Sur: 26 mi S Loreto, 95°37'N, 111°17'W, 1.i1.1977 (C. E. Griswold & L. Vincent, CAS); Isla Magdalena, Puerto Magdalena, 24°38'N, 112°9'W, 16.iii.1957 (R. Zweifel, AMNH); Isla San Francisco, South Side, 24°50'N, 110°35’W, 19.v.1970 (S. C. Williams & V. F. Lee, CAS); Sierra San Nicolas, 26°32'N, 111°36’W [?] (Eisen & Vaslit, MCZ). Sonora: Isla San Pedro Nolasco, 27°58’N, 111°25'W, 17.iv.1921 (G. C. Chamberlin, MCZ); Si- erra de Alamos, 30°51'N, 112°2'W [?], 19.i.1968 (V. Roth, AMNH). USA California: 1 mi NW Winches- ter (Double Butte), in web between Artemisia cali- fornica bushes, 33°43'N, 117°6'W, 4.xii.1976 (W. Icenogle, MCZ). 34. Metepeira chilapae Chamberlin and lvie Figures 264-270, 333; Map 13 Metepeira chilapae Chamberlin and Ivie, 1936: 45, figs. 119-121, d. Male holotype from Chilapa, Guerrero, Mexico, in the ZMB. Roewer, 1942: 868. Metepeira chilapica:—Bonnet, 1957: 2820. Unjusti- fied emendation. Note. Examination of Chamberlin and_Ivie’s (1936) and Levi's (personal illustrations) figures of the holotype was sufficient to identify M. chilapae accurately. Description. Female from Cocoyoc, Morelos, Mexico. Brownish black cara- pace, paler around and just posterior to lateral eyes (Fig. 269). Distal halves of ventral leg articles black, elsewhere yel- lowish. Femur I with row of four macro- setae on anterior side; zero to three setae on anteroventral side. Dorsal folium a white fleur-de-lis marking set on golden- METEPEIRA ® Piel 79 petatlan (35) Figures 257-263. Metepeira crassipes Chamberlin and lvie (sp. 33 [257-261] 33°43’N, 117°6’W; [262,263] 27°58’N, 111°25’W). 257, male palpus, mesal. 258, epigynum, posterior. 259, epigynum, ventral. 260, male, dorsal. 261, male, ventral. 262, female, dorsal. 263, female, ventral. Figures 264-270. Metepeira chilapae Chamberlin and Ivie (sp. 34 [264] 17°39'N, 99°22’W; [265-270] 17°39’N, 99°22’W). 264, male palpus, mesal. 265, epigynum, posterior. 266, epigynum, ventral. 267, male, dorsal. 268, male, ventral. 269, female, dorsal. 270, female, ventral. Figures 271-277. Metepeira petatlan new species (sp. 35 [271,274,275] 17°14’N, 100°53’W; [272,273,276,277] 17°17'32'N, 101°2'40”"W). 271, male palpus, mesal. 272, epigynum, posterior. 273, epigynum, ventral. 274, male, dorsal. 275, male, ventral. 276, female, dorsal. 277, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 80 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 brown speckled pattern over white. Foli- um darkens posteriorly (Fig. 269). Ante- rior shoulders black. Venter black with wide median white line and pair of white spots on either side of spiracle (Fig. 270). Sternum brownish black with wide, white line widening anteriorly (Fig. 270). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.1, posterior laterals 1.1. Anterior median eyes separated by 1.3 diameters, posterior median eyes by 0.8, anterior median eyes separated from anterior laterals by 2.2 di- ameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. To- tal length 5.8 mm. Carapace 2.7 mm long, 2.3 wide. First femur 2.9 mm, patella and tibia 3, metatarsus 2.6, tarsus 1. Second patella and tibia 2.4 mm, third 1.6, fourth Dheore Male from 6 mi northeast of Tixtla de Guerrero, Guerrero, Mexico. Carapace, dorsum, venter, sternum darker and more contrasty version of female (Figs. 267, 268). Distal halves of ventral leg articles black, elsewhere yellowish. Femur I with row of four macrosetae on anterior side: three on anteroventral side. Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.3. Anterior median eyes separat- ed by 1.7 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 1.9 diameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. Total length 3.5 mm. Carapace 1.8 mm long, 1.4 wide. First femur 2.3 mm, patella and tibia 2.1, metatarsus 1.8, tarsus 0.8. Second patella and tibia 1.8 mm, third 0.9, fourth 1.4. Diagnosis. The openings to the epigyn- um resemble those of M. crassipes be- cause they are shaped like squares with rounded edges (compare Fig. 266 with Fig. 259). However, the dark, comma- shaped mark inside each opening resem- bles that of M. ventura (compare Fig. 266 with Fig. 231). As in M. ventura (Fig. 229) but unlike all other species in the M. ven- tura species group, the embolus tapers strongly then curves sharply up and inward right at the tip (Fig. 264). The bend in the embolus in other M. ventura group species is less pronounced (Figs. 236, 257) or not so near the tip (Figs. 243, 250). Unlike M. ventura, the median apophysis in M. chi- lapae is slimmer (Fig. 264) and the sepa- ration between the embolus and the basal embolic apophysis is relatively greater. Variation. Average body length of eight females examined 6.5 mm, range 5.7 to 8.3 mm. Average body length of three males examined 2.9 mm, range 2.3 to 3.5 mm. Natural History. Mature specimens have been collected in July through Oc- tober (Fig. 333). Distribution. Southern Mexican states between Nayarit and Oaxaca; elevations from 1,000 to 2,000 m (Map 13). Records Examined. MEXICO Guerrero: 6 mi NE Tixtla de Guerrero, 17°39'N, 99°22’W, 16.vii.1984 (J. B. Woolley, ADC). Morelos: Cocoyoc, 18°52'N, 98°59'W, 28.vii.1956 (W. Gertsch & V. Roth, AMNH); Cuernavaca, 18°55'N, 99°15’W, 15.x.1944 (N. L. H. Krauss, AMNH). Nayarit: 30 mi SE Tepic, 21°12'N, 104°33'W, 23.xi.1948 (E. S. Ross, CAS): 5 mi NW Tepic, 21°32'N, 104°57'W, 13.v.1963 (W. J. Gertsch & W. Ivie, AMNH). Oaxaca: 6 mi NE Mitla, 16°59'N, 96°21'W, 20.viii.1985 (J. Woolley & G. Zolnerowich, ADC); Huajuapan, 17°48'N, 97°46’W, 1.x.1946 (Ee Wagner, AMNH); Oaxaca, 17°3'N, 96°43'W, 17.vii.1955 (C. & P. Vaurie, AMNH), 12.viii.1991 (W. H. Piel & G. S. Bodner, MCZ); San Balt. Chichica- pan, 16°45'N, 96°29'W, 4.viii.1991 (W. H. Piel & G. S. Bodner, MCZ). Metepeira minima Group Female spiders in the M. minima group (Metepeira petatlan, Metepeira minima, Metepeira pacifica, Metepeira jamaicensis) have a thin scape with epigynal openings that are shaped from longitudinal slits (Figs. 273, 280, 281) to ovals that are lon- ger than wide (Figs. 288, 295). In males, the flagellae on the median apophysis are set off on a distinctly narrower stalk (Figs. SHTAL. SATS. DASTG,,. OXSIS}). 35. Metepeira petatlan new species Figures 271-277, 334; Map 10 Holotype. Male from 50 km southeast of Petatlan, Guerrero, Mexico, 14.viii.1984, J. B. Woolley, in MCZ. The specific name is a noun in apposition after the locality. Description. Female paratype from Pa- panoa, Guerrero. Light region around pos- terior eye row; carapace darkens posteri- orly, then lightens under overhang of ab- domen (Fig. 276). Slight rings on patella and tibia. Femur I with three macrosetae on anterior side, none on anteroventral side. Dorsum of abdomen with typical fo- lium, except that white oak leaf pattern narrower than most species and set on a narrow, remarkably darker brownish dove- tail (Fig. 276). Dark lateral band wraps around abdomen and stretches up over an- terior dorsal portion. Venter with two spots on either side of spiracle and slight indi- cation of a transverse bar. Wide, white lon- gitudinal mark ends at the epigynal groove (Fig. 277). Sternum with wide, white lon- gitudinal mark widening anteriorly (Fig. 277). Ratio of eye diameters: posterior me- dians and anterior medians 1.0, anterior laterals 1.3, posterior laterals 1.2. Anterior median eyes separated by 1.1 diameters, posterior median eyes by 0.7, anterior me- dian eyes separated from anterior laterals by 1 diameter of anterior lateral eyes, lat- eral eyes almost touching. Total length 5 mm. Carapace 2.2 mm long, 1.7 wide. First femur 2.3 mm, patella and tibia 2.5, metatarsus 2, tarsus 0.9. Second patella and tibia 2 mm, third 1.3, fourth 1.9. Male holotype. Carapace uniform brown with a lighter median streak just an- terior to thoracic depression (Fig. 274). Weakly ringed on tibia; distal half of fem- ora dark. Femur I with two macrosetae on anterior side. Abdomen is a lighter version of the female (Fig. 274). Venter with a much reduced and shorter longitudinal line, compared to the female (Fig. 275). Sternum as in female. Ratio of eye diam- eters: posterior medians and anterior me- dians 1.0, anterior laterals 1.4, posterior laterals 1.4. Anterior median eyes separat- ed by 1.1 diameters, posterior median eyes by 0.7, anterior median eyes separated from anterior laterals by 1.1 diameters of METEPEIRBA ° Piel Sl anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 2 mm. Carapace 1 mm long, 0.7 wide. First fe- mur 1.2 mm, patella and tibia 1.0, meta- tarsus 0.8, tarsus 0.5. Second patella and tibia 0.9 mm, third 0.5, fourth 0.8. Diagnosis. The female’s dorsum can be distinguished from other species by the unusually dark brown color of the dovetail- shaped mark (Fig. 276). Both this mark and the white oak leaf pattern have nar- rower margins that are more parallel and not as wedge-shaped as they are in other species (Fig. 276). The epigynum resem- bles that of M. minima by the narrow scape and the parallel slitlike openings on either side. These openings differ from those of M. minima because they have conspicuous sclerotized spheres just ante- rior to the lateral edge of the slits (Fig. 273) but which are much farther away in M. minima (Fig. 280). The male is unusu- ally small and its palp is unique and easily distinguished from other species. The lon- ger flagellum on the median apophysis is needlelike over its entire length, not grad- ually tapering (compare Fig. 271 with Fig. 278). In addition, the long needlelike fla- gellum has a sharper elbow. At the elbow it projects away from the palp for a short distance and then strongly curves around the palp (Fig. 271). Natural History. The female paratype was found at eye level on a hot, dry, wood- ed hillside overlooking the Pacific Ocean at an altitude of about 200 m. Her web differed considerably from the usual Me- tepeira web: instead of an elaborate barrier web, it consisted of a much reduced Y- shaped structure with a curled leaf in the center. The leaf served as a retreat for the spider and protected and hid three egg sacs. The retreat was suspended very near the hub, as opposed to farther away and above it, as found in most other species. Mature specimens have been collected in August through October (Fig. 334). Distribution. Western Mexico: Guerrero and Sinaloa (Map 10). 82 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Records Examined. MEXICO Guerrero: 32 mi SE Petatlan, 17°14’N, 100°53’W, 14.viii.1984 (J. B. Wool- ley, MCZ); Microondas Tamarindos, S. Papanoa, 17°17'32"/N, 101°2’40’W, 18.x.1994 (W. H. Piel, MCZ). Sinaloa: 2 mi S Elota, 23°55'N, 106°48’W, 11.ix.1966 (Jean & Wilton Ivie, AMNH). 36. Metepeira minima Gertsch Figures 278-285, 328; Map 15 Metepeira minima Gertsch, 1936: 10, fig. 31, ¢. Male holotype from Edinburg, Texas, in the AMNH, ex- amined. Roewer, 1942: 869. Chamberlin and Ivie, 1942: 67, fig. 174, 2. Levi, 1977: 206, fig. 70-73, 76-77, 2; 74-75, 6. Bonnet, 1957: 2822. Brignoli, 1983: 275. Description. Female from Celestun, Yu- catan, Mexico. Carapace dark brown with light region surrounding the eyes, some- times extending posteriorly behind the lat- eral eyes. Faint pair of darker feather- shaped patches in center of carapace (Fig. 284). Legs same color as carapace; darker on distal ends of articles. Femur I with row of four macrosetae on anterior side; rarely any on anteroventral side. Dorsum of abdomen with usual Metepeira folium (Fig. 284); venter wide, with long white median line surrounded by faint, thin, white U-shaped line (Fig. 285). Pair of large white spots on either side of spiracle. Sternum black with wide, white, uneven median line (Fig. 285). Ratio of eye di- ameters: posterior medians and anterior medians 1.0, anterior laterals 1.2, posterior laterals 1.1. Anterior median eyes separat- ed by 1.2 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 1.9 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 6.5 mm. Carapace 2.7 mm long, 2 wide. First femur 2.8 mm, patella and tibia 2.8, meta- tarsus 2.5, tarsus 0.9. Second patella and tibia 2.4 mm, third 1.5, fourth 2. Male from Celestun, Yucatan, Mexico. Carapace dark with light mark anterior to thoracic furrow (Fig. 282). Femur I with row of four macrosetae on anterior side; row of two to five macrosetae on anter- oventral side. Distal two-thirds same color as carapace, proximal third white. Folium of abdomen as in female, except posterior half darker than anterior half (Fig. 282). Venter of abdomen and sternum as in fe- male, except median white line of sternum often broken (Fig. 283). Ratio of eye di- ameters: posterior medians and anterior medians 1.1, anterior laterals 1.4, posterior laterals 1.1. Anterior median eyes separat- ed by 1.2 diameters, posterior median eyes by 0.8, anterior median eyes separated from anterior laterals by 1.5 diameters of anterior lateral eyes, lateral eyes separated by 0.1 their diameters. Total length 3 mm. Carapace 1.5 mm long, 1.2 wide. First fe- mur 2 mm, patella and tibia 1.9, metatar- sus 1.8, tarsus 0.8. Second patella and tibia 1.6 mm, third 0.8, fourth 1.3. Diagnosis. Unlike other members of the M. minima species group, the openings to the epigynum of M. minima are narrow slits around a scape of variable size (Figs. 280, 281). The openings of M. petatlan are only slightly wider than M. minima, but the internal darker sclerotized spheres in M. minima sit much farther apart (com- pare Fig. 280 with 273). Unlike M. pacifica (Fig. 286) and M. jamaicensis (Fig. 293), the bent embolus on M. minima tapers strongly and is therefore not like a needle (Fig. 278). Unlike M. petatlan, the longer flagellum on M. minima tapers (Fig. 278) and is not thin over its entire length (Fig. DHT Sly) Variation. Average body length of 13 fe- males examined 5.4 mm, range 3.7 to 6.5 mm. Average body length of 17 males ex- amined 3.4 mm, range 2.5 to 4.5 mm. The epigyna of females from the Yucatan pen- insula differ noticeably from all others, but the males hardly show any distinguishing features. These differences can be seen in the much wider scape and the greater sep- aration between openings in a specimen from the Yucatan (Fig. 281), compared to a specimen from Tamaulipas (Fig. 280) or one from Texas (Levi, 1977: 209, fig. 71). In addition, the posterior view of the epi- gynum from a Yucatan specimen shows the lobes converging to form a V-shape (Fig. 279), whereas females outside of the METEPEIRA ® Piel 83 2338 jamaicensis (38) &: 294 296 \ 207] Figures 278-285. Metepeira minima Gertsch (sp. 36 [278,279,281—285] 20°56’N, 90°21'W; [280] 22°30’N, 99°4’W). 278, male palpus, mesal. 279, epigynum, posterior. 280,281, epigynum, ventral. 282, male, dorsal. 283, male, ventral. 284, female, dorsal. 285, female, ventral. Figures 286-292. Metepeira pacifica new species (sp. 37; 10°27’N, 85°9’W). 286, male palpus, mesal. 287, epigynum, posterior. 288, epigynum, ventral. 289, male, dorsal. 290, male, ventral. 291, female, dorsal. 292, female, ventral. Figures 293-299. Metepeira jamaicensis Archer (sp. 38; 18°17'N, 76°48’W). 293, male palpus, mesal. 294, epigynum, posterior. 295, epigynum, ventral. 296, male, dorsal. 297, male, ventral. 298, female, dorsal. 299, female, ventral. Scale bars: dorsum and venter figures 1.0 mm. 84 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 Yucatan show largely parallel lobes (Levi, 1977: 209, fig. 71). Natural History. Adults have been ob- served on tree trunks and on bushes at and above 150 cm. They can be found in shad- ed areas, which is unusual for Mexican Metepeira. Most mature specimens have been collected in May through September (Fig. 328). Elevations range from sea level to just under 2,000 m. Distribution. Southern Texas to Hon- duras (Map 15). Records Examined. HONDURAS Copan: 14°50'N, 89°9'W, 7.ii1.1939 (AMNH): Ruinas, 14°50’N 89°9'W, 7.iii.1939 (AMNH). MEXICO Campeche: Becan, 18°33'N, 89°30’/W, 31.vii.1991 (W. H. Piel & G. S. Bodner, MCZ); Campeche, 19°51'N, 90°32’W, 14.vii.1948 (C. J. Goodnight, AMNH); Champotén beach, 19°21'N, 90°43’W, 22.vii.1991 (W. H. Piel & G. S. Bodner, MCZ); Seybaplaya, 19°39’N, 90°40’'W, 2.viii.1949 (C. J. Goodnight, AMNH). Chiapas: 16 mi W Cintalpa on rt 190, 16°36’N, 93°51’W, 15.vi.1982 (F. Coyle, MCZ). Nuevo Léon: 20 km E Montemo- relos, Camino Q. Rayones, 25°16’N, 99°41'W, 22.vi.1981 (L. Stange, FSCA): Los Cristales, 25°33’N 100°12'W, 15.viii.1972 (A. F. Archer, AMNH); Villa de Santiago, Hacienda Vista Hermosa, 25°25'N, 100°9'W, 19.vi.1940 (H. Hoogstraal, MCZ); Villa San- tiago, Horsetail Falls, 25°25’N, 100°9'W, 19.vi.1940 (H. Hoogstraal, MCZ). San Luis Potost: 20 km W Cd. Valles, 21°58'N, 99°11’W, 18.iii.1972 (J. A. L. Cooke, AMNH): Cd. Valles, 21°59’N, 99°1’'W, 6.vii.1940 (P. Rau, MCZ), 19.vii.1956 (W. J. Gertsch & V. Roth, AMNH), 15.vii.1959 (L. Steude, AMNH): Cd. Valles, Hotel Covadonga, 21°59'N, 99°1'W (L. Steude, AMNH); Huichichuyan, 21°25'N, 98°55’W, 19.v.1952 (M. Cazier, W. Gertsch, & R. Schrammel, AMNH): Medina, 24°1’N, 100°24'W, 9.ix.1956 (A. F. Archer, AMNH): N section of San Luis Potosi, 22°13’N, 100°58'W, 8.ix.1956 (A. F. Archer, AMNH); Venado Arroyo, 22°56'N, 101°5’W [?], 27.vii.1934 (MCZ). So- nora: 8 mi W Alamos, 29°13'’N, 110°1L0’W, 23.viii. 1965 (W. J. Gertsch & R. Hastings, AMNH). Tamaulipas: 60 km N Cd Valles, 22°30'N, 99°4'W, 10.viii.1991 (W. H. Piel & G. S. Bodner, MCZ): Cd. Victoria, 23°44'N, 99°8’W, 17.v.1952 (M. Cazier, W. Gertsch, & R. Schrammel, AMNH): Laredo road near Cd. Victoria, 23°44'N, 99°8’W, 20.viii.1947 (C. J. & M. Goodnight, AMNH); rl01 26km S Tula, 22°49'N, 99°55’W, 8.ix.1991 (W. H. Piel & G. S. Bod- ner, MCZ): Sisal, 15 mi S Cd. Victoria, 23°38'N, 99°12'W, 22.vii.1966 (Jean & Wilton Ivie, AMNH). Veracruz: 15 mi E Panuco, 22°10'N, 98°3'’W, 29.xi.1941 (A. M. & L. I. Davis, AMNH):; Plan del Rio, 19°6'N, 96°6’W [?], 26.vii. 1956 (W. ile Gertsch & V. Roth, AMNH). Yucatan: 20 km E Valladolid, 20°41'N, 88°2’W, 26.vii.1991 (W. H. Piel & G. S. Bodner, MCZ); 3 km S San Helipers 2iles2uNe 88°14'W, 25.vii.1991 (W. H. Piel & G. S. Bodner, MCZ); 5 mi E Sisal salt flat, 21°9’N, 90°5’W, 9.1.1984 (V. & B. Roth, CAS): Balankanche Cave, 2 km E Chi- chen Itza, 20°40'N, 88°33’W, 19.vii.1983 (W. Mad- dison, MCZ); beach north of Celesttin, 20°56’N 90°21'W, 24.vii.1991 (W. H. Piel & G. S. Bodner, MCZ); Chichén Itza, 20°40'N, 88°34'W (C. J. Good- night, AMNH), 15.vii.1981 (C. Gold, CAS): Cordil- leria Mayapan, 20°28'N, 89°11’W, 6.ix.1952 (J. & D. Pallister, AMNH); Uxmal, 20°22'N, 89°46’W, 7.ix.1970 (A. F. Archer, AMNH), 23.vii.1991 (W. H. Piel & G. S. Bodner, MCZ). USA Texas: 29 mi S Sarita, 26°47’N, 97°47'W, 14.xi.1958 (A. Brady, MCZ); 1 mi S Pharr, 26°10’N, 98°11’W, 14.xi.1958 (A. Brady, MCZ); 1 mi S Pharr on U.S. HW 281, 26°10'N, 98°L1’W, 14.xi.1958 (A. Brady, MCZ). 37. Metepeira pacifica new species Figures 286-292, 302; Map 10 Holotype. Male from La Pacifica, Guanacaste, Costa Rica, 1.ii.1975-2.iii.1975, R. E. Coville, in MCZ. The specific name is a noun in apposition after the locality. Description. Female paratype from La Pacifica, Guanacaste, Costa Rica. Carapace tan, lighter around eyes. Legs white. Fe- mur I with row of three macrosetae on an- terior side; none on anteroventral side. Dorsum of abdomen white with faint, in- distinct folium, darker distally (Fig. 291). Gravid alee often slightly marbled. Venter wide, with long white median line surrounded by faint, thin, white U-shaped line on black. Pair of large white spots on each side of spiracle (Fig. 292). Sternum black with wide, white, uneven median line (Fig. 292). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.2, posterior laterals 1.2. Anterior median eyes separated by 1.2 diameters, posterior median eyes by 0.9, anterior median eyes separated from an- terior laterals by 1.8 diameters of anterior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 4.5 mm. Car- apace 2.2 mm long, 1.7 wide. First femur 2.3 mm, patella and tibia 2.4, metatarsus 1.9, tarsus 0.8. Second patella and tibia 2 mm, third 1.2, fourth 1.8. Male holotype. Carapace, legs, abdo- men as in female, though often darker (Figs. 289, 290). Femur I with row of four Jan Feb July I June gressa, n = 26 July June galatheae, n = 304 Jan ee i Feb i Sep July Apr “May June calamuchita, n = 15 July June datona, n = 66 July June roraima, n= 11 July June revillagigedo, n = 2 July June tarapaca, n = 23 July June crassipes, n= 8 July June jamaicensis, n = 24 July June maya, n= 13 Jan \ Mar \ i | July pacifica, n= 15 Oct Apr June Sep July XC May June rectangula, n= 11 July June nigriventris, n = 44 July glomerabilis, n = 33 Nov al =~ Jes Oct IS June July June inca, n = 29 METEPEIRA ° Piel 85 July wee" June karkii, n = 9 "July June compsa, n = 279 July June arizonica, n = 45 Figures 300-319. Circular histograms depicting relative seasonal abundance of collecting events for mature spiders. 300-308. Primarily collected during the northern hemisphere winter and spring seasons. 300, Metepeira gressa; 301, Metepeira revillagi- gedo; 302, Metepeira pacifica; 303, Metepeira karkii, 304, Metepeira galatheae, 305, Metepeira tarapaca; 306, Metepeira rec- tangula, 307, Metepeira desenderi, 308, Metepeira calamuchita. 309-314. Generally collected throughout. 309, Metepeira cras- sipes; 310, Metepeira nigriventris; 311, Metepeira compsa; 312, Metepeira datona; 313, Metepeira jamaicensis; 314, Metepeira glomerabilis. 315-319. Primarily collected during the Northern Hemisphere summer season. 315, Metepeira cajabamba; 316, Metepeira roraima; 317, Metepeira maya; 318, Metepeira inca; 319, Metepeira arizonica. 86 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 macrosetae on anterior side; two to three on anteroventral side. Median white line on sternum often broken (Fig. 290). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.6, posterior laterals 1.6. Anterior median eyes separated by 1.1 diameters, posterior median eyes by 0.6, anterior median eyes separated from anterior laterals by 1.4 di- ameters of anterior lateral eyes, lateral eyes separated by 0.3 their diameters. To- tal length 3.4 mm. Carapace 1.7 mm long, 1.2 wide. First femur 2.2 mm, patella and tibia 2, metatarsus 1.7, tarsus 0.8. Second patella and tibia 1.6 mm, third 0.9, fourth eS. Diagnosis. Unlike other members of the M. minima species group, M. pacifica and M. jamaicensis share very light pigmenta- tion, and the embolus in both species is needle-shaped (Figs. 286, 289, 291, 293, 296, 298). Unlike M. jamaicensis, the fla- gellae on the median apophysis of M. pa- cifica are set off on a short, wide stalk (Fig. 286) rather than a long, thin one (Fig. 293). A ventral view of the epigynum in M. pacifica shows a ridge under the scape that is more V-shaped (Fig. 288), com- pared to a straighter line (Fig. 295). Variation. Average body length of three females examined 5.3 mm, range 4.5 to 5.8 mm. Average body length of five males ex- amined 3 mm, range 2.8 to 3.4 mm. Most specimens appear whitish, but the loss of pigment is variable, especially among males. Natural History. Specimens have been collected from the wasp nests of Trypar- gilum nitidum, T. tenoctitlan, and T. ben- soni. In Costa Rica, mature specimens have been collected in November through February; Honduras and Nicaragua in July (Fig. 302). Altitudes range from 100 to 2,600 m. Distribution. Costa Rica, Honduras, and Nicaragua (Map 10). Records Examined. COSTA RICA Guanacaste: 4 km NW Cajfias, La Pacifica, prey of Trypargilum ben- soni, 10°27'N, 85°9'W, 291.1975 (R. E. Coville, MCZ); 4 km NW Caijias, La Pacifica, prey of Trypar- gilum nitidum, 10°27'N, 85°9’W, 1.ii.1975—2.iii.1975 (R. E. Coville, MCZ): 4 km NW Canas, La Pacifica, prey of Trypargilum tenoctitlan, 10°27'N, 85°9'W, 1.ii.1975 (R. E. Coville, MCZ); 4 km NW Canas, La Pacifica, wasp collected, 10°27'N, 85°9'W, 16.ii.1975 (R. E. Coville, MCZ); Bagaces, Palo Verde, 10°21’N, 85°21'W, 19.1.1978 (W. Eberhard, MCZ); ca. Cafias, 10°25'N, 85°7'W, 15.xi.1982 (W. Eberhard, MCZ): Canias, 10°25'N, 85°7'W, 15.xi.1982 (W. Eberhard, MCZ). Puntarenas: Finca San Miento-Sialas, 10°9’N, 84°54'W, 5.ii.1976 (Roth & Schroepfer, AMNH). San José [?]: Santa Maria, 9°39'N, 83°58’W, 15.i.1930 (Dodge, MCZ). HONDURAS Tegucigalpa: Teguci- galpa, 14°6’N, 87°13/W, L.vii.1948 (Clarke, AMNH). NICARAGUA Managua: Laguna de Jiloa, SW Ma- nagua, campsite, 12°13'N, 86°19’W, 8.vii.1970 (S. Riechert, SR). 38. Metepeira jamaicensis Archer Figures 293-299, 313; Map 15 Metepeira jamaicensis Archer, 1958: 16, fig. 33, °. Female holotype from Port Henderson, St. Cath- erine Parish, Jamaica, in the AMNH, examined. Metepeira minima:—Levi, 1977: 206, 208. Brignoli, 1983: 275. Erroneous synonymy. Description. F emale from Saint Mary's Parish, Strawberry Fields near Robin’s Bay and Green Castle, Jamaica. Carapace dirty brown, white around eyes, central white wedge (Fig. 298). Legs whitish yellow; slight rings on legs II and III. Femur I with row of three to four macrosetae on anterior side; none or only a few very fine setae on anteroventral side. Dorsal folium lighter than in most species; fleur-de-lis white on speckled light gray (Fig. 298). Venter brownish gray with white margins. Wide median white line with pair of large white spots on either side of spiracle (Fig. 299). Sternum brownish black with wide, white line widening anteriorly, sometimes broken in center (Fig. 299). Ratio of eye diameters: posterior medians and anterior medians 1.0, anterior laterals 1.2, posterior laterals 1.1. Anterior median eyes separat- ed by 1.4 diameters, posterior median eyes by 0.7, anterior median eyes separated from anterior laterals by 2 diameters of an- terior lateral eyes, lateral eyes separated by 0.2 their diameters. Total length 5.1 mm. Carapace 2.1 mm long, 1.5 wide. First fe- mur 2.1 mm, patella and tibia 2.2, meta- 320 Dec Jan Nov Feb Oct Mar Sep Apr Aug May July June ventura, n = 35 324 Dec July olmec, n= 6 328 Dec Oct June Jan Feb Sep July minima, n = 39 May June July June lacandon, n = 10 July atascadero, n = 17 June 321 Dec Jan Nov Feb Oct Mar Sep Apr Aug May July June grandiosa grandiosa, n = 4 July June grandiosa alpina, n= 5 July uncata, n=6 June 333 Dec Jan Nov Feb Oct Mar Sep Apr Aug May July June chilapae, n = 15 337 Dec Jan Nov Feb Oct Mar Sep Apr Aug May July June triangularis, n = 19 July gosoga, n= 2 June July celestun, n= 8 330 Dec June Jan Sep July pimungan, n= 6 June 334 Dec Jan Nov Feb Oct Mar Sep Apr Aug May July June petatlan, n = 3 METEPEIRA ® Piel S87 323 Dec Jan Nov Feb Oct Mar Sep Apr Aug May July June incrassata, n = 33 327 Dec July vigilax, n= 10 June 331 Dec Jan Nov Feb Oct Mar Sep Apr Aug May July June comanche, n = 2 335 Dec Jan Nov Feb Oct Mar Sep Apr Aug May July June spinipes, n= 116 Figures 320-337. Circular histograms depicting relative seasonal abundance of collecting events for mature spiders. 320-331, Primarily collected during the northern hemisphere summer season. 320, Metepeira ventura; 321, Metepeira grandiosa gradiosa; 322, Metepeira gosoga; 323, Metepeira ventura; 324, Metepeira olmec; 325, Metepeira grandiosa alpina; 326, Metepeira celes- tun, 327, Metepeira vigilax, 328, Metepeira minima; 329, Metepeira uncata; 330, Metepeira pimungan, 331, Metepeira comanche. 332-337. Primarily collected during the northern hemisphere fall season. 332, Metepeira lacandon; 333, Metepeira chilapae; 334, Metepeira petatlan; 335, Metepeira spinipes; 336, Metepeira atascadero; 337, Metepeira triangularis. 88 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 tarsus 1.8, tarsus 0.8. Second patella and tibia 1.9 mm, third 1.2, fourth 1.7. Male from same locality as female. Male carapace, dorsum, venter, sternum darker and more contrasty version of female (Figs. 296, 297). All legs ringed. Femur I with row of three macrosetae on anterior side; none on anteroventral side. Ratio of eye diameters: posterior medians and an- terior medians 0.9, anterior laterals 1.2, posterior laterals 1.2. Anterior median eyes separated by 1.3 diameters, posterior median eyes by 0.9, anterior median eyes separated from anterior laterals by 1.2 di- ameters of anterior lateral eyes, lateral eyes separated by 2.3 their diameters. To- tal length 2.3 mm. Carapace 1.2 mm long, 0.9 wide. First femur 1.4 mm, patella and tibia 1.3, metatarsus 1.0, tarsus 0.5. Sec- ond patella and tibia 1.2 mm, third 0.6, fourth 0.9. Diagnosis. Unlike other members of the M. minima species group, M. jamaicensis and M. pacifica share very light pigmen- tation and the embolus in both species is needle-shaped (Figs. 286, 289, 291, 293, 296, 298). Unlike M. pacifica, the flagellae on the median apophysis of M. jamaicensis are set off on a long, thin stalk (Fig. 293) rather than a short, wide one (Fig. 286). A ventral view of the epigynum in M. jamai- censis shows a ridge under the scape that almost forms a straight line (Fig. 295), compared to a V-shape (Fig. 288). Variation. Average body length of elev- en females examined 5 mm, range 4.2 to 6.1 mm. Average body length of four males examined 2.4 mm, range 2.3 to 2.6 mm. Natural History. Mature specimens have been collected in July through March (Fig. 313). Distribution. Primarily in Jamaica and Haiti (Map 15), near sea level. Records Examined. BRITISH WEST INDIES Grand Cayman Island: 19°20'N, 81°10’W, 15.ii.1960 (R. A. Lewin, MCZ). HAITI Departement de L’Ouest: Port-au-Prince, 18°32’N, 72°20'W, 19.vii.1955 (A. F. Archer, AMNBH), 20.vii.1955 (A. F. Archer, AMNH). Dept. de L’Artibonite: Saint-Marc, 19°7'N, 72°42'W, 15.i.1913 (W. M. Mann, MCZ). JA- MAICA Cornwall: Montego Bay, 18°28'N, 77°55’'W, 1.iii.1984 (L. E. Schulten Jr, MCZ). Middlesex: 3 mi E Old Harbor, 17°56'N, 77°10’W, 21.x.1957 (A. M. Chickering, MCZ); Christiana, 18°10'N, 77°29'W, 13.xi.1957 (A. M. Chickering, MCZ), 15.vii.1960 (C. & P. Vaurie, AMNH), 17.vii.1960 (C. & P. Vaurie, AMNH); Strawberry Fields near Robin’s Bay and Green Castle, 18°17'N, 76°48'W, 23.iii.1972 (H. W., L. & F. Levi, MCZ), 25.iii.1972 (H. W., L. & F. Levi, MCZ), 26.iii.1972 (H. W., L. & F. Levi, MCZ). Saint Ann: Roaring River, 18°24'N, 77°9’W [?], 8.ii.1946 (B. Heineman, AMNH); Saint Ann’s Bay, 18°26'N, 77°8'W, 20.xi.1959 (A. M. Nadler, AMNH). Saint Catherine: E Green Harbour, S slope of Healthshire, 17°53'N, 76°51'W, 12.viii.l1958 (A. F. Archer, AMNH). St. Andrews: Ferry, 9/10 mi on Spanishtown Road, 18°2’N, 76°53'W, 26.vii.1955 (A. F. Archer, AMNBH). Surrey: Kingston, Mona Road, pasture, 17°59'N, 76°24'W, 10.x.1957 (A. M. 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The Vegetation of the Galapagos Islands. Zierikzee, The Netherlands: Drukkerij Lakenman & Ochtman. INDEX METEPEIRA ° Piel 91 VIERA, C. 1986. Comportamiento de captura de Me- tepeira sp. A (Araneae, Araneidae) sobre Acro- myrmex sp. (Hymenoptera, Formicidae) en con- diciones experimentales. Arachnologia, 6: 1-8. . 1989. Caracteristicas de la tela orbicular de Metepeira sp. A (Araneae, Araneidae). Boletin de la Sociedsad Zoologica del Uruguay, 5: 5-6. . 1992. Comparacion de telas de hembras y juveniles de Metepeira seditiosa (Araneae, Ara- neidae). Boletin de la Sociedad Zoologica del Uruguay, 7: 17-18. VIERA, C., AND F. G. Costa. 1988. Analisis del com- portamiento de captura de presas por machos adultos de Metepeira sp. A (Araneae, Araneidae), utilizando telas de juveniles y hembras adultas coespecificos. Journal of Arachnology, 16: 141— 152. WISE, D. H. 1983. Competitive mechanisms in a food-limited species: relative importance of in- terference and exploitative interactions among labyrinth spiders (Araneae: Araneidae). Oecolo- gia, 58: 1-9. Valid taxon names are printed in italics. Page numbers in bold refer to illustrations; those in italics refer to species descriptions. acostai, Metepeira 63 Aculepeira 8 alpina, Metepeira 23, 24 Amazonepeira 8 Arachnidomyia 10 Araneus 8 arizonica, Metepeira 9, 19, 65, 66 atascadero, Metepeira 9, 16, 18, 67, 69 bani, Metepeira 63 biogeography 9, 10 cajabamba, Metepeira 9, 12, 17, 26, 31 calamuchita, Metepeira 14, 18, 39, 42 cancriformis, Gasteracantha 10 celestun, Metepeira 9, 10, 16, 19, 74, 75 cereicola, Metepeira 43 chilapae, Metepeira 9, 16, 19, 78, 79 chilapica, Metepeira 78 comanche, Metepeira 14, 19, 60, 61 compsa group, Metepeira 47 compsa, Aranea 48 compsa, Metepeira 7, 9, 10, 11, 14, 17, 48, 49 crassipes, Metepeira 7, 9, 10, 16, 19, 77, 79 Cyrtophora 8, 9 dakota, Metepeira 23, 24 daytona, Metepeira 6, 7, 8, 9, 10, 12, 17, 20 desenderi, Metepeira 6, 10, 12, 17, 19, 21, 25 digital photography 4 dominicana, Metepeira 30 dorsal folium 6 douglasi, Metepeira 34 ensenada, Metepeira 71 epigynum 8 foxi group, Metepeira 19 foxi, Metepeira 8, 9, 10, 12, 17 galatheae, Araneus 43 galatheae, Epeira 43 galatheae, Metepeira 9, 14, 18, 43, 45 glomerabilis, Araneus 28 glomerabilis, Epeira 28 glomerabilis, Metazygia 28 glomerabilis, Metepeira 9, 12, 17, 28, 31 gosoga, Metepeira 14, 18, 59, 61 grandiosa alpina, Metepeira 9, 12, 17, 24, 25 grandiosa grandiosa, Metepeira 9, 10, 12, 17, 23, 25 grandiosa palustris, Metepeira 9, 12, 17 grandiosa, Metepeira 23 gressa, Epeira 54 gressa, Metazygia 54 gressa, Metepeira 9, 18, 54, 57 gressus, Araneus 54 grinnelli, Metepeira 34 habitats 9 inca, Metepeira 6, 14, 17, 18, 57, 58 incrassata group, Metepeira 55 incrassata, Metepeira §, 9, 10, 11, 16, 19, 68, 69 inerma, Metepeira 20 jamaicensis, Metepeira 9, 17, 19, 83, 86 josepha, Metepeira 77 Kaira 6, 8 karkii, Araneus 46 karkii, Metepeira 9, 13, 18, 45, 46 koepckeorum, Araneus 6 92 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 labyrinthea grinnelli, Aranea 34 rectangula, Metepeira 5, 6, 9, 12, 17, 32, 35 labyrinthea grinnelli, Epeira 34 rectangulata, Metepeira 32 labyrinthea grinnelli, Metepeira 34 revillagigedo, Metepeira 16, 19, 73, 75 labyrinthea group, Metepeira 33 roraima, Metepeira 9, 12, 18, 49, 53 labyrinthea, Epeira 21 Saléi, Epeira 68 labyrinthea, Metepeira 7, 9, 10, 14, 17, 32, 34, 46, 48 salei, Metepeira 70 labyrintheus, Araneus 48 sallei, Aranea 70 lacandon, Metepeira 9, 14, 17, 35, 37 sallei, Araneus 70 latigyna, Metepeira 48 santa, Aranea 28 lindae, Arachnidoyia 10 scitulus, Araneus 54 maya, Metepeira 9, 14, 18, 56, 57 seditiosa, Epeira 54 measurements 5 seditiosa, Eustala 54 seditiosa, Metepeira 54 Mecynogea 8, 9 3 seditiosus, Araneus 54 median apophysis § Metepeira 5 Singa 6 minima group, Metepeira 80 Seer ne et 10 As minima, Metepeira 9, 10, 12, 16, 19, 81, 83, 86 spinipes, Araneus 34 ) nigriventris group, Metepeira 38 spinipes, Metepeira 5, 6, 8, 9, 14, 17, 34, 35 nigriventris, Araneus 38 suspended remeat © : nigriventris, Epeira 38 tarapaca, Metepeira 7, 9, 14, 18, 39, 40 nigriventris, Metepeira 9, 10, 11, 12, 18, 38, 39 ae a A wae Shy Ue LS G8), CD ocosingo, Mecynogea 10 GU ores ; olmec “Metepeira 9, 16, 19, 59, 61 uncata, Metepeira 9, 16, 19, 75, 76 ; : ; uncatus, Araneus 76 i ae ee a ed S304 vaurieorum, Metepeira 48 paomata, Metepenes ventura group, Metepeira 71 palp oy = ventura, Metepeira 10, 16, 19, 69, 71 palustris, Metepeira ao vigilax group, Metepeira 26 perezi, Metepeira 48 vigilax, Araneus 30 petatlan, Metepeira 16, 19, 79, SO vigilax, Epeira 29 pimungan, Metepeira 8, 9, 16, 19, 62, 65 vigilax, Metepeira 9, 10, 12, 17, 26, 30, 31 predation 10 virginensis, Metepeira 48 rayorae, Arachnidomyia 10 web 8 rectangula, Epeira 32 Zygiella 6 idee Pe Pe at its Piet i Wily ‘Type Specimens of Recent Mammals in the Museum of Comparative Zoology K. M. HELGEN ANDT. L. McFADDEN na 9 mf) ~y th U qg cvwy = ND fe Al GD LA A TUNG : paw 4 Gt 3 Bs fy oe a. Pees ‘ a A ee OP LS 4 Dee \ \ 5 { | HARVARD UNIVERSITY VOLUME 157, NUMBER 2 _ CAMBRIDGE, MASSACHUSETTS, U.S.A. 8 JUNE 2001 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY BrevioraA 1952— BULLETIN 1863— Memoirs 1865-1938 Jounsonia, Department of Mollusks, 1941-1974 | OccCASIONAL PAPERS ON MoLLusks, 1945— SPECIAL 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 ‘llinsietisd Catalogue of the Tere- . dinidea (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4. Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology 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 of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the 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 of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter's Check-list of Birds of the World, vols. 1-16. es of the New England Zoological Club 1899-1947. (Complete sets only.) Proceedings of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2001. TYPE SPECIMENS OF RECENT MAMMALS IN THE MUSEUM OF COMPARATIVE ZOOLOGY K. M. HELGEN AND T. L. MCFADDEN' CONTENTS INO Stacia oes Ue CE EES ba Ps SAR ee 94 MintnOGuctiOny eats hich ys se Lunes Sibi VU 94 Authors of (ivpesD escripuonss ae 94 Definitions and Organization 96 Taxa Included in This Catalogue __............. 97 ANG SRREN ANC Woy aR Oe he ee 98 @ISINON AG GAGES Se eee 98 Pareallecosy oe SEES Ga 98 mecounts Of Type Specimens 2 99 Order Didelphimeonpiiay se aes 99 HamilysCalucouyidae ees 99 amotly,Maxmiosidact 222.2). eee 99 HannilyaDicdel plidac hase tse = ses eee 100 Order Dasyuromorphia —_......-.-.---------------- 100 [Seilh\y IDE tials: A 100 OxdemPerame lia ee ee 100 Hamnilyaeeronyctidac sss ka sleelee olla 100 Ordem@ingulatae ee sais wearer 101 HamulyDasypodidacs a ee 101 OrdemAtnosoncida == = ame ee 101 Hamnily Chrysochlonidae == a. - 101 Hannilyem@emkecidae 2:0 ee 101 Orders Rodentiay= ==. ee ee 103 HamiulyeAplodontidacy== 2s" 2 ssnurs ls 103 HamulysSciumGac, a. sai e te Be 103 Hamulya@astondac ge 2. Bees 109 anulvaGcommidac. =. sere) eee 110 annilvalteteromiyidae sess eee ee 111 lSeneotllhy ID Mhovorettoleyss Was 112 I Semanullhy, Ilona es: eee Ea) Sulotfannilysarvicolimae sus eenen nee ILD Sulbtaunilyz@nicetinac =e ees LIE Subfamily Cricetomyinae ____..-_- LAL Subfamily Dendromurinae —__-- LAL Subfamulya@erbillinae = ILE 7e SWlonehoanllhys IY Munabieves Se 117 Subtamuily Nesomyimae 9 == 120 Subtamulya@ tomy aes eee ee 120 Subfamily Sigmodontinae _....._....-.-...- 121 Hamnilyakedetidac)s ==) eee 128 ‘Mammal Department, Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- setts 02138. HamnilySNiyOxiG. ae ee eeeeete sae En mL anenun me HamulysBathyercidac =u ln stm aes amnilyeHraithiZomtiGdacr re ear eee HamilysDasyproctidac: 2 = a ane HamnilyeAcoutiGaey= sane fae es ae rneE ee Harnily; Octodomtidacise= set ee arailyais chimayi dace. eels eee n a HamnilysCapnomnyi dae jes ee Orders Bagomompnay se ee ee Mamnilys@chotomidae = == snsss men Benrmllyy JUsteroa CHS) oe ee Order Scandentiay ee arnilyaslupaidacy ues asses eaeeeneen Ovrcler IPatmbiveg a Hammill dae gas oe, oe eee en ee careers HamuilysDaubentoniidac) 9. sss os HannilysG alagonidac sass: eee eee Hamnilya@ebidac geass eo ee eee: Hamulyaetylobatidac ttm. semen ie ee EamnilyarlomuniGd acy 22s salsa ine eee @rderplaipotyplila ss8 os eae eee Hamily)Nesophontidac (22. ts ee Family Solenodontidae 02.2228 [stangaullyy: SYO;atGNG Vs) Ue ee al tenooulby IRalhonGhys)) es eee Order Chiropterass -iiey i ees amilyaPteropodidae) = aasaes .. slr aasnmens anuly SH mlbjallonuridac) parents. ero Jenene hyowerate evenee eS a Hamilynninolophidac === == HannilyaNormnoopidac === s=aesananenenas Hamulysehyllostomidac === === HamailygMolossiclac yams see sean HamulyaVespenrtilionidae) =a: sass aa mewaavilye TU oyjanoyoueratolevey Se et Order Antiodac ty ats. ss eee nee Family Tayassuidae ESR ORY Ae EAE ES Hamuilya Monod ontidac yas es seen Hannilyathocoemicd ac eee maen aaleene Iaeioalhy CRIA: a IRlemodtly” JEN ANCEYS ie es Orrgler Carnivora) se Banari (CRNA YS) op HanailyaWinsidac heen senate: seen eece sees Bull. Mus. Comp. Zool., 157(2): 93-181, June, 2001 94 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Family Procyonidae PSS late Aare Nc nate TO 151 HamulvaMiustelicacmems a: sa ennasenenes 152 |@eronvilhy: IN iejo) owt ORS 154 | Bertani hyn WaKeranKC ENS) Qo 155 Mamaily Eero eS ti ac aes eee 155 Benoa iy JSC RS =a 155 OxdemEinolestaya eee 156 HanailyaNiani ac aes ass see ee eee ane 156 RE TENE TNC Sp eisie a tas tees a cect ce an ORE COLE IN 157 IIFas Vey: co poh Bee SN Wf eer ets ties etal Tae Toes cha 171 ABSTRACT. The Mammal Department at the Mu- seum of Comparative Zoology houses name-bearing types of 342 species-group taxa of Recent mammals. The type collection consists of 327 holotypes, 2 lec- totypes, 4 complete syntype series, and 9 partial syn- type series. This catalogue notes infor mation on the type locality, collector, dae of collection, present con- dition, original publication, and synonyms for all name-bearing types in the Mammal Department. Comments on the taxonomic and historical impor- tance of many type specimens are included. Lecto- types for a number of taxa are designated for pur- poses of taxonomic consistency. INTRODUCTION “Some of them are more or less historic specimens, ” wrote the gteat mammalogist Glover M. Allen in 1931, “whose location has undoubtedly in many cases been lost sight of, so that it may be of value to make the present record” (1931: 230). Allen, then Curator of Mammals at the Museum of Comparative Zoology at Harvard Uni- versity, was referring to the type speci- mens of mammals in the museum, the subject of a listing he published that year. Allen’s words resound with even greater truth today than when they were written 70 years ago. Since that time, the muse- um’s holdings of mammal type specimens have grown considerably. Furthermore, type specimens of mammals housed in the MCZ have sometimes been credited to the collections of other institutions or consid- ered to no longer exist, making the need for a new catalogue obvious. Type specimens are biological reference points that lend objectivity to taxonomy and are thus of critical importance in sys- tematic investigations; accordingly, “every institution in which name-bearing types are deposited should publish lists Be name- bearing types in its possession or custody” (International Commission on Zoological Nomenclature, 1999: 79). Our duty in this regard is thus long overdue, and this cat- alogue should serve to fulfill that obliga- tion. Without doubt, it will alleviate con- fusion in the scientific community as to the whereabouts of certain specimens, thought to be lost, especially those type specimens that were acquired from Guillaume Gran- didier. In addition, it should bring to light information, formerly unavailable, on this museum's very notable holdings of name- bearing types. All taxonomic judgments in this work, including new name combinations and lec- totype designations, reflect the decision of the first author (KMH) alone and should be cited accordingly. AUTHORS OF TYPE DESCRIPTIONS Four mammalogists authored the over- whelming majority of names based on type specimens in the MCZ: Glover Allen, Out- ram Bangs, Guillaume Grandidier, and Barbara Lawrence. Following are brief bi- ographies of these four outstanding mam- malogists. Glover M. Allen (1879-1942) Curator of Mammals at the MCZ from 1924 until his death in 1942, Glover Allen began his work in the Mammal Depart- ment in 1907. He was known as a careful, dependable researcher and an outstand- ing, patient teacher. His goal in all en- deavors was to increase the sum total of knowledge about the mammals of the world. Although much of his career was spent in the museum studying, as he put it, “the dried remains of animals” (Barbour et al., 1943: 300), he also traveled widely throughout the world collecting specimens as well as observing and learning about liv- ing animals. He once commented that the actual knowledge of living creatures could all too often be summed up by saying “when we found it, it ran like hell, where- upon we shot it!” (Barbour et al., 1943: 300). He wrote prolifically; his bibliogra- phy of publications is 81 pages long, the TYPE SPECIMENS OF RECENT MAMMALS °* Helgen and McFadden 95 first of which he published when he was only 11 years old (Lawrence 1947a: 1). Holotypes for 96 taxa, which Allen de- scribed alone or with colleagues, are de- posited in the MCZ. Outram Bangs (1863-1932) Outram Bangs, Curator of Mammals from 1899 to 1924, “was one of those for- tunate mortals, born with a love of nature and the outdoors which rule their entire lives” (Peters 1933: 265). Bangs authored 135 MCZ mammal names (only one co- authored!). He and brother Edward began collecting as boys using slingshots and horsehair nooses. He was an early ecolo- gist, saving Microtus breweri by killing the feral cat population on Massachusetts’ Muskeget Island and restocking the mouse from a tiny islet across the channel. Al- though his greatest passion was the natural history of birds, he was also fascinated with mammals and served as curator of both departments in the MCZ. He decided to systematically collect the mammals of east- ern North America in about 1890. He be- gan by trapping in New England and later made trips to the southeastern United States and Canada. Other collectors assist- ed Bangs, expanding the collection area to western North America and south to Cen- tral America. His precision and organiza- tion were legendary, and today his mam- mal collection of more than 10,000 speci- mens, donated to the Museum in 1899, re- mains one of the best curated and most informative in the department. A complete list of Bangs’ scientific publications was compiled by Porter (1943). Guillaume Grandidier (1873-1957) French explorer and scientist Guillaume Grandidier, son of naturalist Alfred Gran- didier, authored and coauthored descrip- tions of 13 taxa whose types were donated to the MCZ in 1947, along with his exten- sive personal collection of. Malagasy mam- mals. The collection was purchased by Robert Barbour and donated to the MCZ in honor of his brother, MCZ director Thomas Barbour. Between 1898 and 1902, Grandidier explored the center and south- ern portions of Madagascar, collecting specimens and describing the geography of the area. Through his writings he brought the unique fauna of this remote region to the attention of the scientific world. He was known for his devotion to the careful acquisition of knowledge, whether it was geographical, historical, or scientific. His work was honored by Bonn the scientific community and the French government (Chapus, 1953). Barbara Lawrence (1909-97) After graduating from Vassar College in 1931, Barbara Lawrence became a volun- teer at the MCZ. She was encouraged by Dr. Glover Allen to do her own research, and in the late 1930s she made field trips to the Philippines and Sumatra to collect mammals. In 1952 she was appointed Cu- rator of Mammals, a position she held un- til her retirement in 1976. Her areas of scientific interest were many, ranging from echolocation in whales to zooarchaeology, as well as the more traditional mammalog- ical pursuit of taxonomy. “She once wrote . to know and love a bit of the world so well that you can give it to someone else . is a rare talent’ ” (Rutzmoser, 1999: 1049), certainly a talent Barbara Lawrence had in abundance. She authored or coau- thored 20 MCZ names. The specimens described by these au- thors and others are the result of world- wide collecting by numerous expeditions and individuals, listed in the accounts that follow. A handful of type specimens in the MCZ were formerly in the collection of the National Museum of Natural History in Washington, D.C. According to Glover Allen, these were obtained via exchanges “at a time when ‘duplicates’ were more freely disposed of, |and would] prove to be cotypes or even actual types” (1931: 229). The collections of the Boston Society of Natural History, once contained in the Boston Museum of Science, were trans- 96 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 ferred to the Museum of Comparative Zo- ology over the course of the 20th century. Several types are among these specimens, which consist primarily of mammals from the New England region. DEFINITIONS AND ORGANIZATION Type Categories Type specimens are categorized as one of the following: Holotype. The single specimen desig- nated as name-bearer in the original pub- lished description of the species-group tax- on (Article 73.1, International Commission on Zoological Nomenclature, 1999: 79). Syntype. One of multiple specimens on which a species-group name is equally based, when no holotype is specified in the original description and no subsequent designation of a lectotype has been pub- lished (Article 73.2, International Com- mission on Zoological Nomenclature, 1999: 81). Lectotype. One of multiple specimens upon which a species-group name is orig- inally based, designated in a publication subsequent to the original description to become the unique name-bearer (Article 74, International Commission on Zoologi- cal Nomenclature, 1999: 82). Neotype. A specimen chosen as_ the name-bearer of a species-group taxon, “if no holotype, lectotype, syntype, or prior neotype is believed to exist” (Article 75, International Commission on Zoological Nomenclature, 1999: 84). There are no neotypes in the MCZ mammal collection. Paratype. A specimen other than the holotype (if designated originally) that is mentioned in the original description of a species-group taxon. Paratypes of species- group taxa whose holotypes are housed in other institutions are not mentioned in this catalogue, although many such specimens exist in this museum. Paralectotype. If no holotype is desig- nated, a specimen other than the lectotype (if designated subsequently) that is men- tioned in the original description of a spe- cies-group taxon. Paralectotypes of spe- cies-group taxa whose lectotypes are housed in other institutions are discussed below, before the accounts for name-bear- ng types. Locality This category includes country, second- level geopolitical division (state, depart- ment, territory, province, or district), col- lection site, and altitude where available. The geographic name given for a type lo- cality is that found in the original publi- cation. When the description of the col- lection locality does not include current geopolitical divisions, that information is provided in parentheses. If an original name is no longer used, an equals sign (=) is included within the parentheses to des- ignate an equivalent modern name. Where altitude was originally given in feet, it has been converted to meters and included in parentheses. Sources used for the current names are from the most current available Gazetteers of the United States Board on Geographic Names, the 10th comprehensive edition of the Times Atlas of the World, the Colum- bia Gazetteer of the World, and the Or- nithological Gazetteers of South America. Account Organization The type locality, collector, date of col- lection, and present condition of each specimen are noted. The publication of the original description is cited for each specimen. Many names have changed in rank or synonymy since their origin; in these cases, the name by which a taxon is known today is noted, with a citation of the publication in which that name combina- tion was first employed for that taxon. Comments are offered for most entries to provide additional information or to dispel potential sources of confusion. The format of this catalogue is largely borrowed from the most recent type cat- alogue of mammals in the American Mu- seum of Natural History (Lawrence, 1993). For systematic consistency, the tax- TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Of onomic judgments at the species-level by the authors of the chapters in Mammal Species of the World (Wilson and Reeder, 1993) are largely adhered to. However, in many cases alternate views are explored, and subsequent work by other authors is noted. The sequence of mammalian orders presented here is as follows: Didelphimor- phia, Dasyuromorphia, Peramelia, Cingu- lata, Afrosoricida, Rodentia, Lagomorpha, Scandentia, Primates, Lipotyphla, Chirop- tera, Artiodactyla, Carnivora, and Cimoles- ta. This sequence represents the ongoing understanding of higher mammalian rela- tionships being produced by research in molecular phylogenetics (Waddell et al., 1999) as well as paleontological studies (McKenna and Bell, 1997). A number of these names are rather nontraditional; use of the names Cingulata (for armadillos) and Cimolesta (for pangolins) at ordinal rank follows McKenna and Bell (1997). Af- rosoricida is used for an order including tenrecs and golden-moles (following Stan- hope et al., 1998: 9971-9972). We use the ordinal name Lipotyphla in a restricted sense to refer to the Recent families Ne- sophontidae, Solenodontidae, Soricidae, and Talpidae (others have used the term “Eulipotyphla,” e.g., Waddell et al., 1999). Cetaceans are included here within the or- der Artiodactyla (an assemblage often re- ferred to as “Cetartiodactyla” in recent lit- erature; see Graur et al., 1997). To avoid unnecessary complexity, or- ders, families, and genera are the only ranks above the level of species that are listed, except for the large family Muridae, for which subfamilial distinctions are pro- vided, in alphabetical order. Within each order, the sequence of families generally follows Simpson (1945), but Wilson and Reeder’s (1993) order of rodent families and Simmons’ (1998: 12) arrangement of the bats are observed. Within each genus, taxa are presented in alphabetical order by original name. The format of this catalogue is as fol- lows, with all the following information provided when possible: Original binomen. Name of describer, date of description. Citation of original publication. =Presently used name, if different from original. Citation of publication in which this name combination was initially used for this taxon. Type Category. Number of specimen.’ Preparation of specimen (skin, skull, alco- hol, etc.), age and sex. Locality. Type locality. Date of collec- tion. Collector. Name of collector. Original number of specimen. Condition. Current condition of the type material. Type Series. Any paratypes, paralecto- types, or additional syntypes in existence are mentioned, with their preparation, sex, and age. Comments. Additional comments re- garding the systematic status or the history of the specimen. TAXA INCLUDED IN THIS CATALOGUE Unlike Allen’s original type catalogue, fos- sil mammals are not considered here, but type specimens of Recent mammals known only from subfossil remains are discussed. Several type specimens of Recent mam- mals that were included in Allen’s cata- logue are not considered here as name- bearing types. A number of syntype series in the MCZ have since Allen’s time been rendered paralectotypes by the designa- tion of a lectotype preserved in another institution; these are discussed in the sec- tion below on paralectotypes. Additionally, MCZ 14929, listed as a “cotype” of Nyc- teris revoili Robin, 1881 by G. M. Allen (1931: 235), is a paratype rather than a syntype and is not considered here. * Mammal specimens in the MCZ bear any of three kinds of numbers. A number preceded by “MCZ” can be found in the general collection in the Mammal Department. A number preceded by a “B” is part of the collection of E. A. and O. Bangs, also housed in the Mammal Department. A number preceded by “VP” designates that the specimen is stored in the Vertebrate Paleontology Department. 98 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 One other enigmatic specimen in the col- lection deserves some mention. When it was received by the Mammal Department, Guil- laume Grandidier’s personal collection con- tained specimens marked as types for 16 taxa, described by either Grandidier or M. L. Lavauden. Fifteen of these holotypes are discussed within this catalogue in separate accounts. The remaining specimen, a fruit bat, MCZ 45073, is marked exactly like the other type specimens, and both its skin and its skull tags bear the name “Eidolon saka- lava nov. spec., G. Grandidier” in the script- ed handwriting typical of Grandidier’s spec- imens. The locality given for the specimen is Ankavandra, in west-central Madagascar. Though this specimen has been curated as a type specimen, I (KMH) can find no pub- lished description of this taxon or any ref- erence to this name in the literature. This name is therefore invalid, barring a discov- ery in the future that it was indeed pub- lished and has since been overlooked. What- ever the published status of the name, it ap- pears to me that this specimen should be considered as a young specimen of Eidolon dupreanum rather than a distinct species, and discussion of this mysterious binomial here should in no way be construed as a formal description of a new taxon. ABBREVIATIONS Abbreviations are used in the text to designate the following institutions. AMNH American Museum of Natural History, New York BMNH_ Natural History Museum, Lon- don BSNH — Boston Society of Natural His- tory, Boston FMNH_ Field Museum of Natural His- tory, Chicago MNHN Muséum National d Histoire Naturelle, Paris MVZ Museum of Vertebrate Zoology, Berkeley, California RMNH_ Rijksmuseum van Natuurlijke Historie, Leiden USNM_ National Museum of Natural History, Washington, D.C. YPM Yale Peabody Museum of Nat- ural History, New Haven, Con- necticut ACKNOWLEDGMENTS We would like to thank the faculty and staff of the Mammal Department at the MCZ—A. W. Crompton, Andrew Biewe- ner, Judith Chupasko, Jane Harrison, and especially Maria Rutzmoser for access to the type specimens. Mary Sears, Timothy McNeece, and Ronnie Broadfoot at the Ernst Mayr Library, Harvard University, were extremely helpful in pinning down difficult references. We also thank Alison Pirie for allowing us to make use of the MCZ Ornithology Department locality references; Carolyn Kirdahy of the Boston Museum of Science, who granted us ac- cess to the archives of the Boston Society of Natural History; and the many individ- uals associated with other museums who assisted by answering any questions that arose. Finally, we are thankful for the sug- gestions we received from two anonymous reviewers, which helped us improve this catalogue. PARALECTOTYPE SERIES Hesperomys eremicus Baird, 1858 =Peromyscus er- emicus eremicus (Baird, 1858). Lectotype, USNM 2575, designated by Osgood (1909: 241). MCZ 4310 and 5273 are paralectotypes. Neotoma fuscipes Baird, 1858 =Neotoma fuscipes fus- cipes Baird, 1858. Lectotype, USNM 22026, des- ignated by Lyon and Osgood (1909: 99). MCZ 4336 and 5264 are paralectotypes. Mus bairdii Hoy and Kennicott, 1857 =Peromyscus maniculatus bairdii (Hoy and Kennicott, 1857). Lectotype, number 750 in the Collection of the Academy of Natural Sciences of Philadelphia, des- ignated by Osgood (1909: 80). MCZ 8073 is a par- alectotype. Pteropus lanigera [sic] H. Allen, 1890 =Pteropus in- sularis Hombron and Jacquinot, 1842. The cor- rected spelling of the original name is Pteropus lan- iger (see Andersen 1912: 297). H. Allen based the description of Pteropus laniger on two syntypes, USNM 19066 (skin)/37815 (skull) and MCZ 7023, a skin. Andersen (1912: 297-298) was unaware of TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden 99 the whereabouts of this latter specimen and based his evaluation of the systematic status of P. laniger solely on the USNM specimen in his taxonomic review of the Megachiroptera. Because it is rep- resented by both a skin and a skull in good con- dition and because it has been used in past taxo- nomic treatments, USNM 19066/37815 is hereby designated as the lectotype of Pteropus laniger to ensure consistency between past and future taxo- nomic treatments of this name. MCZ 7023 is thus a paralectotype. The type locality of laniger is the Caroline Islands, as emended by Andersen (1912: 298), not Samoa, as originally described. Sciurus castanotus Baird, 1855 and Sciurus castan- onotus Baird, 1858 =Sciurus aberti aberti Wood- house, 1853. Baird (1858: 266) noted that the name castanotus, used in his original description of this taxon, was a misprint for castanonotus. Lectotype, USNM 121/1107, designated by Lyon and Osgood (1909: 183). Though not explicitly stated by Lyon and Osgood, this specimen should serve as a lec- totype for both names (Sciwrus castanotus Baird, 1855 and Sciuwrus castanonotus Baird, 1858), for taxonomic consistency. MCZ 4692 is a paralecto- type. Spermophilus obsoletus Kennicott, 1863 =Spermo- philus spilosoma obsoletus Kennicott, 1863. Lec- totype, USNM 3222/27998, designated by A. H. Howell (1938: 130). See G. M. Allen (1931: 252) for a list of paralectotypes (then considered synty- pes) in the MCZ. Spermophilus parryi var. kodiacensis J. A. Allen, 1874 =Spermophilus parryi kodiacensis J]. A. Allen, 1874. Lectotype, USNM 9242/38543, designated by A. H. Howell (1938: 103). See G. M. Allen (1931: 252) for a list of paralectotypes (then con- sidered syntypes) in the MCZ. Spermophilus tridecemlineatus var. pallidus J. A. Al- len, 1874 =Spermophilus tridecemlineatus pallidus J. A. Allen, 1874. Lectotype, USNM 16237, des- ignated by A. H. Howell (1938: 112). See G. M. Allen (1931: 253) for a list of paralectotypes (then considered syntypes) in the MCZ. Tamias quadrivittatus var. pallidus J. A. Allen, 1874 =Tamias minimus pallidus J. A. Allen, 1874. Lec- totype, USNM _ 11656/38311, designated by Cary (1906: 88). G. M. Allen (1931: 255) provided a list of paralectotypes. ACCOUNTS OF NAME-BEARING TYPE SPECIMENS Order DIDELPHIMORPHIA Gill, 1872 Family CALUROMYIDAE Kirsch and Reig, 1977 Genus CALUROMYS J. A. Allen, 1900 Philander cicur Bangs, 1898k Proc. Biol. Soc. Washington, 12: 161, 10 August. =Caluromys lanatus cicur (Bangs, 1898). See Cabrera (1958: 2). Holotype. BS8114. Skin and skull. Adult female. Locality. Colombia: (Magdalena), Santa Marta Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 27 March 1898. Collector. W. W. Brown, Jr. Original number 123. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; B8036, skin and skull, adult male; B8115, skin and skull, adult male; B8116, skin and skull, adult male. Family MARMOSIDAE Hershkovitz, 1992 Genus MARMOSA Gray, 1821 Marmosa robinsoni Bangs, 1898i Proc. Biol. Soc. Washington, 12: 95, 30 April. =Marmosa robinsoni robinsoni Bangs, 1898. See Cabrera (1958: 24). Holotype. B7749. Skin and skull. Adult male. Locality. Venezuela: (Nueva Esparta), Margarita Island. 12 July 1895. Collector. W. Robinson. Original number 506. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 paratypes, both in the USNM; USNM 63209, skin and skull, adult male; USNM 63210, skin and skull, adult female. Comments. M. robinsoni was considered a valid species by Gardner (1993: 18) and Nowak (1999: DIL), Marmosa mitis Bangs, 1898k Proc. Biol. Soc. Washington, 12: 162, 10 August. =Marmosa robinsoni robinsoni Bangs, 1898. See Cabrera (1958: 24). Holotype. B8123. Skin and skull. Adult male. Locality. Colombia: (Magdalena), Santa Marta Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 25 March 1898. Collector. W. W. Brown, Jr. Original number 91. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 26 paratypes; B8117—B8122, B8124— B8143; all represented by skin and skull, 15 fe- males and 11 males. 5 paratypes are no longer in the MCZ (BS8118 is at Wellesley College, B8124 and B8141 are at FMNH, and B8136 and B8138 are at USNM). 100 Marmosa fulviventer Bangs, 1901b Amer. Nat., 35: 632, 22 August. =Marmosa robinsoni fulviventer Bangs, 1901. See Handley (1966: 775). Holotype. B8435. Skin and skull. Adult male. Locality. Panama: (Panama), Gulf of Panama, San Miguel Island. 28 April 1900. Collector. W. W. Brown, Jr. Original number 123. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; B8436, skin and skull, fe- male (exchanged to FMNH in 1931); B8437, skin and skull, female; B8438, skin and skull, male. Comments. M. r. fulviventer was retained as a valid subspecies by Hall (1981: 14) and O'Connell (1983: 1). Family DIDELPHIDAE Gray, 1821 Genus D/IDELPHIS Linnaeus, 1758 Didelphis marsupialis particeps Goldman, 1917 Proc. Biol. Soc. Washington, 30: 107, 23 May. Holotype. B8439. Skin and skull. Adult male. Locality. Panama: (Panama), Gulf of Panama, San Miguel Island. 8 May 1900. Collector. W. W. Brown, Jr. Original number 165. Condition. Skin and skull complete. Type Series. 1 paratype; B8440, skin and _ skull, adult female. Comments. Retained as a valid subspecies by Hall (1981: 4). Didelphis virginiana pigra Bangs, 1898b Proc: Boston’ Soc. Nat. Hist., 28: 172, 15 March. Holotype. B3500. Skin and skull. Adult female. Locality. (United States): Florida, Brevard County, Oak Lodge, East Peninsula opposite Micco. 31 Jan- uary 1895. Collector. O. Bangs. Condition. Skin and skull complete. Type Series. 11 paratypes; all represented by skin and skull, most still in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 5) and McManus (1974: 1). The type de- scription lists 31 January 1896 as the date of col- lection, but the date is written as “January 31, 1895” on the original specimen label and in Bangs’ accession catalogue. Order DASYUROMORPHIA Gill, 1872 Family DASYURIDAE Goldfuss, 1820 Genus ANTECHINUS Macleay, 1841 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Antechinus mayeri misim Tate, 1947 Bull. Amer. Mus. Nat. Hist., 88: 130, 20 February. =Antechinus naso misim Tate, 1947. See Laurie and Hill (1954: 7). Holotype. MCZ 29924. Skin and skull. Adult male. Locality. Papua New Guinea: Morobe Province, Mount Misim (=Missim), 5,850 ft (1,784 m). 14 April 1933. Collector. H. Stevens. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; MCZ 29923, skin and skull, adult female. Comments. Tate, in the original description, erro- neously listed 24 April 1933 as the date of collec- tion. A. n. misim was retained as a valid subspecies by Flannery (1995a: 80), who also noted that the New Guinean species assigned to the genus Ante- chinus are not closely related to the Australian spe- cies of that genus and will be reassigned at the generic level pending a full taxonomic revision of the group. Genus MYOICTIS Gray, 1858 Myoictis melas wavicus Tate, 1947 Bull. Amer. Mus. Nat. Hist., 88: 140, 20 February. =Myoictis melas wallacei Gray, 1858. See Flannery (1990: 56). Holotype. MCZ 28082. Skin and skull. Adult male. Locality. Papua New Guinea (S. Morobe), Wau, 3,800 ft (1,159 m). 27 March 1932. Collector. H. Stevens. Original number 1. Condition. Skin and skull complete. Type Series. Holotype only. Comments. Stevens’ original field label bears the following lament—*To my intense anguish I failed to retrieve the female, shot on recumbent, decayed log in undergrowth.” Order PERAMELIA Ameghino, 1889 Family PERORYCTIDAE Groves and Flannery, 1990 Genus ECHYMIPERA Lesson, 1842 Suillomeles hispida G. M. Allen and Barbour, 1909 Proc. New England Zool. Club, 4: 44, 12 July. =Echymipera kalubu kalubu (Fischer, 1829). See Laurie and Hill (1954: 11). Holotype. MCZ 7006. Skin and skull. Adult. TYPE SPECIMENS OF RECENT MAMMALS ® Helgen and McFadden Locality. (Indonesia): Dutch New Guinea (=Irian Jaya), Doreh Bay, Manokwari, “not far from the foot of Mt. Arfak.” 23 February 1907. Collector. T. Barbour. Condition. Skin complete. Skull partial (occipital region and two posterior upper molars on each side missing). Type Series. Holotype only. Comments. S. hispida is the type species of Swil- lomeles G. M. Allen and Barbour, 1909. Echymipera rufescens australis Tate, 1948 Bull. Amer. Mus. Nat. Hist., 92: 334, 25 November. Holotype. MCZ 29214. Skin and skull. Adult male. Locality. Australia: Queensland, Cape York, near Coen, east slope of McIlwraith Ranges, Rocky Riv- er, “Rocky Scrub.” 20 June 1932. Collector. P. J. Darlington, Jr., Harvard Australian Expedition. Original number 209. Condition. Skin complete, but tip of tail worn. Skull complete. Type Series. Holotype only. Comments. Tate (1952: 582) reported that the ex- act type locality is “in the dense rain forests of the upper Nesbit River on the east slopes of the Mcllwraith Range.” Retained as a valid subspecies by Flannery (1995a: 111). Darlington noted in his field notebook, regarding this specimen, “Animal about the fattest I have skinned—I shall dream of it!” Order CINGULATA Illiger, 1911 Family DASYPODIDAE Gray, 1821 Genus DASYPUS Linnaeus, 1758 Dasypus novemcinctus hoplites G. M. Allen, 1911a Bull. Mus. Comp. Zool., 54: 195, July. Holotype. MCZ 8116. Skin, skull, and_posteranial skeleton. Adult female. Locality. Grenada: hills back of Gouyave. 7 Sep- tember 1910. Collector. G. M. Allen. Original number 26. Condition. Skin, skull, and postcranial skeleton complete. Type Series. 2 paratypes; MCZ 8117, skin and skull, adult male; MCZ 8118, skin and skull, adult male. Comments. Retained as a valid subspecies by Hall (1981: 283) and McBee and Baker (1982: 1). 101 Order AFROSORICIDA Stanhope et al., 1998 Family CHRYSOCHLORIDAE Gray, 1825 Genus CHRYSOCHLORIS Lacépéde, 1799 Chlorotalpa tropicalis G. M. Allen and Loveridge, 1927 Proc. Boston Soc. Nat. Hist., 38: 418, 23 December. =Chrysochloris stuhlmanni tropicalis (G. M. Allen and Loveridge, 1927). See Meester (1974: 3). Holotype. MCZ 22435. Skin and skull. Adult female. Locality. Tanganyika Territory (=Tanzania): Ulu- guru Mountains, Bagilo. 5 October 1926. Collector. A. Loveridge. Condition. Skin complete. Skull slightly damaged (coronoid processes missing from both mandibular rami). Mandible disarticulated. Type Series. Holotype only. Comments. Included in Chrysochloris stwhlmanni by Meester (1974: 3) but recognized as distinct by Simonetta (1968: 42). Hutterer (1993: 75) noted that the systematic status of tropicalis merits fur- ther study. Family TENRECIDAE Gray, 1821 Genus GEOGALE Milne-Edwards and A. Grandidier, 1872 Cryptogale australis G. Grandidier, 1928 Bull. Mus. Hist. Nat. Paris, 34: 64, 26 January. = Geogale aurita Milne-Edwards and A. Grandidier, 1872. See Genest and Petter GiOi5:23)) Holotype. MCZ 45057. Skull fragments. Locality. Madagascar: (Toliary), south of Fort Dau- phin (=Tolanaro), Andrahomana grotto. 1927. Collector. R. Decary. Condition. MCZ 45047 includes 18 partial crania and § mandibular rami. Fragmentary. Type Series. All the type material of C. australis bears a single accession number. Comments. C. australis is the type species of the genus Cryptogale G. Grandidier, 1928. Geogale aurita orientalis G. Grandidier and Petit, 1930 Faune des Colonies Frangaises, 4: 446. Holotype. MCZ 45660. Body in alcohol, cranium sep- arate. 102 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Locality. Madagascar: (Toamasina), east coast, Fe- nerive (=Fenoarivo Atsinanana). April 1928. Collector. R. Decary. Original number 12. Condition. Alcoholic, cranium complete. Type Series. Holotype only. Comments. Retained as a valid subspecies by Ge- nest and Petter (1975: 3). The holotype of G. a. orientalis seems to be the only record of Geogale aurita from the east coast of Madagascar. Genus MICROGALE Thomas, 1882 Microgale decaryi G. Grandidier, 1928 Bull. Mus. Hist. Nat. Paris, 34: 69, 26 January. =Microgale principula Thomas, 1926. See MacPhee (1987a: 9). Holotype. MCZ 45049. Cranium. Adult. Locality. Madagascar: (Toliary), south of Fort Dau- phin (=Tolanaro), Andrahomana caves. 1926. Collector. R. Decary. Condition. Cranium partial (back of cranium miss- ing posterior to parietals). Type Series. Paratype material consists of MCZ 45408, which includes 3 partial crania and 5 man- dibular rami, and MAD-1649, a partial skull in the collections of the Institut de Paléontologie, MNHN. Microgale drouhardi G. Grandidier, 1934 Bull. Mus. Hist. Nat. Paris, 6: 474, 29 November. Holotype. MCZ 45034. Body in alcohol, skull extract- ed. Juvenile female. Locality. Madagascar; (Antsiranana), east coast, Di- ego-Suarez (=Antsiranana). May 1934. Collector. M. E. Drouhard. Original number A. Condition. Alcoholic, skull complete. Type Series. 6 paratypes; MCZ 46007-MCZ 46012, all in alcohol. MCZ 46017, represented by a skull and postcranial skeleton, was collected at the same time and place as the type series but is not men- tioned in the original description. Comments. MCZ 45034 represents an immature animal (MacPhee 1987a: 7), not an adult as claimed in the original description. MacPhee (1987a: 9) synonymized drouhardi with Microgale cowani, an approach followed by Hutterer (1993: 71), but Jen- kins et al. (1997: 6) argued that M. drouhardi is a distinct species. Microgale parvula G. Grandidier, 1934 Bull. Mus. Hist. Nat. Paris, 6: 476, 29 November. Holotype. MCZ 45465. Body in alcohol, skull extract- ed. Juvenile male. Locality. Madagascar: (Antsiranana), east coast, Di- ego-Suarez (=Antsiranana). May 1934. Collector. M. Drouhard. Condition. Alcoholic, skull complete. Type Series. Holotype only. Comments. MCZ 45465 represents an immature animal (MacPhee 1987a: 7), not an adult as claimed in the original description. The holotype of Micro- gale pulla Jenkins, 1988 actually represents an adult specimen of M. parvula (Jenkins et al., 1996: 204). Considered a valid species by Hutterer (1993: 71) and Nowak (1999: 190). Microgale prolixacaudata G. Grandidier, 1937 Bull. Mus. Hist. Nat. Paris, 9: 348, 25 November. =Microgale longicaudata Thomas, 1882. See MacPhee (1987a: 9). Holotype. MCZ 45035. Body in alcohol, skull extract- ed. Juvenile. Locality. Madagascar: (Antsiranana), east coast, Di- ego-Suarez (=Antsiranana). May 1934. Collector. M. Drouhard. Condition. Alcoholic; skull partial (left tympanic bulla missing). Mandible disarticulated. Type Series. 1 paratype; MCZ 46020, in alcohol. Comments. This specimen represents an immature animal (MacPhee 1987a: 8), not an adult as claimed in the original description. Paramicrogale occidentalis G. Grandidier and Petit, 1931 Bull. Soc. Zool. France, 56: 129, 15 June. =Microgale brevicaudata G. Grandidier, 1899. See MacPhee (1987a: 9). Holotype. MCZ 45047. Body in alcohol, skull extract- ed. Juvenile male. Locality. Madagascar: (Antananarivo), northwest of Maintirano, Andriafeuelo. 1930. Collector. M. A. de la Rue. Condition. Alcoholic, skull complete. Type Series. Holotype only, but see comments. Comments. P. occidentalis is the type species of the genus Paramicrogale G. Grandidier and Petit, 1931. This specimen represents an immature ani- mal (MacPhee 1987a: 7), not an adult as claimed in the original description. The original description mentions only a single specimen but describes ex- tensively the skeleton of P. occidentalis. This is puz- zling, as the skeleton has not been extracted from MCZ 45047 (MacPhee 1987a: 7). TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden 103 Genus SETIFER Froriep, 1806 Dasogale fontoynonti G. Grandidier, 1930a Bull. Acad. Malgache, n. ser., 11: 85 (for 1928). = Setifer setosus (Schreber, 1777). See Poduschka and Poduschka (1982: 261). Holotype. MCZ 45016. Skin, skull, and posteranial skeleton. Juvenile. Locality. Madagascar: east coast. 1917. Collector. Received by G. Grandidier from the Académie Malgache in 1917. Condition. Skin represented by a small patch of fur. Skull partial (occipital region missing). Postcra- nial skeleton complete, partially articulated. Type Series. 1 paratype; MCZ 45532, in alcohol. Comments. Walker (1975: 110) commented that “the only specimen known of D. fontoynonti .. . is in the Paris Museum”; actually, the only material attributed to Dasogale is in the MCZ, a fact first noted in publication by Poduschka and Poduschka (1982: 253). Dasogale was often considered to be an extremely rare or recently extinct species until Poduschka and Poduschka (p. 261) and MacPhee (1987b: 135) demonstrated that the holotype is probably a juvenile Setifer setosus. D. fontoynonti is the type species of the genus Dasogale G. Gran- didier, 1930. Order RODENTIA Bowdich, 1821 Family APLODONTIDAE Brandt, 1855 Genus APLODONTIA Richardson, 1829 Aplodontia californica columbiana Taylor, 1916 Univ. California Publ. Zool., 12: 499, 6 May. =Aplodontia rufa rainieri Merriam, 1899. See Dalquest (1948: 369). Holotype. B1899. Skin and skull. Adult male. Locality. (Canada): British Columbia, Hope, Roab’s Ranch. 14 June 1894. Collector. W. C. Colt. Original number 479. Condition. Skin and skull complete. Type Series. 8 paratypes; B1892—B1898, B1900; all represented by skin and skull, 3 females and 5 males. Family SCIURIDAE Fischer de Waldheim, 1817 Genus CALLOSCIURUS Gray, 1867 Callosciurus baluensis medialis G. M. Allen and Coolidge, 1940 Bull. Mus. Comp. Zool., 87: 156, 31 December. Holotype. MCZ 22265. Skin and skull. Adult female. Locality. (Indonesia): Dutch Borneo, (Kalimantan), Mount Tibang (possibly =Bukit Tungun). 1925. Collector. E. Mjoberg. Original number 6. Condition. Skin complete. Skull partial (right pa- rietal, right tympanic bulla, and lachrymal_ bro- ken—bulla present). Mandible disarticulated. Type Series. Holotype only. Comments. Retained as a valid subspecies by Med- way (1977: 90). Callosciurus ferrugineus primus G. M. Allen and Coolidge, 1940 Bull. Mus. Comp. Zool., 87: 157, 31 December. = Callosciurus erythraeus (Pallas, 1778). See Corbet and Hill (1992: 283). Holotype. MCZ 35352. Skin and skull. Adult female. Locality. Siam (=Thailand): Mae Wan River near Doi, Mount Souket (=Saket), 1,500 ft (45S m). 20 February 1937. Collector. J. A. Griswold, Jr., Asiatic Primate Ex- pedition. Original number 5. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; MCZ 35353, skull, adult male. skin and Sciurus castaneoventris haemobaphes G. M. Allen, 1912c Proc. Biol. Soc. Washington, 25: 177, 24 December. =Callosciurus erythraeus haemobaphes (G. M. Allen, 1912). See Hayman and Holt (1940: 359). Holotype. MCZ 13693. Skin and skull. Male. Locality. China: southeastern Yunnan, Chih-ping (=Shiping). 26 February 1911. Collector. Kobayashi Collection. Original number AG. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Only the holotype is mentioned in the original publication, but Allen had four other spec- imens of these squirrels at the time of description (MCZ 13692, 13694-13696; all represented by skin and skull, 3 males, 1 female). 104 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Genus DREMOMYS Heude, 1898 Dremomys pernyi flavior G. M. Allen, 1912c Proc. Biol. Soc. Washington, 25: 178, 24 December. Collector. Harbison and Bargamin. Condition. Skin and skull complete. Type Series. 1 paratype; B6960, skin and _ skull, adult male. Comments. G. s. bangsi was retained as a valid sub- species by Hall (1981: 450). Holotype. MCZ 13691. Skin and skull. Male. Locality. China: southeastern Yunnan, Mongtz (=Mengzi). 1911. Collector. Kobayashi Collection. Original number 6/8. Condition. Skin complete. Skull partial (squamo- sals missing, supraoccipital damaged, right jugal missing, palatine missing, tympanic bulla dam- aged). Type Series. Holotype only. Sciuropterus alpinus lascivus Bangs, 1899} Proc. New England Zool. Club, 1: 69, 31 July. =Glaucomys sabrinus lascivus (Bangs, 1899). See A. H. Howell (1918: 55). Holotype. B9186. Skin and skull. Adult female. Locality. (United States): California, El Dorado County, Tallac. 28 August 1898. Collector. W. W. Price and P. O. Simons. Original Dremomys senex G. M. Allen, 1912b Mem. Mus. Comp. Zool., 40: 229, August. =Dremomys pernyi senex G. M. Allen, 1912. See Hayman and Holt (1940: 382). Holotype. MCZ 7582. Skin and skull. Adult female. number 1722. Condition. Skin and skull complete. Type Series. 2 paratypes, B9187, skin and skull, fe- male; B9188, skin and skull, female. Comments. G. s. lascivus was retained as a valid subspecies by Hall (1981: 451). Locality. China: Hupeh (=Hubei), Ichanghsien, Nantou. 5 February 1909. Collector. W. R. Zappey. Original number 373. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; MCZ 7583, skin and skull, adult male. Sciuropterus sabrinus makkovikensis Sornborger, 1900 Ottawa Nat., 14: 48, 6 June. =Glaucomys sabrinus makkovikensis (Sornborger, 1900). See A. H. Howell (1918: 34). Syntypes. MCZ 10476: Skin and skull. Adult. MCZ Genus FUN/ISCIURUS Trouessart, 1880 Funisciurus pyrrhopus victoriae G. M. Allen and Loveridge, 1942 Bull. Mus. Comp. Zool., 8: 180, February. =Funisciurus pyrrhopus akka De Winton, 1899. See Amtmann (1975: 8). Holotype. MCZ 39199. Skin and skull. Adult male. Locality. Uganda: Toro, Kibale Forest, 4,200 ft (1,281 m). 16 December 1938. Collector. A. Loveridge. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Genus GLAUCOMYS Thomas, 1908 Sciuropterus alpinus bangsi Rhoads, 1897 Proc. Acad. Nat. Sci. Philadelphia, 1897, p. 321 (footnote), July. =Glaucomys sabrinus bangsi (Rhoads, 1897). See A. H. Howell (1918: 38). Holotype. B6959. Skin and skull. Adult male. Locality. (United States): Idaho, Idaho County. March 8, 1897. 10477: Skin and skull. Adult. MCZ 10478: Skin and skull. Adult. Locality. (Canada): Labrador Peninsula, Makkovik. 1899. Collector. W. W. Perrett. Original number 1540. Condition. MCZ 10476: Skin complete. Skull par- tial (base of skull missing). Mandible disarticulated. MCZ 10477: Skin partial (tail broken but present). Skull partial (base of skull missing). Mandible dis- articulated. MCZ 10478: Skin partial (missing left hind foot and tail). Skull partial (occipital chipped). Mandible disarticulated. Type Series. 3 syntypes only. Comments. Sornborger’s original description was based on three specimens, original numbers 1540, 1541, and 1542, now MCZ 10476, 10477, 10478, respectively. G. s. makkovikensis was retained as a valid subspecies by Hall (1981: 453). Sciuropterus silus Bangs, 1896j Proc. Biol. Soc. Washington, 10: 163, 28 December. =Glaucomys volans volans (Linnaeus, 1758). See A. H. Howell (1918: 20). Holotype. B4931. Skin and skull. Adult male. Locality. (United States): West Virginia, Greenbri- TYPE SPECIMENS OF RECENT MAMMALS ® Helgen and McFadden er County, White Sulphur Springs, top of Katis Mtn. 3,200 ft (976 m). 2 September 1895. Collector. T. Surber. Original number 19. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Sciuropterus volans querceti Bangs, 1896} Proc. Biol. Soc. Washington, 10: 166, 28 December. =Glaucomys volans querceti (Bangs, 1896). See A. H. Howell (1918: 26). Holotype. B2451. Skin and skull. Adult female. Locality. (United States): Florida, Citrus County, Citronelle. 17 September 1894. Collector. F. L. Small. Original number 1363. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 paratypes; B2452, skin and skull, adult male; B2453, skin and skull, adult female. Comments. G. v. querceti was retained as a valid subspecies by Hall (1981: 449). Genus HYLOPETES Thomas, 1908 Pteromys (Hylopetes) alboniger orinus G. M. Allen, 1940 The Mammals of China and Mongolia, Natural History of Central Asia, 11: 723, 3 September. =Hylopetes alboniger alboniger (Hodgson, 1836). See Ellerman and Morrison-Scott (1951: 469). Holotype. MCZ 28086. Skin and skull. Adult female. Locality. China: Yunnan, Likiang Range (=Li- jiang), 7,800 ft (2,379 m). December 1931. Collector. J. F. Rock. Condition. Skin complete. Skull partial (most of right tympantic bulla missing, supraoccipital dam- aged). Mandible disarticulated. Type Series. Allen examined 10 specimens in ad- dition to the type, including 4 specimens from the BMNH, 3 from AMNH, and MCZ 28087, skin and skull, an unsexed adult. Pteromys phayrei anchises G. M. Allen and Coolidge, 1940 Bull. Mus. Comp. Zool., 87: 153, 31 December. =Hylopetes phayrei anchises (G. M. Allen and Coolidge, 1940). See Ellerman and Morrison-Scott (1951: 469). Holotype. MCZ 35776. Skin and skull. Adult male. Locality. Siam (=Thailand): (Chiang Mai) Mount 105 Angka (=Doi Inthanon), 4,300 ft (1,312 m). 27 February 1937. Collector. J. A. Griswold, Jr., Asiatic Primate Ex- pedition. Original number 24. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; MCZ 35775, skin and skull, adult female; MCZ 35777, skin and _ skull, adult male; MCZ 35778, skin and skull, subadult male. Genus MARMOTA Blumenbach, 1779 Arctomys flaviventer avarus Bangs, 1899} Proc. New England Zool. Club, 1: 68, 31 July. =Marmota flaviventris avara (Bangs, 1899). See A. H. Howell (1915: 41). Holotype. B7299. Skin and skull. Juvenile female. Locality. (Canada): British Columbia, Okanagan. 17 July 1897. Collector. A. C. Brooks. Original number 969. Condition. Skin and skull complete. Type Series. 2 paratypes; B7298, skin and skull, ju- venile male; B7300, skin and skull, juvenile female. Comments. M. f. avara was retained as a valid sub- species by Hall (1981: 371). Arctomys ignavus Bangs, 1899d Proc. New England Zool. Club, 1: 13, 28 February. =Marmota monax ignava (Bangs, 1899). See A. H. Howell (1915: 29). Holotype. B7971. Skin and skull. Adult male. Locality. (Canada): Labrador Peninsula, Black Bay. 13 July 1898. Collector. E. Doane. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 5 paratypes; B7968—B7970, B7972, B7973 (juvenile); all represented by skin and skull, 4 females and 1 male. Comments. M. m. ignava was retained as a valid subspecies by Hall (1981: 370) and Kwiecinski (1998: 1). Genus MICROSCIURUS J. A. Allen, 1895 Sciurus (Microsciurus) browni Bangs, 1902b Bull. Mus. Comp. Zool., 39:24, April. = Microsciurus alfari browni (Bangs, 1902). See J. A. Allen (1914:151). Holotype. MCZ 10404. Skin and skull. Adult male. Locality. Panama: Chiriqui, Bogaba, 600 ft (183 m). 15 July 1901. 106 Collector. W. W. Brown, Jr. Original number 631. Condition. Skin and skull complete. Type Series. 4 paratypes; MCZ 10405, adult fe- male, 10406, adult female, 10407, adult female, 10408, juvenile male; all represented by skin and skull. Comments. M. a. browni was retained as a valid subspecies by Hall (1981: 439). Genus PARAXERUS Forsyth Major, 1893 Aethosciurus byatti laetus G. M. Allen and Loveridge, 1933 Bull. Mus. Comp. Zool., 75: 96, February. = Paraxerus vexillarius byatti (Kershaw, 1923). See Amtmann (1975: 11). Holotype. MCZ 26198. Skin and skull. Adult male. Locality. Tanganyika Territory (=Tanzania): north end of Lake Nyasa, Ukinga Mountains, Madehani, 7,000 ft (2,135 m). 22 February 1930. Collector. A. Loveridge. Condition. Skin and skull complete. Type Series. 9 paratypes; MCZ 26196, 26197, 26199-26202, 26204-26206; all represented by skin and skull, 4 females and 5 males. Genus SCIUROTAMIAS Miller, 1901 Sciurotamias davidanus thayeri G. M. Allen, 1912b Mem. Mus. Comp. Zool., 40: 231, August. = Sciurotamias davidianus consobrinus (Milne-Edwards, 1868). See Moore and Tate (1965: 308). Holotype. MCZ 8008. Skin and skull. Adult male. Locality. China: western Szechwan (=Sichuan), Washan (=Wushan), 6,000 ft (1,830 m). 17 May 1908. Collector. W. R. Zappey. Original number 163. Condition. Skin complete. Skull partial (basioccip- ital and left maxilla damaged). Mandible disartic- ulated. Type Series. Holotype only. Comments. G. M. Allen (1912b: 231) noted in the original description of thayeri that “unfortunately, the skull of the type was lost.” The skin and skull have been reunited subsequently. Genus SC/URUS Linnaeus, 1758 Sciurus (Guerlinguetus) aestuans chiriquensis Bangs, 1902b Bull. Mus. Comp. Zool., 39: 22, April. = Sciurus granatensis chiriquensis Bangs, 1902. See Hershkovitz (1947: 7). Holotype. MCZ 10044. Skin and skull. Adult male. Locality. Panama: Chiriqui, Divala. 18 November 1900. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Collector. W. W. Brown, Jr. Original number 10. Condition. Skin and skull complete. Type Series. There is a large series of paratypes in the MCZ. Comments. S. g. chiriquensis was retained as a val- id subspecies by Hall (1981: 437) and Nitikman (1985: 1). Sciurus carolinensis extimus Bangs, 1896j Proc. Biol. Soc. Washington, 10: 158, 28 December. Holotype. B4519. Skin and skull. Adult female. Locality. (United States): Florida, Dade County, Miami. 12 March 1895. Collector. L. Brownell. Original number 59. Condition. Skin complete (with bald spots on ven- trum). Skull complete. Type Series. 7 paratypes; B3406, B4517, B4518, (all represented by skin and skull), B4520—B4523 (skins only); 4 females and 3 males. Comments. Retained as a valid subspecies by Hall (1981: 417) and Koprowski (1994: 1). Sciurus carolinensis var. yucatanensis J. A. Allen, 1877 In Coues and J. A. Allen, Monogr. N. Amer. Rodentia, U.S. Geol. Geograph. Survey Terr., Rep. 11: 705, August. = Sciurus yucatanensis J. A. Allen, 1877. See Elliot (1896: 80). Syntype. MCZ 5398. Skin. Adult male. Locality. (Mexico): Yucatan, Merida. March 1865. Collector. A. Schott. Original number 228. For- merly USNM 8502. Condition. Skin complete. Type Series. J. A. Allen referred to “four specimens of this variety before me” in the original descrip- tion (1877: 705). Three specimens are mentioned by number: USNM 8502 (now MCZ 5398), 8503, and 8505. A juvenile referred to in the description (but not by number) corresponds to USNM 8504. Comments. S. yucatanensis was considered a valid species by Hoffman et al. (1993: 443) and Nowak (1999: 1265). The syntype of yucatanensis in the MCZ was received from the USNM in March 1877. Poole and Schantz (1942: 554) state that USNM 8505 is no longer in the USNM. Sciurus ludovicianus vicinus Bangs, 1896) Proc. Biol. Soc. Washington, 10: 150, 28 December. = Sciurus niger vulpinus Gmelin, 1896. See Barkalow (1954: 25). Holotype. B5215. Skin and skull. Adult female. Locality. (United States): West Virginia, Greenbri- TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden er County, White Sulphur Springs. 29 January 1896. Collector. T. Surber. Original number 55. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Sciurus nesaeus G. M. Allen, 1902b Proc. Biol. Soc. Washington, 15: 93, 25 April. = Sciurus granatensis nesaeus G. M. Allen, 1902. See Hershkovitz (1947: 37). Holotype. MCZ 10744. Skin. Adult female. Locality. Venezuela: (Nueva Esparta), Margarita Island, El Valle. 8 July 1901. Collector. A. H. Clark. Original number 619. Condition. Skin complete. The lower incisors are present in the skin. Type Series. Holotype only. Comments. S. g. nesaeus was retained as a valid subspecies by Cabrera (1961: 367) and Nitikman (1985: 1). Sciurus variabilis morulus Bangs, 1900d Proc. New England Zool. Club, 2: 43, 20 September. = Sciurus granatensis morulus Bangs, 1900. See Miller and Kellogg (1955: DT): Holotype. B8420. Skin and skull. Adult female. Locality. Panama: Canal Zone, Loma del Leon. 13 March 1900. Collector. W. W. Brown, Jr. Original number 5. Condition. Skin complete. Skull slightly damaged (left bulla broken). Type Series. 5 paratypes; B8418, B8419, B8421— B8423 (juvenile); all represented by skin and skull, 3 females and 2 males. Comments. S. g. morulus was retained as a valid subspecies by Hall (1981: 437) and Nitikman (1985: 1). Sciurus variabilis saltuensis Bangs, 18980 Proc. Biol. Soc. Washington, 12: 185, 16 November. = Sciurus granatensis saltuensis Bangs, 1898. See Hershkovitz (1947: 15). Holotype. B8144. Skin and skull. Adult female. Locality. Colombia: Magdalena, Santa Marta Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 26 March 1898. Collector. W. W. Brown, Jr. Original number 112. Condition. Skin and skull complete. Type Series. 2 paratypes; B8145, skin and skull, adult male; B8244, skin and skull, adult female. Comments. S. g. saltwensis was retained as a valid 107 subspecies by Cabrera (1961: 368) and Nitikman (1985: 1). Genus SPERMOPHILUS F. Cuvier, 1825 Citellus obscurus siccus G. M. Allen, 1925 Amer. Mus. Novitates, 163: 3, 2 April. = Spermophilus alashanicus Buchner, 1888. See Hoffmann et al. (1993: 444). Holotype. MCZ 19924. Skin and skull. Adult female. Locality. China: Shansi (=Shenxi), 10 miles (16.1 km) west of Taiyuanfu. August 1921. Collector. F. R. Wulsin. Original number 146. Condition. Skin complete. Skull partial (parietals and basiooccipital missing, right tympanic bulla broken but present). Mandible disarticulated. Type Series. Holotype only. Spermophilus armatus Kennicott, 1863 Proc. Acad. Nat. Sci. Philadelphia, 15: 158, June. Syntypes. MCZ 297: Skull. Male. Collected 11 April 1858. Original number 167, formerly USNM 4799. MCZ 4793: Skin and skull. Male. Collected 2 April 1858. Original number 140, formerly USNM 3478 (3373). MCZ 4790: Skin. Female. Collected 26 May 1858. Original number 455, formerly USNM 3470 (3470). MCZ 4794: Skull only (skin spoiled in repreparation, discarded). Male. Collected April 14, 1858. Original number 215, formerly USNM 3474 (3373). Locality. (United States): Utah (now Wyoming), (Uinta County), near Fort Bridger, foothills of the Uinta Mountains, Camp Scott. Collector. C. Drexler. Type Series. See comments below. Comments. Considered a valid species by Hoff- mann et al. (1993: 444) and Nowak (1999: 1254). Kennicott does not designate a type in the original description; thus, “all the specimens from Fort Bridger collected by C. Drexler and in the collec- tions [of the USNM] prior to 1863 are evidently cotypes [=syntypes] of this species” (Lyon and Os- good 1909: 163). The syntypes of armatus in the MCZ were received from the USNM in January 1874. These specimens possess USNM labels that bear numbers in discrepancy with the USNM cat- alogue; the numbers listed above are taken from the catalogue of the USNM, followed in parenthe- ses by the number on the specimen label. A com- plete list of syntypes, most of the remainder of which are in the USNM, can be found in Poole and Schantz (1942: 504-505). The skin of MCZ 297, which formerly bore the number USNM 3472, is not to be found in the collections of either museum. 108 Spermophilus elegans Kennicott, 1863 Proc. Acad. Nat. Sci. Philadelphia, 15: 158, June. Syntypes. MCZ 4791: Skin. Female. Collected 11 April 1858. Original number 168. Formerly USNM 3468. MCZ 4792: Skin. Male. Collected 11 April 1858. Original number 166. Formerly USNM 3473. Locality. (United States): Utah (now Wyoming), (Uinta County), Fort Bridger. Collector. C. Drexler. Condition. Following their receipt, these speci- mens were spoiled in repreparation and discarded. Type Series. See comments below. Comments. These specimens are no longer in ex- istence. Kennicott does not designate a type in the original description; thus, “all the specimens from Fort Bridger collected by C. Drexler and in the collections [of the USNM] prior to 1863 are evi- dently cotypes [=syntypes] of this species” (Lyon and Osgood 1909: 163). These syntypes of elegans were received from the USNM in January 1874. A complete list of syntypes can be found in Lyon and Osgood (1909: 166). G. M. Allen (1931: 251) er- roneously included MCZ 4791 in a list of the type series of Spermophilus armatus. S. elegans is con- sidered a valid species by Hoffmann et al. (1993: 446) and Nowak (1999: 1254). Spermophilus (Ictidomys) tridecemlineatus badius Bangs, 1899c Proc. New England Zool. Club, 1: 1, 8 February. = Spermophilus tridecemlineatus texensis Merriam, 1898. See Hall and Kelson (1959: 347). Holotype. B1682. Skin and skull. Adult male. Locality. (United States): Missouri (Vernon Coun- ty), Stotesbury. 17 April 1894. Collector. T. Surber. Original number 81. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 paratypes; B1683, skin and skull, adult male; B5609, skin and skull, adult male. Comments. A. H. Howell (1938: 110) first synon- ymized badius with texensis under the genus Ci- tellus. Genus SYNTHEOSCIURUS Bangs, 1902 Syntheosciurus brochus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 25, April. Holotype. MCZ 10402. Skin and skull. Adult male. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 7,000 ft (2,135 m). 30 April 1901. Collector. W. W. Brown, Jr. Original number 415. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Condition. Skin complete. Skull partial (left supra- occipital damaged) Type Series. 1 paratype; MCZ 10403, skin and skull, adult female. Comments. Type species of the genus Syntheosciu- rus Bangs, 1902. Considered a valid species by Hoffman et al. (1993: 452) and Nowak (1999: 1268). Following a discussion with a collector em- ployed by W. W. Brown, Jr., and a biological survey of Boquete, Enders (1953b: 509) recommended that the type locality be considered “the Cordillera about 8 miles [12.9 km] north of Boquete and not at Boquete which is on the lower slopes of El Vol- can de Chiriqui.” Genus TAMIAS llliger, 1811 Tamias cooper! Baird, 1855 Proc. Acad. Nat. Sci. Philadelphia, 7: 334, 24 April 24. = Tamias townsendii cooperi Baird, 1855. See Baird (1858: 737). Syntype. MCZ 4754. Skin only. Adult, unsexed. Locality. (United States): Washington (Skagit County), Clickitat (=Klickitat) Pass, Cascade Mountains, 4,500 ft (1,373 m). July 1853. See com- ments. Collector. J. G. Cooper. Formerly USNM 211/ 1182. Condition. Skin complete. Type Series. The other syntype is USNM 212/1183, skin and skull, unsexed adult. Comments. No type is designated in the original description, but Baird subsequently explained that the two specimens listed above were those on which cooperi was based (1858: 301). Cooper (1869: 531) emended the type locality. The syntype of cooperi in the MCZ was received from the USNM in January 1874. Tamias dorsalis Baird, 1855 Proc. Acad. Nat. Sci. Philadelphia, 7: 332, 24 April. =Tamias dorsalis dorsalis Baird, 1855. See Hayman and Holt (1940: 435). Syntype. MCZ 4759. Skin and mandible. Locality. (United States): New Mexico (Grant County), “Fort Webster, Coppermines of the Mim- bres” (near present Georgetown). 32°47’'N, 108°41'W. 1851. See A. H. Howell (1929: 131). Collector. J. H. Clark. Formerly USNM 119/3151. Condition. Skin complete. Mandible partial (pos- terior of right mandibular ramus broken off and missing). Type Series. The other syntype is USNM 120, skull within skin. Comments. No type is designated in the original description, but Baird subsequently listed the two TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden specimens above as those on which dorsalis was based (1858: 300). The syntype of dorsalis in the MCZ was received from the USNM in January 1874. Tamias quadrivittatus neglectus J. A. Allen, 1890 Bull. Amer. Mus. Nat. Hist., 3: 106, June. = Tamias minimus neglectus (J. A. Allen, 1890). See Hayman and Holt (1940: 430). Holotype. MCZ 1575. Skin and cranium. Locality. (Canada): (Ontario), eastern end of Lake Superior (near mouth of Montreal River). 5 July 1848. Collector. L. Agassiz. Condition. Skin complete. Cranium partial (zygo- matic arches, parietals, left tympanic bulla, and mandible missing). Type Series. The description mentions 6 paratypes, including MCZ 1567, skin and skull; and 4 others at USNM. Comments. Following A. H. Howell (1929: 54), Hayman and Holt (1940: 430) considered T. m. neglectus a synonym of T. m. borealis. However, neglectus was retained as a valid subspecies of min- imus by Hall (1981: 346) under the genus Euta- mias. Tamias striatus venustus Bangs, 1896h Proc. Biol. Soc. Washington, 10: 137, 28 December. Holotype. B5478. Skin and skull. Adult male. Locality. (United States): Indian Territory (=Oklahoma) (Adair County), Stilwell. 13 August 1896. Collector. T. Surber. Original number 63. Condition. Skin complete. Skull partial (condyle of left mandibular ramus missing). Type Series. 2 paratypes; B5479, skin and _ skull, adult female; B5605, skin and skull, adult male. Comments. Retained as a valid subspecies by Hall (1981: 340). Genus TAMIASCIURUS Trouessart, 1880 Sciurus hudsonicus gymnicus Bangs, 1899f Proc. New England Zool. Club, 1: 28, 31 March. = Tamiasciurus hudsonicus gymnicus (Bangs, 1899). See Osgood (1938: 438). Holotype. B4914. Skin and skull. Adult female. Locality. (United States): Maine, Piscataquis 109 County, Greenville, near Moosehead Lake. 1 De- cember 1895. Collector. C. H. Goldthwaithe. Original number 2. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. T. h. gymnicus was retained as a valid subspecies by Hall (1981: 442) and Steele (1998: 1). Sciurus hudsonicus loquax Bangs, 1896} Proc. Biol. Soc. Washington, 10: 161, 28 December. = Tamiasciurus hudsonicus loquax (Bangs, 1896). See A. H. Howell (1936: Ih) Holotype. B4270. Skin and skull. Adult male. Locality. (United States): Connecticut, New Lon- don County, Liberty Hill. 24 December 1895. Collector. O. Bangs. Original number 3. Condition. Skin and skull complete. Type Series. There is a large series of paratypes in the MCZ. Comments. T. h. loquax was retained as a valid sub- species by Hall (1981: 443) and Steele (1998: 1). Sciurus hudsonicus orarius Bangs, 1897i Proc. Biol. Soc. Washington, 11: 281, 30 December. = Tamiasciurus douglasii mollipilosus (Audubon and Bachman, 1841). See Hayman and Holt (1940: 347). Holotype. B4978. Skin and skull. Adult female. Locality. (United States): California, Mendocino County, Philo. 9 December 1895. Collector. C. A. Allen. Original number 887. Condition. Skin complete. Skull partial (right tym- panic bulla broken). Type Series. 13 paratypes; B4832, B4979-B4989, B5462; all represented by skin and skull, 9 females and 4 males. Family CASTORIDAE Hemprich, 1820 Genus CASTOR Linnaeus, 1758 Castor caecator Bangs, 1913 Bull. Mus. Comp. Zool., 54: 513, July. = Castor canadensis caecator Bangs, 1913. See G. M. Allen (1942: 62). Holotype. B6979. Skull. Adult male. Locality. (Canada): Newfoundland, near Bay St. George. 1896. Collector. E. Doane. Condition. Skull complete. Mandible disarticulat- ed. 110 Type Series. Holotype only. Comments. C. c. caecator was retained as a valid subspecies by Hall (1981: 602). Family GEOMYIDAE Bonaparte, 1845 Genus GEOMYS Rafinesque, 1817 Geomys colonus Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 178, 15 March. =Geomys pinetis pinetis Rafinesque, 1817. See Williams and Genoways (1980: 444), Holotype. B5001. Skin and skull. Adult male. Locality. (United States): Georgia, Camden Coun- ty, Arnot Plantation, about 4 miles (6.4 km) west of St. Mary’s. 21 March 1896. Collector. O. Bangs. Original number 8. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. Hall (1981: 505) recognized Geomys colonus as a valid species. Geomys cumberlandius Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 180, 15 March. =Geomys pinetis pinetis Rafinesque, 1817. See Williams and Genoways (1980: 444). Holotype. B5016. Skin and skull. Adult male. Locality. (United States): Georgia, Camden Coun- ty, Cumberland Island, Stafford Place. 17 April 1896. Collector. O. Bangs. Original number 1. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. In the original description, Bangs lists B5015 as the type and gives its data. The data given fit B5016, not B5015. B5016 has “Type” written after the entry in Bangs’ catalogue and should be considered the holotype, rather than B5015. Hall (1981: 505) and Laerm (1981: 150) supported the specific status of Geomys cumberlandius. Geomys floridanus austrinus Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 177, 15 March. =Geomys pinetis pinetis Rafinesque, 1817. See Williams and Genoways (1980: 444). Holotype. B6983. Skin and skull. Adult male. Locality. (United States): Florida, Pinellas County, Belleair. 3 August 1897. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Collector. W. S. Dickinson. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype, in the USNM. Geomys tuza goffi Sherman, 1944 Proc. New England Zool. Club, 23: 38, 30 August. =Geomys pinetis goffi Sherman, 1944. See Harper (1952: 37). Holotype. B7222. Skin and skull. Adult male. Locality. (United States): Florida, Brevard County, Eau Gallie. 1S March 1897. Collector. O. Bangs. Original number 1. Condition. Skin and skull complete. Type Series. 12 paratypes; B7212—B7217, B7219— B7221; all represented by skin and skull, 5 females and 7 males. Comments. Williams and Genoways (1980: 444) synonymized gofft with G. pinetis pinetis; however, Hall (1981: 504) maintained G. p. goffi as a valid subspecies. The IUCN designates goffi as extinct. Genus ORTHOGEOMYS Merriam, 1895 Macrogeomys cavator Bangs, 1902b Bull. Mus. Comp. Zool., 39: 42, April. = Orthogeomys cavator cavator (Bangs, 1902). See Russell (1968a: 532). Holotype. MCZ 10381. Skin and skull. Adult male. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 4,800 ft (1,464 m). 9 March 1901. Collector. W. W. Brown, Jr. Original number 212. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a large series of paratypes in the MCZ. Comments. O. cavator was considered a valid spe- cies by Patton (1993: 470) and Nowak (1999: 1314). Macrogeomys pansa Bangs, 1902b Bull. Mus. Comp. Zool., 39: 44, April. = Orthogeomys cavator pansa (Bangs, 1902). See Russell (1968a: 532). Holotype. MCZ 10364. Skin and skull. Adult female. Locality. Panama: Chiriqui, Bogaba, 600 ft (183 m). 6 July 1901. Collector. W. W. Brown, Jr. Original number 567. Condition. Skin complete. Skull partial (right tym- panic bulla missing). Type Series. There are 7 paratypes; MCZ 10362— 10363, 10365-10369, 4 females and 3 males, all represented by skin and skull. MCZ 10362 and 10366 are now in the FMNH, and MCZ 10365 is in the USNM. Comments. O. c. pansa was retained as a valid sub- species by Hall (1981: 513). TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Orthogeomys grandis pluto Lawrence, 1933a Proc. New England Zool. Club, 13: 66, 8 May. Holotype. MCZ 29040. Skin and skull. Adult female. Locality. Honduras: Francisco Morazan, north of Tegucigalpa, Cerro Cantoral. 20 July 1932. Collector. C. F. Underwood. Original number 1100. Condition. Skin and skull complete. Type Series. 2 paratypes; MCZ 29035, skin and skull, juvenile female; MCZ 29039, skin and skull, unsexed juvenile. Comments. Retained as a valid subspecies by Hall (1981: 509). Genus PAPPOGEOMYS Merriam, 1895 Cratogeomys castanops rubellus Nelson and Goldman, 1934a Proc. Biol. Soc. Washington, 47: 147, 13 June. =Pappogeomys castanops rubellus (Nelson and Goldman, 1934). See Russell (1968b: 682). Holotype. MCZ 20507. Skin and skull. Adult male. Locality. Mexico: San Luis Potosi, near San Luis Potosf, Soledad, 6,400 ft (1,952 m). 1 August 1923. Collector. W. W. Brown, Jr. Original number 231. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a large paratype series in the MCZ. Comments. P. c. rubellus was retained as a valid subspecies by Hall (1981: 520). Genus THOMOMYS Wied-Neuwied, 1839 Thomomys umbrinus atrodorsalis Nelson and Goldman, 1934b J. Mammal., 15: 111, 15 May. Holotype. MCZ 20487. Skin and skull. Adult male. Locality. Mexico: San Luis Potosi, Alvarez, 8,000 ft (2,440 m). 7 November 1923. Collector. W. W. Brown, Jr. Original number 338. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a large series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 476). JEILIL Family HETEROMYIDAE Gray, 1868 Genus DIPODOMYS Gray, 1841 Dipodomys californicus pallidulus Bangs, 1899) Proc. New England Zool. Club, 1: 65, 31 July. =Dipodomys californicus californicus Merriam, 1890. See Kelt (1988: 1). Holotype. B9147. Skin and skull. Adult female. Locality. (United States): California, Colusa Coun- ty, Sites. 27 June 1896. Collector. P. O. Simons. Original number 222. Condition. Skin complete. Skull partial (left jugal missing). Type Series. 1 paratype; B9148, skin and skull, adult female. Dipodops ordii palmeri J. A. Allen, 1891 Bull. Amer. Mus. Nat. Hist., 3: 276, 30 June. =Dipodomys ordii palmeri (J. A. Allen, 1891). See Grinnell (1921: 96). Syntypes. MCZ 5886: Skin and skull. Juvenile male. MCZ 5887: Skin and skull. Unsexed juvenile. Locality. Mexico: San Luis Potosi, San Luis Potosi. 1 May 1878. Collector. EF. Palmer. Condition. MCZ 5886: Skin complete. Skull partial (part of parietal, right occipital condyle, and tym- panic bulla missing). MCZ 5887: Skin complete. Skull partial (left squamosal process missing). Man- dible disarticulated. Type Series. 2 syntypes only. Comments. Allen did not specify a holotype in the original description, which he based on the two specimens above. D. 0. palmeri was retained as a valid subspecies by Hall (1981: 569). Genus HETEROMYS Desmarest, 1817 Heteromys repens Bangs, 1902b Bull. Mus. Comp. Zool., 39: 45, April. =Heteromys desmarestianus repens Bangs, 1902. See Goldman (1920: 115). Holotype. MCZ 10356. Skin and skull. Adult female. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 4,000 ft (1,220 m). 8 April 1901. Collector. W. W. Brown, Jr. Original number 264. Condition. Skin complete. Skull partial (left jugal and process of squamosal missing). Type Series. 5 paratypes, MCZ 10359, adult male, 10355, adult male, 10358, adult male, 10361, ju- venile female, 10360, juvenile female. Comments. H. d. repens was retained as a valid subspecies by Hall (1981: 597). 112 Genus LIOMYS Merriam, 1902 Heteromys alleni Coues, 1881 In J. A. Allen, Bull. Mus. Comp. Zool., 8: 187, March. =Liomys irroratus alleni (Coues, 1881). See Goldman (1911: 56). Holotype. MCZ 5889. Skin and skull. Male. Locality. (Mexico): San Luis Potosi, Rio Verde, Ha- cienda Angostura. 26 February 1878. Collector. E. Palmer. Condition. Skin complete, skull within skin. Type Series. Holotype only. Comments. L. i. alleni was retained as a valid sub- species by Hall (1981: 590). Genus PEROGNATHUS Wied-Neuwied, 1839 Perognathus longimembris bangsi Mearns, 1898 Bull. Amer. Mus. Nat. Hist., 10: 300, 31 August. Holotype. B5304. Skin and skull. Adult female. Locality. (United States): California, (Riverside County), Colorado Desert, Palm Springs, 450 ft (137 m). 13 April 1896. Collector. E. C. Thurber. Original number 644. Condition. Skin complete. Skull partial (teeth sep- arate from skull). Mandible disarticulated. Type Series. 2 paratypes; B5302, adult male, skin and skull; B5303, adult female, skin and skull. Comments. Retained as a valid subspecies by Hall (1981: 537). Family DIPODIDAE Fischer de Waldheim, 1817 Genus ZAPUS Coues, 1875 Zapus hudsonius hardyi Batchelder, 1899 Proc. New England Zool. Club, 1: 5, 8 February. : =Zapus hudsonicus acadicus (Dawson, 1856). See Krutzsch (1954: 432). Holotype. MCZ 41681. Skin and skull. Adult female. Locality. (United States): Maine, Hancock County, Mount Desert Island. 24 August 1898. Collector. C. F. Batchelder. Original number 1597. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Zapus hudsonius ladas Bangs, 1899d Proc. New England Zool. Club, 1: 10, 28 February. Holotype. B4169. Skin and skull. Adult female. Locality. (Canada): Labrador, Hamilton Inlet, Ri- goulette. 15 July 1895. Collector. C. H. Goldthwaite. Original number 2. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a large series of paratypes, most of which are in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 843). Zapus orarius Preble, 1899 N. Amer. Fauna, 15: 29, 8 August. =Zapus trinotatus orarius Preble, 1899. See Hooper (1944: 67). Holotype. B250. Skin and skull. Adult male. Locality. (United States): California, Marin Coun- ty, Point Reyes. 14 May 1893. Collector. C. A. Allen. Original number 618. Condition. Skin and skull complete. Type Series. 3 paratypes, not in MCZ; mentioned by locality in original description. Comments. Z. t. orarius was retained as a valid sub- species by Hall (1981: $46) and Gannon (1988: 1). Family MURIDAE Illiger, 1811 Subfamily ARVICOLINAE Gray, 1821 Genus CHIONOMYS Miller, 1908 Hypudaeus nivicola Schinz, 1845 Syst. Verzeichniss Saiugethiere Synopsis Mammalium, 2: 236. =Chionomys nivalis (Martins, 1842). See Musser and Carleton (1993: 507). Syntype. MCZ 1291. Skin. Juvenile female. Locality. Switzerland: St. Gotthard. Collector. Received from L. Agassiz, Condition. Skin complete. Type Series. Schinz wrote that he had examined 16 specimens, “junge und alte.” Comments. G. M. Allen discussed the somewhat uncertain type status of this specimen (1931: 263). The original label bears the following: “Hypodaeus [sic] nivicola juv. 2 sp. nov. du St. Gotthard et du Faulhorn.” Louis Agassiz, a student of Schinz’s, de- posited the specimen in the MCZ. TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden LES Genus CLETHRIONOMYS Tilesius, 1850 Evotomys proteus Bangs, 1897g In Bailey, Proc. Biol. Soc. Washington, 11: 137, 13 May. =Clethrionomys gapperi proteus (Bangs, 1897). See Jackson (1938: 433). Holotype. B4081. Skin and skull. Adult female. Locality. (Canada): Labrador, Hamilton Inlet. 27 August 1895. Collector. C. H. Goldthwaite. Original number 10. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a large series of paratypes in the MCZ. Comments. C. g. proteus was retained as a valid subspecies by Hall (1981: 783). Genus DICROSTONYX Gloger, 1841 Dicrostonyx chionopaes G. M. Allen, 1914b Proc. New England Zool. Club, 5: 62, 9 April. = Dicrostonyx torquatus chionopaes G. M. Allen, 1914. See Ognev (1948: 507). Holotype. MCZ 15263. Skin and skull. Adult male. Locality. U.S.S.R. (=Russian Federation): eastern Siberia, Nijni Kolymsk (Nizhnekolymsk), near mouth of Kolyma River. 15 October 1911. Collector. J. Koren. Original number 257. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Dicrostonyx exsul G. M. Allen, 1919a Bull. Mus. Comp. Zool., 62: 532, February. Holotype. MCZ 11885. Skin and skull. Adult male. Locality. (United States): (Alaska), St. Lawrence Is- land. 24 June 1913. Collector. J. Dixon. Original number 3267. Condition. Skin complete. Skull partial (basioccip- ital and right tympanic bulla missing). Mandible disarticulated. Type Series. 3 paratypes; MCZ 11883, skin and skull, female; MCZ 11884, skin and skull, female: USNM 232007, skin and skull, female. Comments. Considered a valid species by Musser and Carleton (1993: 510) but included in D. groen- landicus by Nowak (1999: 1479). Genus EOTHENOMYS Miller, 1896 Craseomys aquilus G. M. Allen, 1912b Mem. Mus. Comp. Zool., 40: 216, August. =Eothenomys eva eva (Thomas, 1911). See G. M. Allen (1940: 837). Holotype. MCZ 7190. Skin and skull. Adult male. Locality. China: Hupeh (=Hubei), Showlungtan. 17 May 1907. Collector. W. R. Zappey. Original number 10. Condition. Skin complete. Skull partial (hole in left parietal, occiput missing). Type Series. 5 paratypes; MCZ 7189, 7191-7194, 7196; all represented by skin and skull, all female. Microtus (Eothenomys) aurora G. M. Allen, 1912b Mem. Mus. Comp. Zool., 40: 211, August. =Eothenomys melanogaster aurora (G. M. Allen, 1912). See Hinton (1923: 149). Holotype. MCZ 7788. Skin and skull. Male. Locality. China: Hupeh (=Hubei), Changyangh- sieh. 2 February 1909. Collector. W. R. Zappey. Original number 372. Condition. Skin and skull complete. Type Series. 3 paratypes; MCZ 7185, skin and skull, female; MCZ 7186, skin and skull, male; MCZ 7188, skin and skull, male. Comments. E. m. aurora was retained as a valid subspecies by Zhang et al. (1997: 229). Microtus (Eothenomys) mucronatus G. M. Allen, 1912b Mem. Mus. Comp. Zool., 40: 214, August. =Eothenomys melanogaster melanogaster (Milne-Edwards, 1871). See G. M. Allen (1940: 806). Holotype. MCZ 7789. Skin and skull. Adult female. Locality. China: western Szechwan (=Sichuan), Tachiao, 12,000 ft (3,660 m). 11 August 1908. Collector. W. R. Zappey. Original number 269. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; MCZ 7790, skin and skull, adult female (exchanged to the BMNH); MCZ 7791, skin and skull, adult female; MCZ 7803, skin and skull, juvenile female. 114 Genus LEMMUS Link, 1795 Lemmus paulus G. M. Allen, 1914b Proc. New England Zool. Club, 5: 60, 9 April. =Lemmus sibiricus chrysogaster J. A. Allen, 1903. See Ellerman and Morrison- Scott (1951: 656). Holotype. MCZ 15268. Skin and skull. Adult male. Locality. (Russian Federation): northeastern Sibe- ria, Kalaschowo, near mouth of Kolyma River. 22 June 1912. Collector. J. Koren. Original number 152. Condition. Skin complete. Skull partial (left man- dibular ramus missing). Type Series. 1 paratype; MCZ 15267, skin, female. Genus MICROTUS Schrank, 1798 Arvicola breweri Baird, 1858 Mammals, in Repts. Explor. Surveys Railr. to Pacific, 8(1): 525, 14 July. =WMicrotus breweri (Baird, 1858). See Miller (1896: 83). Syntype. MCZ 4365. Body in alcohol. Formerly USNM 2833. Locality. (United States): Massachusetts, Nantuck- et County, off Nantucket Island, Muskeeget (=Mu- skeget) Island. July 1856. Collector. T. M. Brewer. Condition. Alcoholic. Type Series. There are 5 other syntypes (see Poole and Schantz, 1942: 271-272), all of which were at one time in the USNM. Comments. Poole and Schantz (1942: 271) noted that USNM 2829, a syntype of breweri, could not be found. Arvicola riparia var. longipilis Baird, 1858 Mammals, in Repts. Explor. Surveys Railr. to Pacific, 8(1): 524, 14 July. =Microtus pennsylvanicus pennsylvanicus (Ord, 1815). See Bailey (1900: 16). Syntype. MCZ 5292. Skin and skull. Locality. (United States): Illinois, (Cook County), West Northfield. Spring 1855. Collector. R. Kennicott. Formerly USNM 745. Condition. Skin complete. Skull partial (part of pa- rietal, both tympanic bullae and palatine missing). Mandible disarticulated. Type Series. See comments. Comments. G. M. Allen (1931: 260), mentioned that “although no individuals are mentioned [in Baird’s description], ... [this] specimen was un- doubtedly among those examined by Baird in pre- paring his diagnosis and, therefore, is a cotype Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 [=syntype].” Baird’s specimens were collected by Kennicott at West Northfield, [linois, and by Hoy at Racine, Wisconsin, and were apparently depos- ited in the USNM. Lyon and Osgood (1909) and Poole and Schantz (1942) make no mention of oth- er syntypes. Arvicola rufidorsum Baird, 1858 Repts. Explor. Surveys Railr. to Pacific, 8(1): 526, 14 July. = Microtus pennsylvanicus pennsylvanicus (Ord, 1815). See Bailey (1900: 16). Holotype. MCZ 54372. Skin and skull. Locality. (United States): Massachusetts, (Duke’s County), Martha’s Vineyard, Holmes Hole. Collector. D. J. Wyman. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. This specimen has formerly been BSNH 1949 and USNM 901. Arvicola terraenovae Bangs, 1894a Proc. Biol. Soc. Washington, 9: 129, 27 July. =Microtus pennsylvanicus terraenovae (Bangs, 1894). See Davis (1936: 290). Holotype. B1104. Skin and skull. Adult male. Locality. (Canada): Newfoundland, Codroy. 27 No- vember 1893. Collector. E. Doane. Original number 4. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. M. p. terraenovae was retained as a val- id subspecies by Hall (1981: 796). Microtus chrotorrhinus ravus Bangs, 1898p Proc. Biol. Soc. Washington, 12: 188, 16 November. Holotype. B7951. Skin and skull. Adult male. Locality. (Canada): Labrador, Strait of Belle Isle, Black Bay. 15 July 1898. Collector. E. Doane. Original number 4. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Paratype material consists of Bangs’ series from the type locality; B7952—7967, all rep- resented by skin and skull, 10 females, 3 males, and 3 unsexed individuals. B7953 and B7955 were ex- changed to MVZ in 1937; B7964 was exchanged to USNM in 1922. Comments. Retained as a valid subspecies by Hall (1981: 811) and Kirkland and Jannett (1982: 1). TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Microtus enixus Bangs, 1896g Amer. Nat., 30: 1051, 1 December. =Microtus pennsylvanicus enixus Bangs, 1896. See Davis (1936: 290). Holotype. B3973. Skin and skull. Adult female. Locality. (Canada): Labrador, Hamilton Inlet. 15 July 1895. Collector. C. H. Goldthwaite. Original number 4. Condition. Skin and skull complete. Type Series. There is a large series of paratypes in the MCZ. Comments. M. p. enixus was retained as a valid subspecies by Hall (1981: 793). Microtus fontigenus Bangs, 1896d Proc. Biol. Soc. Washington, 10: 48, 9 March. =Microtus pennsylvanicus fontigenus Bangs, 1896. See Miller (1897: 14). Holotype. B3837. Skin and skull. Adult female. Locality. (Canada): Quebec, Lake Edward. 28 Sep- tember 1895. Collector. E. A. and O. Bangs. Original number 9. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 7 paratypes; B3838—B3544; all repre- sented by skin and skull, 2 females and 5 males. Comments. M. p. fontigenus was retained as a valid subspecies by Hall (1981: 793). Microtus koreni G. M. Allen, 1914b Proc. New England Zool. Club, 5: 64, 9 April. =Microtus oeconomus koreni G. M. Allen, 1914. See Ellerman and Morrison-Scott (1951: 706). Holotype. MCZ 15213. Skin and skull. Adult female. Locality. (Russian Federation): northeastern Sibe- ria, Nijni Kolymsk (=Nizhnekolymsk), near mouth of Kolyma River. 1 November 1911. Collector. J. Koren. Original number 132. Condition. Skin complete. Skull partial (left tym- panic bulla broken, right mandibular ramus miss- ing). Type Series. There is a large series of paratypes in the MCZ. Microtus pennsilvanicus [sic] shattucki Howe, 1901 iRnoe Portland Soc) Nat. Iaist.. 2 2012 31 December. =Microtus pennsylvanicus shattucki Howe, 1901. See Wyman (1922: 162). Holotype. MCZ 10011. Skin and skull. Adult female. Locality. (United States): Maine, Penobscot Bay, 115 near Long Island, Tumble Down Dick Island. 10 July 1900. Collector. R. H. Howe and G. C. Shattuck. Original number 31. Condition. Skin and skull complete. Type Series. There is a small series of paratypes in the MCZ. Comments. Wyman (1922: 166) considered M. p. shattucki a synonym of M. p. pennsylvanicus, but Hall (1981: 796) retained it as a separate subspe- cies. Microtus pennsylvanicus acadicus Bangs, 1897c Amer. Nat., 31: 239, 1 March. Holotype. B2155. Skin and skull. Adult female. Locality. Canada: Nova Scotia, Digby. 22 July 1894. Collector. O. Bangs. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 792). Microtus provectus Bangs, 1908 Proc. New England Zool. Club, 4: 20, 6 March. =Microtus pennsylvanicus provectus Bangs, 1908. See Chamberlain (1954: 589). Holotype. B9794. Skin and skull. Adult female. Locality. (United States): Rhode Island, Block Is- land. 5 August 1899. Collector. O. Bangs. Original number 11. Condition. Skin complete. Skull partial (tympanic bullae broken). Mandible disarticulated and angu- lar process chipped. Type Series. There is a series of paratypes in the MCZ. Comments. M. p. provectus was retained as a valid subspecies by Hall (1981: 796). Genus MYOPUS Miller, 1910 Myopus thayeri G. M. Allen, 1914b Proc. New England Zool. Club, 5: 58, 9 April. =Myopus schisticolor thayeri G. M. Allen, 1914. See Ognev (1948: 526). Holotype. MCZ 15264. Skin and skull. Adult male. Locality. (Russian Federation): northeastern Sibe- ria, Yakutsk, Nijni Kolymsk (=Nizhnekolymsk), near mouth of Kolyma River. 28 March 1912. Collector. J. Koren. Original number 264. Condition. Skin and skull complete. Mandible dis- articulated. 116 Comments. 1 paratype; MCZ 15265, skin and skull, male. Genus ONDATRA Link, 1795 Fiber obscurus Bangs, 1894b Proc. Biol. Soc. Washington, 9: 133, 15 September. = Ondatra zibethicus obscurus (Bangs, 1894). See Cameron (1959: 85). Holotype. B1155. Skin and skull. Adult female. Locality. (Canada): Newfoundland, Codroy. 14 May 1894. Collector. E. Doane. Original number 3. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a small series of paratypes in the MCZ. Comments. O. z. obscurus was retained as a valid subspecies by Hall (1981: 827). Fiber zibethicus aquilonius Bangs, 1899d Proc. New England Zool. Club, 1: 11, 28 February. = Ondatra zibethicus aquilonius (Bangs, 1899). See Miller (1912:231). Holotype. B3957. Skin and skull. Adult male. Locality. (Canada): Labrador, Hamilton Inlet, Ri- goulette. 15 August 1895. Collector. C. H. Goldthwaite. Original number 11. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; B7974, skin and skull, adult female; B7975, skin and_ skull, juvenile; B8704, skull, juvenile. Comments. O. z. aquilonius was retained as a valid subspecies by Hall (1981: 825). Fiber zibethicus rivalicius Bangs, 1895b Proce, Bostom Soc. Nat Hist) 26:54" 31 July. = Ondatra zibethicus rivalicius (Bangs, 1895). See Davis and Lowery (1940: DAD) Holotype. B2719. Skin and skull. Adult male. Locality. (Canada): Louisiana, Plaquemines Parish, Burbridge. 31 January 1895. Collector. F. L. Small. Original number 1556/165. Condition. Skin and skull complete. Type Series. 4 paratypes; B2720, skin and skull, adult female; B2721, skin and skull, adult male: B2883, skin and skull, male; B2884, skin and skull, male. Comments. O. z. rivalicius was retained as a valid subspecies by Hall (1981: 827). Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Genus PHENACOMYS Merriam, 1889 Phenacomys celatus crassus Bangs, 1900c Proc. New England Zool. Club, 2: 39, 20 September. =Phenacomys ungava crassus Bangs, 1900. See A. B. Howell (1926: 27). Holotype. B3959. Skin and skull. Adult male. Locality. (Canada): Labrador, Hamilton Inlet, Ri- goulette. 15 August 1895. Collector. C. H. Goldthwaite. Original number 1. Condition. Skin and skull complete. Type Series. There is a small series of paratypes, most of which are no longer in the MCZ. Comments. In the original description, “3946” is a misprint for 3964. Genus SYNAPTOMYS Baird, 1858 Synaptomys fatuus Bangs, 1896d Proc. Biol. Soc. Washington, 10: 47, 9 March. = Synaptomys cooperi cooperi Baird, 1858. See Wetzel (1955:8). Holotype. B3857. Skin and skull. Adult female. Locality. (Canada): Quebec, Lake Edward. 28 Sep- tember 1895. Collector. E. A. and O. Bangs. Original number 3. Condition. Skin and skull complete. Type Series. 8 paratypes; B3854—B3856, B3858— B3862; all represented by skin and skull, 3 females and 5 males. Synaptomys (Mictomys) innuitus medioximus Bangs, 1900c Proc. New England Zool. Club, 2: 40, 20 September. = Synaptomys borealis medioximus Bangs, 1900. See A. B. Howell (1927: 9). Holotype. B8852. Skin and skull. Adult male. Locality. (Canada): Labrador, Strait of Belle Isle, Lance (=Lanse) au Loup. 15 April 1899. Collector. E. Doane. Original number 7. Condition. Skin complete. Skull slightly damaged (hole in left parietal). Type Series. 1 paratype; B3972, skin and skull, adult male. Comments. S. b. medioximus was retained as a val- id subspecies by Hall (1981: 834). TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Subfamily CRICETINAE Fischer de Waldheim, 1817 Genus CANSUMYS G. M. Allen, 1928 Cansumys canus G. M. Allen, 1928 J. Mammal., 9: 245, 9 August. Holotype. MCZ 23779. Skin and skull. Adult female. Locality. China: southern Kansu (=Xinjiang), Choni. 9 December 1925. Collector. R. B. Ekvall. Original number 79. Condition. Skin complete. Skull partial (left jugal and occiput missing). Type Series. 1 paratype; MCZ 23780, skin and skull, juvenile male. Comments. Type species of Cansumys G. M. Allen, 1928. Considered a valid species by Musser and Carleton (1993: 537) and Nowak (1999: 1423). Known only by the type series and one additional specimen, FMNH 36067. Subfamily CRICETOMYINAE Roberts, 1951 Genus SACCOSTOMUS Peters, 1846 Saccostomus cricetulus G. M. Allen and Lawrence, 1936 Bull. Mus. Comp. Zool., 79: 100, January. =Saccostomus mearnsi Heller, 1910. See Hubert (1978: 51). Holotype. MCZ 31475. Skin and skull. Subadult male. Locality. Uganda: Sabei District, due north of Mount Elgon, south bank of Greek River (=Kelim River), 3,000 ft (915 m). 5 December 1933. Collector. A. Loveridge. Condition. Skin complete. Skull partial (right tym- panic bulla missing). Mandible disarticulated. Type Series. 1 paratype; MCZ 31474, skin and skull, adult female. Subfamily DENDROMURINAE G. M. Allen, 1939 Genus STEATOMYS Peters, 1846 Steatomys pratensis nyasae Lawrence and Loveridge, 1953 Bull. Mus. Comp. Zool., 110: 39, June. Holotype. MCZ 44213. Skin and skull. Adult male. Locality. Nyasaland (=Malawi): foot of Mulanje Mountain, Likabula River, 2,100 ft (641 m). 29 July 1948. Collector. A. Loveridge. Condition. Skin and skull complete. Type Series. 14 paratypes; MCZ 44214-44216, 4421844228; all represented by skin and skull, 9 males and 5 females. LI 7 Comments. Retained as a valid subspecies by An- sell (1978: 77). Subfamily GERBILLINAE Gray, 1825 Genus TATERA Lataste, 1882 Tatera flavipes G. M. Allen, 1914d Bull. Mus. Comp. Zool., 58: 331, July. =Tatera valida kempi Wroughton, 1906. See Bates (1988: 277). Holotype. MCZ 14491. Skin and skull. Adult female. Locality. Sudan: Blue Nile, north of (Er) Roseires, Aradeiba. 22 January 1913. Collector. G. M. Allen and J. C. Phillips. Original number 69. . Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Tatera soror G. M. Allen, 1914d Bull. Mus. Comp. Zool., 58: 333, July. = Tatera valida kempi Wroughton, 1906. See Bates (1988: 277). Holotype. MCZ 14492. Skin and skull. Adult female. Locality. Sudan: Blue Nile, Fazogli. 16 January 1913. Collector. G. M. Allen and J. C. Phillips. Original number 53. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; MCZ 14493, skin and skull, juvenile male. Genus TATERILLUS Thomas, 1910 Taterillus melanops G. M. Allen, 1912a Bull. Mus. Comp. Zool., 54: 446, April. = Taterillus harringtoni (Thomas, 1906). See Musser and Carleton (1993: 563). Holotype. MCZ 8132. Skin and skull. Male. Locality. British East Africa (=Kenya): arid plains by the Meru River. 11 August 1909. Collector. G. M. Allen. Original number 108. Condition. Skin and skull complete. Type Series. Holotype only. Subfamily MURINAE Illiger, 1811 Genus APODEMUS Kaup, 1829 Apodemus mystacinus euxinus G. M. Allen, 1915b Bull. Mus. Comp. Zool., 59: 11, February. =Apodemus mystacinus mystacinus (Danford and Alston, 1877). See Corbet (1978: 133). Holotype. MCZ 14887. Skin and skull. Male. Locality. Asia Minor (=Turkey): (Trabzon), near 118 Trebizond (=Trabzon), Scalita, 1,000 m. 25 No- vember 1905. Collector. A. Robert. Original number 2189. Condition. Skin complete. Skull partial (right and left jugal missing, right tympanic bulla separated but present, right and left squamosal missing). Mandible disarticulated. Type Series. Holotype only. Genus DASYMYS Peters, 1875 Dasymys incomtus alleni Lawrence and Loveridge, 1953 Bull. Mus. Comp. Zool., 110: 53, June. Holotype. MCZ 26322. Skin and skull. Adult male. Locality. Tanganyika Territory (=Tanzania): (Mbeya), Iolo, near Rungwe Mountains, 4,600 ft (1,403 m). 31 March 1930. Collector. A. Loveridge. Condition. Skin and skull complete. Mandible dis- articulated and right coronoid process chipped. Type Series. 11 paratypes, all represented by skin and skull, in the MCZ. Comments. Retained as a valid subspecies by An- sell (1978: 83). Genus GRAMMOMYS Thomas, 1915 Thamnomys ochraceus G. M. Allen, 1912a Bull. Mus. Comp. Zool., 54: 442, April 1912. =Grammomys macmillani (Wroughton, 1907). See Musser and Carleton (1993: 594). Holotype. MCZ 8126. Skin and skull. Adult male. Locality. British East Africa (=Kenya): Meru Riv- er, near junction with northern Guaso Nyiro (=Ewaso Ngiro). 8 August 1909. Collector. G. M. Allen. Original number 103. Condition. Skin and skull complete. Type Series. Holotype only. Genus HYLOMYSCUS Thomas, 1926 Hylomyscus alleni simus G. M. Allen and Coolidge, 1930 In Strong, The African Republic of Liberia and the Belgian Congo, 2: 599, October. =Hylomyscus alleni (Waterhouse, 1838). See Heim de Balsac and Aellen (1965: TID" Holotype. MCZ 24028. Skin and skull. Adult male. Locality. Liberia: Merikay. 13 September 1926. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Collector. G. M. Allen and H. J. Coolidge, Jr. Orig- inal number 97. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a series of paratypes in the MCZ. Genus MELOMYS Thomas, 1922 Melomys levipes stevensi Rummler, 1935 Z. Saugetierkunde, 10: 109, 31 December. =Melomys mollis Thomas, 1913. See Flannery (1990: 226). Holotype. MCZ 29890. Skin and skull. Adult male. Locality. Papua New Guinea: Morobe, Mount Mis- im (=Missim), 6,700 ft (2,044 m). 17 April 1933. Collector, H. Stevens. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 11 paratypes; MCZ 29889, 29892— 29899, 29901; all represented by skin and skull, 4 females and 7 males. Melomys moncktoni alleni Rummler, 1935 Z. Séugetierkunde, 10: 112, 31 December. =Melomys rubex alleni Rummler, 1935. See Tate (1951: 299). Holotype. MCZ 29902. Skin and skull. Adult female. Locality. Papua New Guinea: Morobe, Mount Mis- im (=Missim), 6,700 ft (2,044 m). 17 April 1933. Collector, H. Stevens. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; MCZ 29900, skin and skull, adult male. Comments. Flannery (1995a: 308-309) did not list subspecies of M. rubex, noting, “It is a very variable species, and a taxonomic revision is needed in or- der to clarify the taxonomic status of many popu- lations.” Genus MUS Linnaeus, 1758 Leggada bufo ablutus G. M. Allen and Loveridge, 1942 Bull’yMuss Comp. ZooleiSO 199! February. =Mus bufo (Thomas, 1906). See Musser and Carleton (1993: 622). Holotype. MCZ 40745. Skin and skull. Adult male. Locality. Belgian Congo (=Democratic Republic | of Congo): Kivu, Lake Kivu, Idjwi Island, Upper — Mulinga, 6,500 ft (1,983 m). 24 February 1939. Collector. A. Loveridge. TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 paratypes; MCZ 40746, skull (no longer in MCZ); MCZ 40747, skull, female. skin and skin and Leggada gerbillus G. M. Allen and Loveridge, 1933 Bull. Mus. Comp. Zool., February. =Mus tenellus (Thomas, 1903). See Musser and Carleton (1993: 629). Holotype. MCZ 26586. Skin and skull. Adult male. Locality. pei Territory (=Tanzania): Do- doma, Ugogo, 3,700 ft (1, 129 m). 23 December 1929. Collector. A. Loveridge. Condition. Skin complete. Skull partial (right jugal missing). Mandible disarticulated and angular pro- cess of right ramus chipped. Type Series. Holotype only. Tee Aes. Mus bactrianus tantillus G. M. Allen, 1927 Amer. Mus. Novitates 270: 9, 31 May. =Mus musculus tantillus G. M. Allen, 1927. See Marshall (1977: 214). Holotype. MCZ 23476. Skin and skull. Adult female. Locality. China: Sze-chuan (=Sichuan), Wanhsien (=Wanxian). 14 November 1921. Collector. W. W. Granger. Condition. Skin complete. Skull partial (6 frag- mone Mandible disarticulated. Type Series. There is a series of paratypes in the AMNH, anal 1 paratype in the MCZ; MCZ 23475 (formerly AMNH 56403), skin and skull, male. Comments. This specimen was formerly AMNH 56416. The original description lists AMNH 56413 as the holotype, but this specimen is a juvenile, not an adult female, as Allen stated. The collection date and measurements of the holotype in the original description fit MCZ 23476 rather than any other specimens in the type series at the AMNH, which are all immatures (Lawrence 1993: 136). Genus MYOMYS Thomas, 1915 Praomys fumatus oweni Setzer, 1956 Proc. U.S. Nat. Mus., 106: 525, 28 November. =Myomys fumatus (Peters, 1878). See Musser and Carleton (1993: 631). Holotype. MCZ 45883. Skin and skull. Adult male. Locality. Anglo-Egyptian Sudan (=Sudan): Equa- toria Province, Torit District, Murukurun, 50 miles (80.5 km) east of Torit, 2,000 ft (610 m). 9 May 1950. 119 Collector. J. S. Owen. Original number 1030. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Genus NIVIVENTER Marshall, 1976 Epimys zappeyi G. M. Allen, 1912b Mem. Mus. Comp. Zool., 40: 225, August. =WNiviventer confucianus (Milne-Edwards, 1871). See Musser and Carleton (1993: 683): Holotype. MCZ 7607. Skin and skull. Adult male. Locality. China: western Szechwan (=Sichuan), Washan Mountains (=Wu Shan), 9,000 ft (2,745 m). 26 October 1908S. Collector. W. R. Zappey. Original number 305. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Genus OENOMYS Thomas, 1904 Oenomys hypoxanthus talangae Setzer, 1956 Proc. U.S. Nat. Mus., November. 106: 505, 28 Holotype. MCZ 45315. Skin and skull. Adult female. Locality. Anglo- Egyptian Sudan (=Sudan): Equa- toria Province, Imatong Mountains, Talanga For- est, 3,000 ft (915 m). 10 July 1950. Collector. J. S. Owen. Original number 1338. Condition. Skin and skull complete. Type Series. | paratype; MCZ 45284, skull, male. Genus PRAOMYS Thomas, 1915 Praomys tullbergi melanotus G. M. Allen and Loveridge, 1933 Bull. Mus. Comp. Zool., February. =Praomys delectorum (Thomas, 1910). See Musser and Carleton (1993: 642). Holotype. MCZ 26287. Skin and skull. Adult male. Locality. Tanganyika Territory (=Tanzania): north- west end of Lake Nyasa, Poroto Mountains, Nyam- wanga, 6,400 ft (1 952 m). 21 March 1930. Collector. A. Loveridge. Condition. Skin complete. Skull partial (left tym- panic bulla missing). Mandible disarticulated. Type Series. 23 paratypes; MCZ 26259, 26285, 26256, 26288—26293, 26295-26297, 26387-26394, 26411, 26497, 26498: all represented by skin and skull. skin and Tex Alley 120 Comments. Van der Straeten and Dieterlen (1987: 9) and Van der Straeten and Dudu (1990: 81) treat- ed melanotus as a species within the Praomys de- lectorum species complex. Genus PSEUDOHYDROMYS Rummler, 1934 Pseudohydromys murinus Rummler, 1934 Z. Siiugetierkunde, 9: 48, 30 December. Holotype. MCZ 29904. Skin and skull. Adult male. Locality. Papua New Guinea, Morobe, Mount Mis- im (=Missim), 7,000 ft (2,135 m). 8 March 1933. Collector. H. Stevens. Condition. Skin complete, with small bald spots. Skull complete. Mandible disarticulated. Type Series. Holotype only. Comments. Type species of Pseudohydromys Rii- mmier, 1934, Considered a valid species by Flan- nery (1995a: 255), Musser and Carleton (1993: 644), and Nowak (1999: 1612). Genus STENOMYS Thomas, 1910 Stenomys niobe stevensi Rummler, 1935 Z. Séugetierkunde, 10: 117, 31 December. =Stenomys niobe niobe (Thomas, 1906). See Flannery (1995a: 339). Holotype. MCZ 29915. Skin and skull. Adult male. Locality. Papua New Guinea: Morobe, Mount Mis- im (=Missim), 7,000 ft (2,135 m). 16 January 1933. Collector. H. Stevens. Original number 8. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. Taylor et al. (1982: 193) first synony- mized stevensi with niobe, under the genus Rattus. Stenomys verecundus mollis Rummler, 1935 Z. Siugetierkunde, 10: 116, 31 December. Holotype. MCZ 29905. Skin and skull. Adult female. Locality. Papua New Guinea: Morobe, Mount Mis- im (=Missim), 5,850 ft (1,784 m). 14 April 1933. Collector. H. Stevens. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. Retained as a valid subspecies by Flan- nery (1995a: 346). Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Genus THAMNOMYS Thomas, 1907 Thamnomys venustus kivuensis G. M. Allen and Loveridge, 1942 Bull. Mus. Comp. Zool., 89: 192, February. Holotype. MCZ 39151. Skin and skull. Adult female. Locality. Belgian Congo (=Democratic Republic of Congo): (Sud-Kivu), Lake Kivu, Idjwi Island, Upper Mulinga, 6,500 ft (1,983 m). 2 March 1939. Collector. A. Loveridge. Condition. Skin complete. Skull partial (separated between nasal and frontal). Right and left angular processes of mandible chipped. Type Series. Holotype only. Subfamily NESOMYINAE Forsyth Major, 1897 Genus NESOMYS Peters, 1870 Nesomys lambertoni G. Grandidier, 1930c Bull. Acad. Malgache, n. ser., 11: 95 (for 1928). Holotype. MCZ 45941. Skull and postcranial skele- ton. Adult male. Locality. Malagasy Republic (=Madagascar): west coast, vicinity of Maintirano. Collector, No collection data available. Received by G. Grandidier from Académie Malgache, 1928. Condition. Skull partial (left zygomatic arch miss- ing, right jugal missing, several small holes). Post- cranial skeleton complete (small foot bones pre- sumably preserved in skin). Type Series. 2 paratypes, MCZ 45933 and 45934, poorly prepared skins (with partial skulls within), both male. Comments. The skin belonging to the holotype is in the MNHN, Paris (M. Carleton, personal com- munication). Considered a valid species by Nowak — (1999: 1434). Subfamily OTOMYINAE Thomas, 1897 Genus OTOMYS F. Cuvier, 1824 Otomys anchietae lacustris G. M. Allen and Loveridge, 1933 Bull. Mus. Comp. Zool., 75: 120, February. =Otomys lacustris G. M. Allen and Loveridge, 1933. See Dieterlen and Van der Straeten (1992: 385). Holotype. MCZ 26358. Skin and skull. Adult female. | Locality. Tanganyika Territory (=Tanzania): north — end of Lake Nyasa, Ukinga Mountains, Madehani, — 7,000 ft (2,135 m). 21 February 1930. TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Collector. A. Loveridge. Condition. Skin complete. Skull partial (left tym- panic bulla damaged). Mandible disarticulated and left coronoid process chipped. Type Series. 17 paratypes; MCZ 26326, 26344— 26351, 26353-26357, 26359, 26654, 26658; all rep- resented by skin and skull. Comments. Included in O. anchietae by Musser and Carleton (1993: 680) but treated as a full spe- cies by Nowak (1999: 1439). Otomys barbouri Lawrence and Loveridge, 1953 Bull. Mus. Comp. Zool., 110: 63, June. Holotype. MCZ 31369. Skin and skull. Adult male. Locality. Uganda: Mount Elgon, Kaburomi, 1°14’N, 34°31’'E, 10,500 ft (3,203 m). 28 December 1933. Collector. A. Loveridge. Condition. Skin and skull complete. Type Series. 9 paratypes; MCZ 31371-31372, 31376, 31421-31425, 31438 (4 males, 4 females, and 1 unsexed individual). Comments. Considered a valid species by Dieter- len and Van der Straeten (1992: 385) and Nowak (1999: 1439) but included in O. anchietae by Mus- ser and Carleton (1993: 680). Otomys uzungwensis Lawrence and Loveridge, 1953 Bull. Mus. Comp. Zool., 110: 61, June. =Otomys typus uzungwensis Lawrence and Loveridge, 1953. See Hanney (1965: 626). Holotype. MCZ 26645. Skin and skull. Adult female. Locality. Tanganyika Territory (=Tanzania): Iringa District, Uzungwa Mountains, Dadaga, 6,000 ft (1,830 m). 31 December 1929. Collector. A. Loveridge. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. O. t. uzungwensis was retained as a val- id subspecies by Ansell (1978: 78). Subfamily SIGMODONTINAE Wagner, 1843 Genus ISTHMOMYS Hooper and Musser, 1964 Megadontomys flavidus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 27, April. =Isthmomys flavidus (Bangs, 1902). See Hooper and Musser (1964: 12). Holotype. MCZ 10331. Skin and skull. Adult male. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 4,000 ft (1,220 m). 12 April 1901. 121 Collector. W. W. Brown, Jr. Original number 28. Condition. Skin complete, with small bald spots. Skull complete. Mandible disarticulated. Type Series. There is a large series of paratypes in the MCZ. Comments. Type species of Isthmomys Hooper and Musser, 1964. I. flavidus was considered a valid species by Musser and Carleton (1993: 706) and Nowak (1999: 1357). Genus NECTOMYS Peters, 1861 Nectomys squamipes amazonicus Hershkovitz, 1944 Misc. Pub., Mus. Zool., Univ. Mich., 58: 47, 4 January. Holotype. MCZ 30820. Skin and skull. Male. Locality. Brazil: (Para), Rio Tapajos, Tauary (=Tau- ari). 23 January 1934. Collector. A. M. Olalla. Original number 7312. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Hershkovitz examined 28 specimens, in the MCZ, AMNH, and FMNH. Comments. Retained as a valid subspecies by Er- nest (1986: 1). Genus NEOTOMA Say and Ord, 1825 Neotoma abbreviata Goldman, 1909 Proc. Biol. Soc. Washington, 22: 140, 25 June. =Neotoma lepida abbreviata Goldman, 1909. See Burt (1932: 182). Holotype. MCZ 12260. Skin and skull. Adult male. Locality. (Mexico): Lower (=Baja) California, San Francisco Island. 22 February 1909. Collector. W. W. Brown, Jr. Original number 22. Condition. Skin and skull complete. Type Series. 9 paratypes; MCZ 12256-12259, 12261-12265; all represented by skin and skull, 5 females and 4 males. MCZ 12265 is now in the USNM. Comments. N. |. abbreviata was retained as a valid subspecies by Hall (1981: 755). Neotoma bella Bangs, 1899j Proc. New England Zool. Club, 1: 66, 31 July. =Neotoma lepida lepida Thomas, 1893. See Goldman (1932: 62). Holotype. B5308. Skin and skull. Adult male. Locality. (United States): California, Riverside County, Palm Springs. 12 April 1896. Collector. E. C. Thurber. Original number 623. Condition. Skin and skull complete. Mandible dis- articulated and left angular process chipped. IY Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Type Series. Holotype only. Neotoma distincta Bangs, 1903a Proc. Biol. Soc. Washington, 16: 89, 25 ijjunmies =Neotoma mexicana distincta Bangs, 1903. See Hall (1955: 329). Holotype. B9819. Skin and skull. Adult male. Locality. Mexico: Vera Cruz, near Jalapa, Texolo. § March 1899. Collector. S. N. Rhoads. Original number 295. Condition. Skin and skull complete. Type Series. 4 paratypes; B9818, skin and skull, male, B9820, skin and skull, female; B9821, skin and skull, male (exchanged to USNM); B9822, skin and skull, male. Comments. N. m. distincta was retained as a valid subspecies by Hall (1981: 761) and Cornely and Baker (1986: 1). Neotoma floridana rubida Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 185, 15 March. Holotype. B2872. Skin and skull. Adult male. Locality. (United States): Louisiana, Terrebonne Parish, Gibson. 4 April 1895. Collector. F. L. Small. Original number 1751. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 specimens are mentioned by num- ber in the original description: B2871, skin and skull, adult male, and B2873, skin and skull, adult female. Comments. Retained as a valid subspecies by Hall (1981: 749). Genus NYCTOMYS de Saussure, 1860 Nyctomys nitellinus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 30, April. =Nyctomys sumichrasti nitellinus Bangs, 1902. See Goldman (1916: 155). Holotype. MCZ 10249. Skin and skull. Adult female. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 4,000 ft (1,220 m). 8 February 1901. Collector. W. W. Brown, Jr. Original number 119. Condition. Skin and skull complete. Type Series. 5 paratypes; MCZ 10245, skin and skull, adult male; MCZ 10246, skin and skull, adult male; MCZ 10247, adult female, skin and skull: MCZ 10248, skin and skull, adult female; MCZ 10250, skin and skull, juvenile female. Comments. N. s. nitellinus was retained as a valid subspecies by Hall (1981: 630). Genus OECOMYS Thomas, 1906 Oecomys trabeatus G. M. Allen and Barbour, 1923 Bull. Mus. Comp. Zool., 65: 262, February. =Oecomys bicolor trabeatus G. M. Allen and Barbour, 1923. See comments. Holotype. MCZ 19837. Skin and skull. Male. Locality. Panama: Rio Jesusito. 10 April 1922. Collector. T. Barbour and W. S. Brooks. Original number 2027. Condition. Skin complete. Skull partial (right zy- gomatic arch broken). Mandible disarticulated. Type Series. Holotype only. Comments. Following Hershkovitz (1960: 533), O. b. trabeatus was retained as a valid subspecies by Hall (1981: 619) under the genus Oryzomys. The use of Oecomys as a generic name follows Musser and Carleton (1993: 715). Oryzomys flavicans illectus Bangs, 1898k Proc. Biol. Soc. Washington, 12: 164, 10 August. =Oecomys flavicans illectus (Bangs, 1898k). See comments. Holotype. BS101. Skin and skull. Adult female. Locality. Colombia: Magdalena, Santa Marta Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 24 March 1898. Collector. W. W. Brown, Jr. Original number 89. Condition. Skin complete. Skull partial (right jugal missing). Type Series. There is a small series of paratypes in the MCZ. Comments. The use of Oecomys as a generic name follows Musser and Carleton (1993: 715). Genus OLIGORYZOMYS Bangs, 1900 Oryzomys navus Bangs, 1899a Proc. Biol. Soc. Washington, 13: 9, 31 January. =Oligoryzomys fulvescens (Saussure, 1860). See Carleton and Musser (1989: 70). Holotype. BS107. Skin and skull. Adult male. Locality. Colombia: Magdalena, Santa Marta Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 26 March 1898. Collector. W. W. Brown, Jr. Original number 154. Condition. Skin and skull complete. Mandible dis- _ articulated. Type Series. 9 paratypes; BS223-B8231; all repre- sented by skin and skull, except B8228 (skin only); 2 females and 7 males. B8224 is now in the USNM, ° TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden and B8223, B8226, B8230, and B8231 are now in the AMNH. Comments. Oryzomys navus is the type species of Oligoryzomys Bangs, 1899. Oryzomys (Oligoryzomys) vegetus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 35, April. = Oligoryzomys vegetus (Bangs, 1902). See Musser and Carleton (1993: 718). Holotype. MCZ 10298. Skin and skull. Adult female. Locality. Panama: Chiriqui, Volcan de Chiriqui, Boquete, 4,000 ft (1,220 m). 16 April 1901. Collector. W. W. Brown, Jr. Original number 304. Condition. Skin complete. Skull partial (left jugal missing). Mandible disarticulated. Type Series. 12 paratypes; MCZ 10295 (now in USNM), 10297, 10300—10306, 10308—10310, 2 fe- males and 10 males, all represented by skin and skull except for 10308 (skin only). Comments. Considered a valid species by Musser and Carleton (1993: 718) and Nowak (1999: 1368). Carleton and Musser (1995) provided distribution- al information and support of specific status for O. vegetus. Genus ORYZOMYS Baird, 1858 Oryzomys devius Bangs, 1902b Bull. Mus. Comp. Zool., 39: 34, April. Holotype. MCZ 10324. Skin and skull. Adult female. Locality. Panama: Chiriqui, Volcan de Chiriqui, Boquete (=Bajo Boquete), 5,000 ft (1,525 m). 29 January 1901. Collector. W. W. Brown, Jr. Original number 73. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; MCZ 10325, skin and skull, adult female; MCZ 10326, skin and skull, adult female; MCZ 10340, skin and skull, adult male. Comments. Considered a valid species by Musser and Carleton (1993: 722) and Nowak (1999: 1366). Oryzomys palustris coloratus Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 189, 15 March. = Oryzomys palustris natator Chapman, 1893. See Wolfe (1982: 1). Holotype. B4470. Skin and skull. Adult male. Locality. (United States): Florida, Monroe County, Cape Sable. 17 April 1895. Collector. C. L. Brownell. Original number 148. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. 123 Oryzomys rostratus carrorum Lawrence, 1947b Proc. New England Zool. Club, 24: 101, 29 May. Holotype. MCZ 41280. Skin, skull, and postcranial skeleton. Adult male. Locality. Mexico: Tamaulipas, Rio Soto la Marina, Rancho Santa Ana, 8 miles (12.9 km) southwest of Padilla. 21 December 1941. Collector. G. H. Pournelle, Third University of Florida Mexican Expedition. Original number 196. Condition. Skin and skull complete. Mandible dis- articulated. Postcranial skeleton partial (right fore- limb, hindlimbs, pelvis, ribs, vertebral column). Type Series. 2 paratypes; MCZ 41281, skin, skull, and postcranial skeleton, female; MCZ 41282, skin, skull, and postcranial skeleton, male. Comments. Retained as Oryzomys melanotis car- rorum by Hall (1981: 614). Musser and Carleton (1993: 724) regarded O. rostratus as distinct from O. melanotis. — Genus PEROMYSCUS Gloger, 1841 Hesperomys gossypinus LeConte, 1853 Proc. Acad. Nat. Sci. Philadelphia, 6: 411, 25 October. = Peromyscus gossypinus gossypinus (LeConte, 1853). See Rhoads (1896b: 189). Lectotype. MCZ 5275. Skin and skull. Adult male. Locality. (United States): Georgia, Liberty County, Riceboro, probably on the LeConte Plantation near Riceboro. 13 September 1847. Collector. J. E. LeConte. Formerly USNM 546. Condition. Skin complete. Skull partial (basioccip- ital and tympanic bullae broken). Mandible disar- ticulated. Type Series. Poole and Schantz (1942: 320) listed potential “cotypes,” meaning syntypes. Osgood (1909: 136) mentioned that a specimen, number 752 in the collection of the Academy of Natural Sciences in Philadelphia, possibly possessed type status, but this specimen was not mentioned by Koopman (1976). Comments. In the original description of gossypi- nus, measurements were given for a single speci- men only; however, in addition to this specimen, LeConte made reference to the coloration of “younger individuals.” The measurements given correspond to MCZ 5275 rather than to any other of LeConte’s specimens still in the USNM (see G. M. Allen, 1931: 262: Poole and Schantz, 1942: 320). In addition, MCZ 5275 was the only speci- men of Peromyscus gossypinus from Georgia men- tioned in Baird’s Mammals of North America (1858: 469). Because LeConte did not specify a ho- lotype, MCZ 5275, which seems to be the speci- 124 men discussed and described by measurement in the original publication and the only adult speci- men mentioned, is here designated as lectotype. P. gossypinus was considered a valid species by Mus- ser and Carleton (1993: 730) and Nowak (1999: 1361). Hesperomys leucopus arcticus Mearns, 1890 Bull. Amer. February. Name preoccupied by Hesperomys arcticus Coues, 1877 Mus. Nat. Hist., 2: 285, 21 Peromyscus maniculatus borealis Mearns, 1911 Proc. Biol. Soc. Washington, 24: 102, 15 May. (Replacement name for Hesperomys leucopus arcticus Mearns, 1890) Holotype. MCZ 5555. Skin and skull. Adult male. Locality. Canada: Northwest Territories, Mac- Kenzie, Fort Simpson. 7 September 1859. Collector. R. Kennicott. Original number 157. For- merly USNM 4531. Condition. Skin complete. Skull partial (left jugal and process of squamosal missing, right process of squamosal missing, basioccipital and palatine miss- ing). Mandible disarueulated and both angular pro- cess missing. Type Series. Holotype only. Comments. P. m. borealis was retained as a valid subspecies by Hall (1981: 672). Hesperomys sonoriensis nebrascensis Coues, 1877 In Coues and J. A. Allen, Monographs N. Amer. Rodentia, U.S. Geol. Geogr. Survey Terr. Rep. Washington, 11: 79, August. = Peromyscus maniculatus nebrascensis (Coues, 1877). See Osgood (1909: 75). Syntype. MCZ 5528. Skin and skull. Locality. Nebraska (now Wyoming): County), Deer Creek. 19 January 1860. Collector. F. V. Hayden. Original no. 80. Formerly USNM 4310. Condition. Skin complete. Skull partial (posterior missing from parietal to palatine; both jugal and process of squamosal broken). Left angular process of mandible chipped, right side broken at cheek teeth. Type Series. | other syntype, present location un- known, formerly in the USNM. Comments. Hesperomys sonoriensis nebrascensis (Converse Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Baird, 1858 is a nomen nudum. Osgood (1909: 78) and Mearns (1911: 102) designated Deer Creek, Nebraska, as the type locality. Jones (1958: 107— 111) reviewed the systematic history of nebrascen- sis, and Jones and Mursaloglu (1961: 101-103) re- ported the discovery of one oe the two original syn- types in the MCZ. The location of the other syn- type, if it still exists, is unknown. Peromyscus anastasae Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 195, 15 March. = Peromyscus gossypinus anastasae Bangs, 1898. See Osgood (1909: 141). Holotype. B7179. Skin and skull. Adult female. Locality. (United States): Florida, St. Johns Coun- ty, Anastasia Island, Point Romo. 15 February 1897. Collector. O. Bangs. Original number 2. Condition. Skin complete, skull partial (premaxilla to frontal in fragments). Right mandible missing, left present. Type Series. There is a small series of paratypes in the MCZ. Comments. P. g. anastasae was retained as a valid subspecies by Hall (1981: 689). Peromyscus bellus Bangs, 1896h Proc. Biol. Soc. Washington, 10: 137, 28 December. = Peromyscus attwateri J. A. Allen, 1895. See Schmidly (1973: 125). Holotype. B5483. Skin and skull. Adult male. Locality. (United States): Indian Territory (=Oklahoma), (Adair County), Stilwell. 15 August 1896. Collector. T. Surber. Original number 67. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. adult female. 1 paratype; B5484, skin and skull, Peromyscus cacabatus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 29, April. = Peromyscus mexicanus (Saussure, 1860). See Huckaby (1980: 15). Holotype. MCZ 10225. Skin and skull. Adult female. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 5,000 ft (1,525 m). 22 April 1901. Collector. W. W. Brown, Jr. Original number 331. Condition. Skin complete. Skull partial (left jugal missing). Mandible disarticulated. Type Series. There is a very large series of paraty- pes in the MCZ. TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden Peromyscus canadensis abietorum Bangs, 1896d Proc. Biol. Soc. Washington, 10: 49, 9 March. =Peromyscus maniculatus abietorum Bangs, 1896. See Osgood (1909: 45). Holotype. B2205. Skin and skull. Adult female. Locality. (Canada): Nova Scotia, James River. 8 August 1894. Collector. C. H. Goldthwaite. Condition. Skin complete. Skull partial (right jugal missing). Type Series. There is a series of paratypes in the MCZ. Comments. P. m. abietorum was retained as a valid subspecies by Hall (1981: 670). Peromyscus canadensis argentatus Copeland and Church, 1906 Proc. Biol. Soc. Washington, 19: 122, 6 September. = Peromyscus maniculatus argentatus Copeland and Church, 1906. See Osgood (1909: 46). Holotype. MCZ 54627. Skin and skull. Adult male. Locality. (Canada): New Brunswick, Grand Manan Island, Grand Harbor. 19 September 1905. Collector. M. L. Church and M. Copeland. Original number 168. Condition. Skin and skull complete. Type Series. There is a large series of paratypes in the MCZ. Comments. P. m. argentatus was retained as a valid subspecies by Hall (1981: 671). Peromyscus crinitus scitulus Bangs, 1899j Proc. New England Zool. Club, 1: 67, 31 July. =Peromyscus crinitus crinitus (Merriam, 1891). See Osgood (1909: 229). Holotype. B9175. Skin and skull. Adult male. Locality. (United States): Nevada, Douglas County, Gardnerville. 13 July 1898. Collector. W. W. Price and P. O. Simons. Original number 1571. Condition. Skin complete. Skull partial (right and left jugal missing). Type Series. 7 paratypes; B9177—B9180, B9183— B9185; all represented by skin and skull, 4 females and 3 males. 125 Peromyscus gossypinus nigriculus Bangs, 1896e Proc. Biol. Soc. Washington, 10: 124, 5 November. =Peromyscus gossypinus gossypinus (LeConte, 1853). See Osgood (1909: 136). Holotype. B2731. Skin and skull. Adult female. Locality. (United States): Louisiana, Plaquemines Parish, Burbridge. 30 January 1895. Collector. F. L. Small. Original number 1547/156. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a very large series of paraty- pes in the MCZ. Peromyscus gossypinus palmarius Bangs, 1896e Proc. Biol. Soc. Washington, 10: 124, 5 November. Holotype. B3224. Skin and skull. Adult female. Locality. (United States): Florida, Brevard County, Oak Lodge on East Peninsula opposite Micco. 23 February 1895. Collector. O. Bangs. Condition. Skin and skull complete. Type Series. There is a very large series of paraty- pes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 690). Peromyscus insulanus Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 196, 15 March. = Peromyscus gossypinus anastasae Bangs, 1898. See Osgood (1909: 141). Holotype. B6438. Skin and skull. Adult male. Locality. (United States): Georgia, Camden Coun- ty, Cumberland Island, north end. 10 April 1897. Collector. W. W. Brown, Jr. Original number 804. Condition. Skin complete. Skull partial (right jugal missing). Type Series. There is a large series of paratypes in the MCZ. Peromyscus leucopus ammodytes Bangs, 1905a Proc. New England Zool. Club, 4: 14, 28 February. Holotype. B828. Skin and skull. Adult male. Locality. (United States): Massachusetts, Barnsta- ble County, Monomoy Island. 28 December 1893. Collector. G. S. Miller, Jr. and O. Bangs. Original number 8. Condition. Skin and skull complete. 126 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Type Series. There is a series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 685). Peromyscus leucopus fusus Bangs, 1905a Proc. New England Zool. Club, 4: 13, 28 February. Holotype. B9737. Skin and skull. Adult male. Locality. (United States): Massachusetts, Dukes County, Martha’s Vineyard, W. Tisbury. 17 June 1899. Collector. O. Bangs. Original number 1. Condition. Skin and skull complete. Type Series. There is a small series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 686). Peromyscus oreas Bangs, 1898f Proc. Biol. Soc. Washington, 12: 84, 24 March. Holotype. B3696. Skin and skull. Adult female. Locality. Canada: British Columbia, near boundary of Whatcom County, Washington, 49th parallel, Mount Baker Range, 6,500 ft (1,983 m). 29 August 1896. Collector. A. C. Brooks. Original number 745. Condition. Skin and skull complete. Type Series. There is a large series of paratypes in the MCZ. Comments. Considered a valid species by Musser and Carleton (1993: 734) and Nowak (1999: 1361). Peromyscus phasma Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 199, 15 March. =Peromyscus poloniotus phasma Bangs, 1898. See Osgood (1909: 107). Holotype. B7175. Skin and skull. Adult female. Locality. (United States): Florida, St. Johns Coun- ty, Anastasia Island, Point Romo. 14 February 1897. Collector. O. Bangs. Original number 3. Condition. Skin complete. Skull partial (parietal, tympantic bulla, left jugal, and process of squa- mosal broken). Mandible disarticulated. Type Series. There is a series of paratypes in the MCZ. Comments. P. p. phasma was retained as a valid subspecies by Hall (1981: 668). Peromyscus subgriseus arenarius Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 202, March. Name preoccupied by Peromyscus eremicus arenarius Mearns, 1896. Peromyscus subgriseus baliolus Bangs, 1898n Science, my ser.,.6: Zila. lOyAueust (Replacement name for Peromyscus subgriseus arenarius Bangs, 1898) =Peromyscus polionotus polionotus (Wagner, 1843). See Osgood (1909: 104). Holotype. B5925. Skin and skull. Adult male. Locality. (United States): Georgia, Scriven County, Hursman’s Lake (Savannah River), near Bascom. 15 December 1896. Collector. W. W. Brown, Jr. Original number 60. Condition. Skin and skull complete. Type Series. There is a large series of paratypes in the MCZ. Peromyscus subgriseus rhoadsi Bangs, 1898b Proc. Boston Soc, Nat, Hist; 28: 2015 V5 March. =Peromyscus polionotus rhoadsi Bangs, 1898. See Osgood (1909: 107). Holotype. B6980. Skin and skull. Adult male. Locality. (United States): Florida, Hillsborough County, head of Anclote River. 23 May 1895. Collector. W. S. Dickinson. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. P. p. rhoadsi was retained as a valid subspecies by Hall (1981: 669). Peromyscus texanus saturatus Bangs, 1897a Amer. Nat. 31: 75, 1 January. =Peromyscus maniculatus saturatus Bangs, 1897. See Osgood (1909: 61). Holotype. B2581. Skin and skull. Adult male. Locality. Canada: British Columbia, Saturna Is- land. 31 January 1894. Collector. W. C. Colt. Original number 128. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. P. m. saturatus was retained as a valid subspecies by Hall (1981: 682). TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden 1 Genus REITHRODONTOMYS Giglioli, 1873 Reithrodontomys australis vulcanius Bangs, 1902b Bull. Mus. Comp. Zool., 39: 38, April. = Reithrodontomys sumichrasti vulcanius Bangs, 1902. See Hooper (1952: 83). Holotype. MCZ 10281. Skin and skull. Adult male. Locality. Panama: Chiriqui, Volcan de Chiriqui, 10,300 ft (3,142 m). 26 May 1901. Collector. W. W. Brown, Jr. Original number 500. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. R. s. vulcanius was retained as a valid subspecies by Hall (1981: 644). Reithrodontomys creper Bangs, 1902b Bull. Mus. Comp. Zool., 39: 39, April. Holotype. MCZ 10284. Skin and skull. Adult female. Locality. Panama: Chiriqui, Volcan de Chiriqui, 11,000 ft (3,355 m). 2 June 1901. Collector. W. W. Brown. Original number 504. Condition. Skin and skull complete. Mandible dis- articulated. Left lower incisor missing. Type Series. Holotype only. Comments. Considered a valid species by Musser and Carleton (1993: 740) and Nowak (1999: 1364). Reithrodontomys lecontii impiger Bangs, 1898m Proc. Biol. Soc. Washington, 12: 167, 10 August. = Reithrodontomys humulis humulis (Audubon and Bachman, 1841). See Hall and Kelson (1959: 584). Holotype. B7784. Skin and skull. Adult male. Locality. (United States): West Virginia, Green- briar County, White Sulphur Springs, 2,000 ft (610 m). 27 February 1898. Collector. T. Surber. Original number 466. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 paratypes; B6931 (not B6932, as stated in the original description), skin and skull, adult male; B7785, skin and skull, adult female. Genus SCOTINOMYS Thomas, 1913 Akodon teguina apricus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 40, April. = Scotinomys teguina apricus (Bangs, 1902). See Thomas (1913: 409). Holotype. MCZ 10236. Skin and skull. Adult female. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 4,000 ft (1,220 m). 24 February 1901. ho ~ Collector. W. W. Brown, Jr. Original number 192. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 4 paratypes; MCZ 10234, skin and skull, adult male; MCZ 10235, skin, adult female; MCZ 10237, skin and skull, adult female; MCZ 10238, skin and skull, adult female. Comments. S. t. apricus was retained as a valid sub- species by Hooper (1972: 21) and Hall (1981: 734). Akodon xerampelinus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 41, April. = Scotionomys xerampelinus (Bangs, 1902). See Thomas (1913: 409). Holotype. MCZ 10240. Skin and skull. Adult male. Locality. Panama: Chiriqui, Volcan de Chiriqui, 10,300 ft (3,142 m). 26 May 1901. Collector. W. W. Brown, Jr. Original number 501. Condition. Skin complete. Skull partial (right jugal missing). Mandible disarticulated. Type Series. 2 paratypes; MCZ 10239, skin and skull, adult male; MCZ 10241, skin and skull, adult male. Comments. S. xerampelinus was considered a valid species by Musser and Carleton (1993: 746) and Nowak (1999: 1355). Genus SIGMODON Say and Ord, 1825 Sigmodon austerulus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 32, April. = Sigmodon hispidus borucae J. A. Allen, 1897. See Hall and Kelson (1959: 672). Holotype. MCZ 10288. Skin and skull. Adult male. Locality. Panama: Chiriqui, Volcan de Chiriqui, 10,000 ft (3,050 m). 1 June 1901. Collector. W. W. Brown, Jr. Original number 503. Condition. Skin complete. Skull partial—according to original description, “unfortunately it [the skull] was broken by the trap directly across the orbits.” Type Series. Holotype only. Comments. For comments regarding the type lo- cality, see Enders (1953a: 508-509), who suggested that a native helper of W. W. Brown, Jr., transplant- ed a specimen of S. [h.] borucae, trapped during a trip to his home in the lowlands near Boruca, to the top of the Volcan, “after Brown had promised a bottle of wine to any man who captured a Sig- modon at that altitude.” Sigmodon hispidus exsputus G. M. Allen, 1920b J. Mammal., 1: 236, 4 December. Holotype. MCZ 18100. Skin and skull. Adult male. Locality. (United States): Florida, Monroe County, Big Pine Key. 16 April 1920. Collector. W. S. Brooks. Original number 1936. 128 Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; MCZ 18101, skin and skull, adult male. Comments. Retained as a valid subspecies by Hall (1981: 737). Sigmodon hispidus furvus Bangs, 1903b Bull. Mus. Comp. Zool., 39: 158, July. Holotype. MCZ 10665. Skin and skull. Male. Locality. Honduras: Atlantida, La Ceiba. 16 Janu- ary 1902. Collector. W. W. Brown, Jr. Original number 4. Condition. Skin and skull complete. Type Series. Holotype only. Comments. Retained as a valid subspecies by Hall (1981: 738). Sigmodon hispidus fervidus Lydekker, 1904 (p. 34) was a misspelling and thus accidental renaming of Sigmodon hispidus furvus Bangs, 1903. Sigmodon hispidus spadicipygus Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 192, 15 March. Holotype. B4477. Skin and skull. Adult female. Locality. (United States): Florida, Monroe County, Cape Sable. 18 April 1895. Collector. C. L. Brownell. Original number 153. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 739). Sigmodon sanctae-martae Bangs, 1898q Proc. Biol. Soc. Washington, 12: 189, 30 December. = Sigmodon hispidus hirsutus (Burmeister, 1854). See Cabrera (1961: 508). Holotype. B8105. Skin and skull. Adult male. Locality. Colombia: (Magdalena), (Santa Marta Mountains), Pueblo Viejo (not far from the source of Rio Ancho), 8,000 ft (2,440 m). 23 March 1898. Collector. W. W. Brown, Jr. Original number 73. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; B8250, skin and _ skull, adult male. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Genus THOMASOMYS Coues, 1884 Oryzomys (Erioryzomys) monochromos Bangs, 1900b Proc. New England Zool. Club, 1: 97, 23 February. = Thomasomys monochromos (Bangs, 1900). See Gardner and Patton (1976: 26): Holotype. B8348. Skin and skull. Adult male. Locality. Colombia: Magdalena, Santa Marta Mountains, Paramo de Macotama, 11,000 ft (3,355 m). 7 March 1899. Collector. W. W. Brown, Jr. Original number 93. Condition. Skin complete. Skull partial (right jugal missing). Mandible disarticulated. Type Series. 4 paratypes; B8248, juvenile male, B8345, adult female, B8346, adult female, B8347, adult male; all represented by skin and skull. Comments. Considered a valid species by Musser and Carleton (1993: 750) and Nowak (1999: 1362). Paramo de Macotama is in La Guajira (Paynter 1997: 261). Genus ZYGODONTOMYS J. A. Allen, 1897 Zygodontomys seorsus Bangs, 1901b Amer. Nat., 35: 642, 22 August. =Zygodontomys brevicauda cherriei (J. A. Allen, 1895). See Voss (1991: 59). Holotype. B8490. Skin and skull. Adult male. Locality. Panama: Gulf of Panama, San Miguel Is- land, Isla del Rey. 5 May 1900. Collector. W. W. Brown, Jr. Original number 147. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Bangs based the description on a se- ries of 68 specimens, most of which are still in the MCZ. Family PEDETIDAE Gray, 1825 Genus PEDETES llliger, 1811 Pedetes cafer taborae G. M. Allen and Loveridge, 1927 Proc. Boston Soc. Nat. Hist., 38: 438, 23 December. =Pedetes capensis (Forster, 1778). See Misonne (1974: 8). Holotype. MCZ 23080. Skin and skull. Adult female. Locality. Tanganyika Territory (=Tanzania): Ta- bora, Unyamwezi, 4,000 ft (1,220 m). 18 November 1921. Collector. A. Loveridge. Original number R6915. TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Condition. Skin complete, with small bald spots on dorsum. Skull complete. Type Series. Holotype only. Family MYOXIDAE Gray, 1821 Genus GRAPHIURUS Smuts, 1832 Aethoglis hueti argenteus G. M. Allen, 1936 J. Mammal., 17: 293, 14 August. Se eg Cet argenteus (G. M. Allen, 1936). See Hayman and Holt (1940: 608). Holotype. MCZ 17920. Skin and skull. Adult female. Locality. Cameroons (=Cameroon): (Kribi), Lolo- dorf. 17 March 1911. Collector. G. Schwab. Original number 1. Condition. Skin and skull complete. Type Series. 2 paratypes; MCZ 17607, skin and skull, adult female; and a specimen in the USNM, number 125434. Comments. Rosevear considered G. h. “possibly valid as a race” (1969: 501). argen teus Claviglis soleatus collaris G. M. Allen and Loveridge, 1933 Bull. Mus. Comp. Zool., 75: February. = Graphiurus lorraineus Dollman, 1910. See Holden (1993: 764). 122, Mas MCZ 26373. Skin and skull. Adult female. Locality. Tanganyika Territory (=Tanzania): (Irin- ga), north end Lake Nyasa, Ukinga Mountains, Ma- dehani, 7,000 ft (2,135 m). 24 February 1930. Collector. A. Loveridge. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 paratypes; MCZ 26374, skin and skull, male; MCZ 26375, skin and skull, female. Graphiurus microtis griseus G. M. Allen, 1912a Bull. Mus. Comp. Zool., 54: 440, April. =Graphiurus murinus griseus G. M. Allen, 1912. See Hayman and Holt (1940: 610). Holotype. MCZ 8244. Skin and skull. Adult male. Locality. British East Africa (=Kenya): northern a aso Nyiro River (=Ewaso Ngiro), 4,000 ft (1,220 m). 25 July 1909. Gollector. G. M. Allen. Original number 51. Condition. Skin and skull complete. Type Series. 2 paratypes; MCZ 8248, skin and skull, male; MCZ 8249, skin and skull, adult female. 129 Graphiurus schwabi G. M. Allen, 1912a Bull. Mus. Comp. Zool., 54: 441, April. = Graphiurus surdus Dollman, 1912. See Holden (1993: 765). Holotype. MCZ 8607. Skin and skull. Juvenile. Locality. Cameroun (=Cameroon), Kribi. 1911. Collector. G. Schwab. Condition. Skin and skull complete. Type Series. Holotype only. Family BATHYERGIDAE Waterhouse, 1841 Genus CRYPTOMYS Gray, 1864 Cryptomys hottentotus occlusus G. M. Allen and Loveridge, 1933 Bull. Mus. Comp. Zool., 75: 125, February. =Cryptomys hottentotus whytei (Thomas, 1897). See Honeycutt et al. (1991: 51). Holotype. MCZ 26557. Skin and skull. Adult male. Locality. Tanganyika Territory (=Tanzania): (Irin- a Ugungwe Mountains, Kigogo, 6,000 ft (1,830 m) 13) Janu: ary 1930. @pllcciar A. Lov eridge. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 15 paratypes; MCZ 26558-26572, all represented by skin and skull. Genus HETEROCEPHALUS Ruppell, 1842 Heterocephalus stygius G. M. Allen, 1912a Bull. Mus. Comp. Zool., 54: 444, April. = Heterocephalus glaber Ruppell, 1842. See Hollister (1919: 160). Holotype. MCZ 12470. Body in alcohol, skull extract- ed. Adult female. Locality. British East Africa ( Guaso Nyiro ( August 1909. Collector. G. M. Allen. Condition. Alcoholic. disarticulated. Type Series. Holotype only. =Kenya): northern =Ewaso Ngiro), Neumann’s Boma. 6 Skull complete. Mandible 130 Family ERITHIZONTIDAE Bonaparte, 1845 Genus ERITHIZON F. Cuvier, 1822 Erethizon dorsatus picinus Bangs, 1900c Proc. New England Zool. Club, 2: 37, 20 September. =Erithizon dorsatum dorsatum (Linnaeus, 1758). See Harper (1961: 90). Holotype. B8839. Skin and skull. Adult male. Locality. (Canada): Labrador, Strait of Belle Isle, Lance (=Lanse) au Loup. 16 F ebruary 1899. Collector. E. Doane. Condition. Skin and skull complete. Type Series. 15 paratypes; B8832—B8838, B8840— B8847; all represented by skin and skull, 8 females and 7 males. Bangs had also a skull of picinus (B8831) from Black Bay, Labrador, at the time of description, not mentioned in the original publi- cation. Family DASYPROCTIDAE Gray, 1825 Genus DASYPROCTA llliger, 1811 Dasyprocta callida Bangs, 1901b Amer. Nat., 35: 635, 22 August. =Dasyprocta punctata callida Bangs, 1901. See Kellogg (1946: 59). Holotype. B8443. Skin and skull. Adult male. Locality. Panama: (Panama), San Miguel Island. 8 May 1900. Collector. W. W. Brown, Jr. Original number 171. Condition. Skin and skull complete. Type Series. 5 paratypes; B8442, B8444—B8447;, all represented by skin and skull, 3 females and 2 males. Comments. D. p. callida was retained as a valid subspecies by Hall (1981: 860). Dasyprocta colombiana Bangs, 1898k Proc. Biol. Soc. Washington, 12: 163, 10 August. =Dasyprocta punctata colombiana (Gray, 1898). See Cabrera (1961: 589). BS8008. Skin and skull. Adult female. Magdalena, Santa Marta. 6 Holotype. Locality. Colombia: January 1898. Collector. W. W. Brown, Jr. Original number 37. Condition. Skin and skull complete. Type Series. | paratype; B8113, skin and skull, ju- venile male. Comments. D. p. colombiana was retained as a val- id subspecies by Cabrera (1961: 589). Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Dasyprocta noblei G. M. Allen, 1914e Proc. New England Zool. Club, 5: 69, 7 October. =Dasyprocta leporina noblei G. M. Allen, 1914. See Woods (1993: 781). Holotype. MCZ 15936. Skin and skull (and atlas). Adult female. Locality. Guadeloupe Island: Goyave. 22 August 1914. Collector. G. K. Noble. Condition. Skin complete. Skull partial (bone miss- ing from nasal and frontal). Type Series. 1 paratype; MCZ 15937, skull, subadult female. skin and Dasyprocta punctata nuchalis Goldman, A Oi7, Proc. Biol. Soc. Washington, 30: 113, 23 May. Holotype. MCZ 10081. Skin and skull. Adult female. Locality. Panama: Chiriqui, Divala. 30 November 1900. Collector. W. W. Brown, Jr. Original number 17. Condition. Skin and skull comple te. Type Series. 4 paratypes; MCZ 10080, skin and skull, adult male: 10084, skull: 10175, skin and skull, adult female; 10176, skin and skull, adult male. Comments. Retained as a valid subspecies by Hall (1981: S60). Family AGOUTIDAE Gray, 1821 Genus AGOUTI Lacépede, 1799 Agouti paca virgatus Bangs, 1902b Bull. Mus. Comp. Zool., 39: 47, April. Holotype. MCZ 10079. Skin and skull. Adult male. Locality. Panama: Chiriqui, Divala. 16 December 1900. Collector. W. W. Brown, Jr. Original number 21. Condition. Skin and skull complete. Type Series. Holotype only. Comments. Retained as a valid subspecies by Hall (1981: 858) and Pérez (1992: 1). Family OCTODONTIDAE Waterhouse, 1839 Genus TYMPANOCTOMYS Yepes, 1941 Octomys barrerae Lawrence, 1941 Proc. New England Zool. Club, 18: 43, 28 January. = Tympanoctomys barrerae (Lawrence, 1941). See Yepes (1942: 75). Holotype. MCZ 39716. Skin and skull. Adult male. Locality. Argentina: Mendoza Province, La Paz. April 1939. TYPE SPECIMENS OF RECENT MAMMALS ° Helgen and McFadden Collector. J. M. de la Barrera. Original number A381. Condition. Skin and skull complete. Type Series. Holotype only. Comments. Type species of Tympanoctomys Yepes, 1941. T. barrerae was considerd a valid species by Woods (1993: 789) and Nowak (1999: 1683). Family ECHIMYIDAE Gray, 1825 Genus BOROMYS Miller, 1916 Boromys torrei G. M. Allen, 1917a Bull. Mus. Comp. Zool., 61: 6, January. Holotype. VP 9601. Palate. Locality. Cuba: Matanzas Province, cave in Sierra de Hato Nuevo. Collector. C. de la Torre. Condition. Condition as described in original “palate with root of right zygomatic arch, pm‘ and alveolar row of right side, m! and posterior part of alveolar row of left side.” Type Series. Paratype material consists of 8 lower jaws and 2 separate lower molars; in the MCZ. Comments. Considered a valid species by Woods (1993: 799) and Nowak (1999: 1703) but almost certainly extinct. This specimen is stored in the Vertebrate Paleontology Department of the MCZ. Genus DIPLOMYS Thomas, 1916 Loncheres labilis Bangs, 1901b Amer. Nat., 35: 638, 22 August. =Diplomys labilis (Bangs, 1901). See Thomas (1916: 296). Holotype. B8480. Skin and skull. Adult male. Locality. Panama: (Panama), Gulf of Panama, San Miguel Island. 26 April 1900. Collector. W. W. Brown, Jr. Original number 103. Condition. Skin and skull complete. Type Series. There is a small series of paratypes in the MCZ and FMNH. Comments. D. labilis was considered a valid spe- cies by Woods (1993: 791) and Nowak (1999: 1695). Genus PROECHIMYS J. A. Allen, 1899 Proechimys burrus Bangs, 1901b Amer. Nat., 35: 640, 22 August. =Proechimys semispinosus burrus Bangs, 1901. See Goldman (1920: 120). Holotype. B8458. Skin and skull. Adult male. Locality. Panama: (Panama), Gulf of Panama, San Miguel Island. 30 April 1900. Collector. WW. Brown, Jr. Original number 130. Condition. Skin and skull complete. Right dentary broken at pm. e331 Type Series. There is a large series of paratypes in the MCZ. Comments. P. s. burrus was retained as a valid sub- species by Hall (1981: 873). Proechimys gorgonae Bangs, 1905b Bull. Mus. Comp. Zool., 46: 89, June. Holotype. MCZ 10828. Skin and skull. Adult male. Locality. Colombia: (Cauca), Gorgona Island. 2 July 1904. Collector. W. W. Brown, Jr. Original number 25. Condition. Skin and skull complete. Type Series. 6 paratypes; MCZ 10829-10834, all represented by skin and skull, 1 female and 5 males. MCZ 10830 and 10834 are now in the FMNH. Comments. Treated as a separate species by Woods (1993: 796) but included in P. cayennensis by No- wak (1999: 1689). Proechimys guyannensis hyleae Moojen, 1948 Univa Kansas Bulbl) Mus! Nataelelist ml: 361, 10 December. =Proechimys cayennensis hyleae Moojen, 1948. See comments. Holotype. MCZ 30887. Skin and skull. Adult male. Locality. Brazil: Para, Porto de Moz, Rio Tapajoz (=Tapajos), Tauari (=Tauary), 87 km south of San- tarem. 19 January 1934. Collector. A. M. Olalla. Original number 7288. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 20 paratypes (19 at MCZ and 1 at FMNH), represented by skin and skull. Comments. Retained as a valid subspecies by Ca- brera (1961: 521). The use of the specific name cayennensis over guyannensis follows Woods (i993 7795): Family CAPROMYIDAE Smith, 1842 Genus CAPROMYS Desmarest, 1822 Capromys pilorides relictus G. M. Allen, 1911a Bull. Mus. Comp. Zool., 54: 207, July. Holotype. MCZ 10996. Skin and skull. Adult male. Locality. Cuba: Isle of Pines (=Isla de Pinos), Nue- va Gerona, Casas Mountains. 10 March 1902 Collector. W. R. Zappey. Condition. Skin complete. Skull partial (bone miss- ing from palatine to occipital). Type Series. 1 paratype; MCZ 10997, skin and skull. Comments. Retained as a valid subspecies by Hall (1981: 863). 132 Genus GEOCAPROMYS Chapman, 1901 Geocapromys cubanus G. M. Allen, 1917a Bull. Mus. Comp. Zool. 61: 9, January. = Geocapromys columbianus (Chapman, 1892). See G. M. Allen (1918b: 145). Holotype. VP 9602. Part of right lower ramus. Juve- nile. Locality. Cuba: Matanzas Province, cave in Sierra de Hato Neuvo. Collector. C. de la Torre. Condition. Portion of ramus includes incisor and three anterior cheekteeth. Type Series. Allen mentions in the original descrip- tion, “Five palates with teeth, about 15 jaw frag- ments mostly with teeth, and numerous other frag- ments.” This material is in the MCZ. Comments. The type material of cubanus is stored in the Vertebrate Paleontology Department of the MCZ. Geocapromys ingrahami abaconis Lawrence, 1934 Occas. Pap. Boston Soc. Nat. Hist., 8: 190, 7 November. Holotype. VP 2108. Left lower ramus. Adult. Locality. Bahamas: Great Abaco Island, Hole in the Wall, Imperial Lighthouse Caves. 1934. Collector. F. Rainey. Condition. Ramus intact. Type Series. The paratype series is a collection of cranial and mandibular fragments, described in the original description and housed in the MCZ. Comments. G. i. abaconis was retained as a valid subspecies by Hall (1981: 866) under the genus Capromys. Woods (1993: 800) employed the genus Geocapromys. G. i. abaconis is extinct. The type material of abaconis is stored in the Vertebrate Pa- leontology Department of the MCZ. Geocapromys ingrahami irrectus Lawrence, 1934 Occas. Pap. Boston Soc. Nat. Hist., 8: 190, 7 November. Holotype. VP 2107. Right lower ramus. Adult. Locality. Bahamas: Crooked Island, Gordon Hill Caves, “Burial Cave No. 1.” 1934. Collector. F. Rainey. Condition. Ramus intact. Type Series. The paratype series is a collection of cranial and mandibular fragments, described in the original description and housed in the MCZ. Comments. G. i. irrectus was retained as a valid subspecies by Hall (1981: 866) under the genus Capromys. Woods (1993: 800) employed the genus Geocapromys. G. i. irrectus is extinct. The type ma- Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 terial of irrectus is stored in the Vertebrate Pale- ontology Department of the MCZ. Genus MESOCAPROMYS Varona, 1970 Capromys nana G. M. Allen, 1917b Proc. New England Zool. Club, 6: 54, 28 March. =Mesocapromys nanus (G. M. Allen, 1917). See Kratochvil et al. (1978: 15). Holotype. VP 9864. Right mandible. Locality. Cuba: Matanzas, cave in Sierra de Hato Nuevo. March 1917. Collector. T. Barbour. Condition. Mandible partial—coronoid and angu- lar processes broken off. Type Series. A series of paratype material is stored at the MCZ. Comments. G. M. Allen supplemented his descrip- tion of Capromys nanus after Thomas Barbour provided him with a freshly killed specimen in 1918 (1918b: 140-145). Type species of the sub- genus Paracapromys Kratochvil, Rodriguez, and Barus, 1978. M. nanus was considered a valid spe- cies by Woods (1993: 801) and Nowak (1999: 1703). The type series is stored in the Vertebrate Paleontology Department of the MCZ. Genus PLAGIODONTA F. Cuvier, 1836 Plagiodonta araeum Ray, 1964 Breviora, Mus. Comp. Zool., 203: 2, 10 April. Holotype. VP 7675. Left upper cheektooth, originally described as “almost certainly the fourth (decidu- ous) premolar.” Locality. Dominican Republic: San Rafael Prov- ince, Hondo Valle Municipality, unnamed cave 2 km southeast of Rancho de La Guardia. April 1963. Collector. R. Allen and C. E. Ray. Condition. Condition of tooth as originally de- scribed—damaged slightly along the posterolabial wall.” Type Series. Holotype only. Comments. Considered a valid species by Woods (1993: 804) and Nowak (1999: 1708). Complete cranial and dentary material of araewm has been collected in Haiti and deposited in the Florida Mu- seum of Natural History (Woods 1993: 804). This species is extinct. This specimen is stored in the Vertebrate Paleontology Department of the MCZ. o/ TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden Order LAGOMORPHA Brandt, 1855 Family OCHOTONIDAE Thomas, 1897 Genus OCHOTONA Link, 1795 Ochotona cuppes Bangs, 1899g Proc. New England Zool. Club, 1: 40, 5 June. =Ochotona princeps cuppes Bangs, 1899. See A. H. Howell (1924: 27). Holotype. B7389. Skin and skull. Adult male. Locality. (Canada): British Columbia, Gold Range, Monishee Divide, 4,000 ft (1,220 m). 2 August 1897. Collector. A. C. Brooks. Original number 1030. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; MCZ 7390-7392; all rep- resented by skin and skull, all female. MCZ 7390 is now at FMNH. Comments. O. p. cuppes was retained as a valid subspecies by Hall (1981: 289) and Smith and Wes- ton (1990: 1). Ochotona saxatilis Bangs, 1899g Proc. New England Zool. Club, 1: 41, 5 June. =Ochotona princeps saxatilis Bangs, 1899. See A. H. Howell (1924: 27). Holotype. MCZ 2703. Skin. Adult male. Locality. (United States): Colorado, Park County, Snowy Range, near Mount Lincoln, Montgomery. 27 July 1871. Collector. J. A. Allen, Rocky Mountain Expedition. Original number 945. Condition. Skin complete. Type Series. Large paratype series; MCZ 209, 243— 263, skulls; B41 and B42, each represented by skin and skull; MCZ 2673-2703, skins. Comments. O. p. saxatilis was retained as a valid subspecies by Hall (1981: 291) and Smith and Wes- ton (1990: 1). Family LEPORIDAE Fischer de Waldheim, 1817 Genus LEPUS Linnaeus, 1758 Lepus (Macrotolagus) alleni palitans Bangs, 1900a Proc. New England Zool. Club, 1: 85, 23 February. Holotype. B9096. Skin and skull. Adult female. Locality. Mexico: Sinaloa, Aguacaliente (about 40 miles [64.4 km] southeast of Mazatlan). 7 August 1897. 133 Collector. P. O. Simons. Original number 157. Condition. Skin and skull complete. Mandible dis- articulated; left ramus chipped; two holes in right ramus. Type Series. 1 paratype; B9097, skin, adult male. Comments. Retained as a valid subspecies by Hall (1981: 332) and Best and Henry (1993: 1). Lepus americanus struthopus Bangs, 1898e Proc. Biol. Soc. Washington, 12: 81, 24 March. Holotype. B2025. Skin and skull. Adult female. Locality. (Canada): Nova Scotia, Digby. 4 August 1894. Collector. O. Bangs. Original number 9. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 318). Lepus arcticus bangsii Rhoads, 1896a Amer. Nat., 30: 236, 6 March. Holotype. B3752. Skin and skull. Adult female. Locality. (Canada): Newfoundland, Codroy. 3 Au- gust 1895. Collector. E. Doane. Original number 1. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 320). Lepus bairdii Hayden, 1869 Amer. Nat., 3: 115, May. =Lepus americanus bairdii Hayden, 1869. See J. A. Allen (1875: 431). Syntype. MCZ 5501. Skin and skull. Adult. Locality. (United States): Wyoming, (Fremont County), summit of Wind River Mountains, near Fremont Peak. 4 June 1860. Collector. F. V. Hayden. Original number 90. For- merly USNM 4264. Condition. Skin and skull complete. Type Series. 2 other adult syntypes; USNM 4262/ 38001, skin and skull, male: USNM 4263/4273, skin and skull; see Poole and Schantz (1942: 210). Comments. No type is designated in the original description, which is based on 6 individuals col- lected by Hayden—3 adults and 3 juveniles, origi- nally deposited in the USNM (Poole and Schantz 1942: 210). 134 Lepus bairdi cascadensis Nelson, 1907 Proc. Biol. Soc. Washington, 20: 87, 11 December. =Lepus americanus cascadensis Nelson, 1907. See Racey and Cowan (1936: H1S8). Holotype. B1886. Skin and skull. Adult male. Locality. Canada: British Columbia, near Hope, Roab’s Ranch. 12 June 1874. Collector. W. C. Colt. Original number 476. Condition. Skin complete. Skull slightly damaged (supraoccipital and occipital chipped). Type Series. Holotype only. Comments. L. a. cascadensis was retained as a valid subspecies by Hall (1981: 315). Genus SYLVILAGUS Gray, 1867 Lepus (Tapeti) incitatus Bangs, 1901b Amer. Nat., 35: 633, 22 August. = Sylvilagus brasiliensis incitatus (Bangs, 1901). See Hershkovitz (1950: 352). Holotype. B8441. Skin and skull. Adult female. Locality. Panama: (Panama), Bay of Panama, San Miguel Island. 30 April 1900. Collector. W. W. Brown, Jr. Original number 127 Condition. Skin and skull complete. Type Series. Holotype only. Comments. S. b. incitatus was retained as a valid subspecies by Hall (1981: 296). Lepus paludicola Miller and Bangs, 1894 Proc. Biol. Soc. Washington, 9: 105, 9 June. = Sylvilagus palustris paludicola (Miller and Bangs, 1894). See Nelson (1909: 269). Holotype. B1451. Skin and skull. Adult female. Locality. (United States): Florida, (Citrus County), Fort Island, near Crystal River. 25 January 1894. Collector. F. L. Small. Original number 1150. Condition. Skin and skull complete. Type Series. 3 paratypes; B1452, skin and skull, adult male; B1453, skin and skull, adult male; B1454, skin and skull, adult female. Comments. S. p. paludicola was retained as a valid subspecies by Hall (1981: 299) and Chapman and Willner (1981: 1). Lepus sylvaticus alacer Bangs, 1896h Proc. Biol. Soc. Washington, 10: 136, 28 December. = Sylvilagus floridanus alacer (Bangs, 1896). See Lyon (1904: 336). Holotype. B5480. Skin and skull. Adult female. Locality. (United States): Indian Territory Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 (=Oklahoma), (Adair County), Stilwell. 14 August 1896. Collector. T. Surber. Original number 65. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; B1677, skin and skull, B1678, skin and skull, adult male; B5481, skin and skull, adult female. Comments. S. f. alacer was retained as a valid sub- species by Hall (1981: 301) and Chapman et al. (1980: 1). Lepus sylvaticus transitionalis Bangs, 1895a Proc. Boston Soc. Nat. Hist., 26: 405, 31 January. = Sylvilagus transitionalis (Bangs, 1895). See Nelson (1909: 195). Holotype. B2407. Skin and skull. Adult female. Locality. (United States): Connecticut, New Lon- don County, Liberty Hill. 6 November 1894. Collector. O. Bangs. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a series of paratypes in the MCZ. Comments. S. transitionalis was considered a valid species by Hoffman (1993: 827) and Nowak (1999: LW) Order SCANDENTIA Wagner, 1855 Family TUPAIIDAE Gray, 1825 Genus TUPAIA Raffles, 1822 Tana tana griswoldi Coolidge, 1938 Proc. New England Zool. Club, 17: 45, 6 May. =Tupaia tana paitana (Lyon, 1913). See Medway (1965: 76). Holotype. MCZ 36416. Skin, skull, and baculum. Subadult male. Locality. (Malaysia): British North Borneo (=Sa- bah), Mount Kinabalu, Kenokok River, Kiau (=Kampong Kiau), 3,300 ft (1,007 m). 9 August 1937. Collector. J. A. Griswold, Jr., Asiatic Primate Ex- pedition. Original number 684. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden Order PRIMATES Linnaeus, 1758 Family INDRIDAE Burnett, 1828 Genus PROPITHECUS Bennett, 1832 Propithecus perrieri Lavauden, 1931 Compt. Rend. Acad. Sci. Paris, 193: 77, 6 July. =Propithecus diadema perrieri Lavauden, 1931. See W. C. O. Hill (1953: 568). Holotype. MCZ 44857. Skin, skull, and postcranial skeleton. Adult male. Locality. Malagasy Republic (Madagascar): (Toli- ary), [fotaka, vicinity of Ambovombe. Probably 1928. Collector. M. Perrier. Original number 1. Grandi- dier Collection. Condition. Skin and skull complete. Postcranial skeleton partial (right humerus broken, foot bones left in skin). Type Series. 3 paratypes; MCZ 44858, skin, skull, and postcranial skeleton, adult female; MCZ 44859, skin and skull, juvenile; MCZ 46001, skin and leg bones, adult. Comments. P. d. perrieri was retained as a valid subspecies by Tattersall (1982: 103). Family DAUBENTONIIDAE Gray, 1863 Genus DAUBENTONIA E. Geoffroy, 1795 Cheiromys madagascariensis laniger G. Grandidier, 1930d Bull. Acad. Malgache, n. ser., 11: 106 (for 1928). = Daubentonia madagascariensis (Gmelin, 1788). See G. M. Allen (1939: 134). Holotype. MCZ 45947. Skull and postcranial skele- ton. Adult male. Locality. Madagascar: Forest of the East. Collector. Received by G. Grandidier from the Académie Malgache, 1927-28. Condition. Skull complete. Postcranial skeleton in- complete (hands and feet missing, presumably in- tact within mounted skin). Type Series. Holotype only. Comments. The mounted skin of the holotype is in the collection of the Académie Malgache (Anta- nanarivo Museum). Grandidier named laniger as a new subspecies on account of its woolly pelage, but it is probably just a molting individual (Schwarz 1931: 428). Si) Family GALAGONIDAE Gray, 1825 Genus GALAGOIDES A. Smith, 1833 Galago demidovii orinus Lawrence and Washburn, 1936 Occ. Pap. Boston Soc. Nat. Hist., 8: 259, 8 January. = Galagoides orinus (Lawrence and Washburn, 1936). See Honess (1996: 58). Holotype. MCZ 22453. Skin and skull. Adult male. Locality. Tanganyika Territory (=Tanzania): Mor- ogoro, Uluguru Mountains, Bagilo, 5,000 ft (1,525 m). 17 September 1926. Collector. A. Loveridge. Condition. Skin and skull complete. Type Series. Holotype only. Family CEBIDAE Bonaparte, 1831 Genus ALOUATTA Lacépéde, 1799 Alouatta palliata luctuosa Lawrence, 1933b Bull. Mus. Comp. Zool., 75: 337, November. =Alouatta pigra Lawrence, 1933. See Smith (1970: 363). Holotype. MCZ 24059. Skin and skull. Adult male. Locality. British Honduras (=Belize): Cayo Dis- trict, Mount Cow. 12 April 1928. Collector. O. L. Austin, Jr. Original number 723. Condition. Skin and skull complete. Type Series. Holotype only. Alouatta palliata trabeata Lawrence, 1933b Bull. Mus. Comp. Zool., 75: 328, November. Holotype. MCZ 29545. Skin and skull. Adult male. Locality. Panama: Herrera Province, Capina. March 1933. Collector. T. Barbour. Original number 4. Condition. Skin and skull complete. Type Series. Lawrence mentions examining 19 specimens of trabeata and lists them by locality (1933: 330). 9 of these are in the MCZ; other than the holotype, they are MCZ 27784, 27785, 28735, 28736, 29543, 29544, 29546, and 29548. 1 paraty- pe, MCZ 29547, has been exchanged to the Museu Palista, Sao Paolo, Brazil. Comments. Retained as a valid subspecies by Hall (1981: 263). 136 Genus CEBUS Erxleben, 1777 Cebus curtus Bangs, 1905b Bull. Mus. Comp. Zool., 46: 91, June. = Cebus capucinus curtus Bangs, 1905. See Cabrera (1917: 240). Holotype. MCZ 10824. Skin and skull. Adult male. Locality. Colombia: (Cauca), Gorgona Island. 2 July 1904. Collector. W. W. Brown, Jr. Original number 27. Condition. Skin and skull complete. Type Series. 1 paratype; MCZ 10825, skin and skull, adult female. Comments. C. c. curtus was retained as a valid sub- species by Cabrera (1958: 169) and Napier (1976: 36). Family HYLOBATIDAE Gray, 1870 Genus HYLOBATES lIlliger, 1811 Hylobates lar carpenteri Groves, 1968 Proc. Biol. Soc. Washington, 81: 625, 30 December. Holotype. MCZ 41430. Skin and_ skeleton. Adult male. Locality. Thailand: Chiengmai District, Mount Angka (=Doi Inthanon), 3,400 ft (1,037 m). 14 March 1937. Collector. H. J. Coolidge, Jr., Asiatic Primate Ex- pedition. Original numbers 21 and 155. Condition. Skin, skull, and skeleton complete. Type Series. Groves examined a series of 144 skins, skulls, and skeletons of this subspecies that were collected by the Harvard Asiatic Primate Expedi- tion in 1937 from Chiengmai District, Thailand, “mostly in the Museum of Comparative Zoology, Harvard, but a few of the osteological specimens in the Anthropology Department, University of California at Berkeley.” He examined also 18 spec- imens in the USNM and 1 in the AMNH, all rep- resented by skin and skull. Comments. Retained as a valid subspecies by Jen- kins (1990: 15) and Geissman (1995: 474). Family HOMINIDAE Gray, 1825 Genus GORILLA |. Geoffroy, 1853 Troglodytes gorilla Savage and Wyman, 1847 Boston J. Nat. Hist., 5: 417, December. =Gorilla gorilla gorilla (Savage and Wyman, 1847). See Rothschild (1923: 176). Lectotype. MCZ 9587. Skull and postcranial skeleton. Adult male. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Locality. Gabon: Gabon Estuary, Mpongwe coun- try. 1847. Collector. T. S. Savage. Original number 28 (of J. Wyman). Condition. Postcranial skeleton partial (includes pelvis, sacrum, scapulae, humeri, radii, left ulna, femora, left tibia, and 7 vertebrae: 2 cervical, 3 dor- sal, 2 lumbar). Head of right humerus and left fe- mur bisected. Type Series. The original description mentions “four skulls, two males and two females, one of each in a perfect condition, and all of them adult; a male and female pelvis, the long bones of the upper and lower extremities, and a few vertebrae and ribs.” However, measurements and illustrations of only two of these specimens, an adult male and female, each represented by a skull and partial postcranial skeleton, are provided. These two, pre- sumably the specimens originally described as be- ing “in a perfect condition,” are the only specimens Br the original four to be noted in Wyman’s per- sonal notebook of osteology (unpublished, now in the library of the Boston Museum of Science) and the only original specimens of which there is any record at Fil These two specimens were trans- ferred from the Boston Society of Natural History to the MCZ in 1915-16. They bear MCZ numbers 9587 (male) and 9311 (female). The skull of the female was sawed in half (hemisected), probably by Wyman. The adult male specimen, MCZ 9587, possesses an intact skull. MCZ 9587 is the only in- tact specimen of the syntype series known to exist, and its measurements and an illustration of its skull are included in the original publication. Addition- ally, it is the only syntype whose measurements are included in Coolidge’ s revision of the genus Gorilla (1929: 325), although it is erroneously listed in that work as MCZ 9586. For these reasons, MCZ 9587 is hereby designated as lectotype of Troglodytes go- rilla Savage and Wyman, 1847; this should ensure taxonomic consistency between past and future treatments of this name. Comments. Type species of the genus Gorilla I. Geoffroy, 1853. The original description is occa- sionally attributed in error to Wyman (1847). G. gorilla was considered a valid species by Groves (1993: 276) and Nowak (1999: 618). Order LIPOTYPHLA Haeckel, 1866 Family NESOPHONTIDAE Anthony, 1916 Genus NESOPHONTES Anthony, 1916 Nesophontes micrus G. M. Allen, 1917a Bull. Mus. Comp. Zool., 61: 5, January. Holotype. VP 9600. Right ramus. Locality. Cuba: Matanzas Province, cave in Sierra de Hato Neuvo. Collector. C. de la Torre. TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden 1 Condition. Condition as originally described— “posterior half of the right ramus, containing a part of pm, m,, ms, and the roots of m;.” Type Series. Holotype only. Comments. Considered a valid species by Hutterer (1993: 70) and Nowak (1999: 201); almost certainly extinct. This specimen is stored in the Vertebrate Paleontology Department of the MCZ. Family SOLENODONTIDAE Gill, 1872 Genus SOLENODON Brandt, 1833 Antillogale marcanoi Patterson, 1962 Breviora, Mus. Comp. Zool., 165: 3, 22 August. = Solenodon marcanoi (Patterson, 1962). See Varona (1974: 8). Holotype. VP 7261. Right lower ramus. Locality. Dominican Republic: San Rafael Prov- ince (=Elias Pina), Hondo Valle Municipality, un- named cave 2 km southeast of Rancho La Guardia. Summer 1958. Collector. C. E. Ray and A. S. Rand. Condition. Condition as originally described—“in- complete right ramus of mandible with P;-M, and alveoli of other teeth.” Type Series. 5 paratypes; 7262, left ramus, 7263, right humerus, 7264, left humerus, 7265, right ulna, 7266, left ramus, juvenile; all partial. Comments. A. marcanoi is the type species of the genus Antillogale Patterson, 1962. Antillogale was first synonymized with Solenodon by Van Valen (1967: 255). Considered a valid species by Hutterer (1993: 69) and Nowak (1999: 199). S. marcanoi is probably extinct. This specimen is stored in the Vertebrate Paleontology Department of the MCZ. Solenodon poeyanus Barbour, 1944 Proc. New England Zool. Club, 23: 6, March 7. = Solenodon cubanus poeyanus Barbour, 1944. See Aguayo (1950: 131). Holotype. MCZ 6597. Mounted skin and skull. Locality. Cuba: Oriente (=Holguin), near Nipi Bay (=Nipe Bay). Collector. Bought by A. Agassiz from H. A. Ward, 1891. Condition. Skin complete. Skull partial (basioccip- ital plate and tympanic bullae missing, parietals damaged). Mandible disarticulated. Type Series. In the original description, Barbour includes a photograph of a “living example of So- lenodon poeyanus now in Zoological Garden, Ha- vana, Cuba, from vicinity of Baracoa.” Comments. The number of this specimen is 6597, not 6957 as stated in the original description. Sh @ oN) ~l poeyanus was retained as a valid subspecies by Hall (1981: 22). Family SORICIDAE Fischer von Waldheim, 1817 Genus BLARINA Gray, 1838 Blarina brevicauda aloga Bangs, 1902a Proc. New England Zool. Club, 3: 76, 31 March. Holotype. B9727. Skin and skull. Adult male. Locality. (United States): Massachusetts, Dukes County, Martha’s Vineyard, West Tisbury. 25 June 1899. Collector. O. Bangs. Original number 2. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 9 paratypes, B9725, B9726, B9728— B9734, all represented by skin and skull, 3 females and 6 females. Comments. Retained as a valid subspecies by Hall (1981: 54) and George et al. (1986: 1). Blarina brevicauda compacta Bangs, 1902a Proc. New England Zool. Club, 3: 77, 31 March. Holotype. B97O5. Skin and skull. Adult male. Locality. (United States): Massachusetts, (Nan- tucket County), Nantucket (Island). 10 July 1899. Collector. O. Bangs. Original number 3. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 6 paratypes; B9701—B9704, B97O06, B9708; all represented by skin and skull, 4 females and 2 males. Comments. Retained as a valid subspecies by Hall (1981: 56) and George et al. (1986: 1). Genus CROCIDURA Wagner, 1832 Crocidura bicolor tephragaster Setzer, 1956 Proc. U.S. Nat. Mus., 106: 458, 28 November. =Crocidura fuscomurina (Heuglin, 1865). See Hutterer (1983: 223). Holotype. MCZ 44773. Skin and skull. Adult male. Locality. Anglo-Egyptian Sudan (=Sudan): (East- ern) Equatoria, Torit. 25 April 1950. Collector. J. S. Owen. Original number 1158. Condition. Skin and skull complete. Type Series. Setzer mentions that he examined 18 specimens of tephragaster, 8 of which are in the MCZ. 138 Crocidura hildegardeae phaios Setzer, 1956 ErOcw Wr SeaNat, November. Mus., 106: 460, 28 Holotype. MCZ 45855. Skin and skull. Adult female. Locality. Anglo-Egyptian Sudan (=Sudan): Equa- toria, Imatong Mts, Gilo, 6,500 ft (1,983 m). 12 June 1950. Collector. J. S. Owen. Original number 1266. Condition. Skin and skull ‘complete. Mandible dis- articulated. Type Series. 3 paratypes; 1 from the MCZ; MCZ 45856, skin and skull, male. Comments. Retained as a valid subspecies, C. gra- cilipes phaios, by Heim de Balsac and Meester (1977: 16): included in Crocidura hilde gardeae by Hutterer (1993: 87). Genus CRYPTOTIS Pomel, 1848 Cryptotis avia G. M. Allen, 1923a Proc. New England Zool. Club, 8: 37, 12 February. =Cryptotis thomasi (Merriam, 1897). See Woodman (1996: 414). Holotype. MCZ 20091. Skin and skull. Adult. Locality. Colombia: (probably Cundinamarca), E] Verjon (see coments). October 1922. Collector. N. Maria. Condition. Skin complete. Skull partial (most of skull from frontals to occiput missing). Left ramus of mandible missing. Type Series. Holotype only. Comments. Considered a valid species by Hutterer (1993: 108) and Nowak (1999: 209) but synony- mized as noted above. The locality “El Verjon” is not shown on any maps av ailable to us but is pos- sibly equivalent to “Paramo Cruz Verde” (Paynter 1997: 463). Genus MYOSOREX Gray, 1838 Crocidura maurisca geata G. M. Allen and Loveridge, 1927 Proc. Boston Soc. Nat. Hist., 3 December. =Myosorex geata (G. M. Allen and Loveridge, 1927). See Heim de Balsac (1967: 610). Holotype. MCZ 22447. Skin and skull. Adult male. Locality. Tanganyika Territory (=Tanzania): Mor- ogoro, Uluguru Mountains, Nyingwa, 7,500 ft (2,288 m). 19 October 1926. Collector. A. Loveridge. Condition. Skin and skull complete. Mandible dis- articulated. 8: 417, 23 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Type Series. 1 paratype; MCZ 22448, skin and skull, adult female. Comments. M. geata was considered a valid species by Hutterer (1993: 99) and Nowak (1999: 217). Genus SOREX Linnaeus, 1758 Neosorex palustris acadicus G. M. Allen, 1915a Proc. Biol. Soc. Washington, 28: 15, 12 February. Name preoccupied by Sorex acadicus Gilpin, 1867. Sorex sere gloveralleni Jackson, 1926 J. Mammal., 7: 57, 15 February. (once ee name for Neosorex palustris acadicus G. M. Allen, 1915) Holotype. B2046. Skin and skull. Adult female. Locality. (Canada): Nova Scotia, Digby. 26 July 1894. Collector. O. Bangs. Original number 3. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; USNM in 1922. Comments. Retained as a valid subspecies by Hall (1981: 41) and Beneski and Stinson (1987: 1). B2053, exchanged to the Sorex araneus ultimus G. M. Allen, 1914b Proc. New England Zool. Club, 5: 51, 9 April. = Sorex tundrensis Merriam, 1900. See Hutterer (1993: 121). Holotype. MCZ 15000. Skin and skull. Adult male. Locality. (Russian Federation): northeastern Sibe- ria, Nijni Kolymsk (=Nizhnekolymsk), near mouth of Kolyma River. 6 November 1911. Collector. J. Koren. Original number 136. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a series of paratypes, most of which are housed in the MCZ. Sorex macropygmaeus koreni G. M. Allen, 1914b Proc. New England Zool. Club, April. = Sorex caecutiens koreni G. M. Allen, 1914. See Yudin (1989: 281). Holotype. MCZ 15085. Skin and skull. Adult female. Locality. (Russian Federation): northeastern Sibe- ria, Nijni Kolymsk (=Nizhnekolymsk), near mouth of Kolyma River. 19 October 1911. Collector. J. Koren! Original number 50. De OOo TYPE SPECIMENS OF RECENT MAMMALS ° Helgen and McFadden Type Series. 5 paratypes; MCZ 15003-15007, all represented by skin and skull; 4 males, 1 female. MCZ 15004, a male, was exchanged to the FMNH in 1931. Condition. Skin and skull complete. Mandible dis- articulated. Sorex macrurus Batchelder, 1896 Proc. Biol. Soc. Washington, 10: 133, § December. Name preoccupied by Sorex macrourus Lehmann, 1822. Sorex dispar Batchelder, 1911 Proc. Biol. Soc. Washington, 24: 97, 15 May. (Replacement name for Sorex macrurus Batchelder, 1896) Holotype. MCZ 41744. Skin and skull. Adult male. Locality. (United States): New York, Essex County, Keene Heights, Beede’s (see comments). 9 Sep- tember 1895. Collector. C. F. Batchelder. Original number 1384. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; MCZ 41745, skin and skull, adult male. Comments. S. dispar was considered a valid species by Hutterer (1993: 114) and Nowak (1999: 205). The type locality was redescribed by Martin (1966: 131) as “0.6 mile south and 0.5 mile east of Saint Huberts, Essex County, New York, lat. 44°09’, long. 73°46'.” Sorex personatus miscix Bangs, 1899d Proc. New England Zool. Club, 1: 15, 28 February. = Sorex cinereus miscix Bangs, 1899. See Jackson (1925: 56). Holotype. B8651. Skin and skull. Adult male. Locality. (Canada): Labrador, Black Bay. 10 Octo- ber 1898. Collector. E. Doane. Original number 1. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Bangs based his description on 39 specimens; corresponding to B7931—B7950 and B8651—B8669; all represented by skin and skull. Comments. S. c. miscix was retained as a valid sub- species by Hall (1981: 29). Sorex sanguinidens G. M. Allen, 1914b Proc. New England Zool. Club, 5: 54, 9 April. = Sorex daphaenodon sanguinidens G. M. Allen, 1914. See Yudin (1989: 198). Holotype. MCZ 15012. Skin and skull. Adult female. Locality. (Russian Federation): northeastern Sibe- 139 ria, Nijni Kolymsk (=Nizhnekolymsk), near mouth Kolyma River. 11 December 1911. Collector. J. Koren. Original number 221. Condition. Tail separate from skin. Skull complete. Mandible disarticulated. Type Series. There is a large series of paratypes, most of which are still in the MCZ. Sorex virG. M. Allen, 1914b Proc. New England Zool. Club, 5: 52, 9 April. = Sorex roboratus Hollister, 1913. See Hoffman (1985: 17). Holotype. MCZ 15068. Skin and skull. Adult female. Locality. (Russian Federation): northeastern Sibe- ria, Nijni Kolymsk (=Nizhnekolymsk), near mouth of Kolyma River. 19 December 1911. Collector. J. Koren. Original number 230. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a large series of paratypes, most of which are still in the MCZ. Genus SUNCUS Ehrenberg, 1832 Suncus ater Medway, 1965 Mammals of Borneo, J. Malay. Branch R. Asiat. Soc., 36: 38, December. Holotype. MCZ 36574. Skin, skull, and postcranial skeleton. Adult female. Locality. Malaysia: northern Borneo, Sabah, Mount Kinabalu, Lumu Lumu, 5,500 ft (1,678 m). 7 July 1937. Collector. J. A. Griswold, Jr. Original number 462. Condition. Skin complete. Skull partial (tympanic bullae missing), and mandible disarticulated. Post- cranial skeleton complete. Type Series. Holotype only. Comments. Considered a valid species by Hutterer (1993: 101) and Nowak (1999: 223). Suncus varilla minor G. M. Allen and Loveridge, 1933 Bull. Mus. Comp. Zool., 75: 57, February. Holotype. MCZ 26754. Skin and skull. Adult female. Locality. Tanganyika Territory (=Tanzania): Urun- gu, Kitungulu, 4,500 ft (1,373 m). 14 May 1930. Collector. A. Loveridge. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. Retained as a valid subspecies by Heim de Balsac and Meester (1977: 6). 140 Family TALPIDAE Fischer de Waldheim, Ws y Genus NEUROTRICHUS Gunther, 1880 Neurotrichus gibbsi hyacinthinus Bangs, 1897d Amer. Nat., 31: 240, 1 March. Holotype. B1240. Skin and skull. Adult female. Locality. U.S.A. (United States): California, Marin County, Nicasio. 10 March 1894. Collector. C. A. Allen. Original number 694. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; B1241, skin and _ skull, adult male. Comments. Retained as a valid subspecies by Hall (1981: 67) and Carraway and Verts (1991: 1). Genus SCALOPUS Desmarest, 1804 Scalops anastasae Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 212, 15 March. = Scalopus aquaticus anastasae (Bangs, 1898). See Jackson (1915: 39). Holotype. B7192. Skin and skull. Adult male. Locality. U.S.A. (United States): Florida, St. Johns County, Anastasia Island, Point Romo. 16 February 1897. Collector. O. Bangs. Original number 10. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 4 paratypes; B7193—B7196; all repre- sented by skin and skull, 2 females and 2 males. Comments. S. a. anastasae was retained as a valid subspecies by Hall (1981: 72). Scalops texanus aereus Bangs, 1896h Proc. Biol. Soc. Washington, 10: 138, 28 December. = Scalopus aquaticus aereus (Bangs, 1896). See Miller (1912: 8). Holotype. B5475. Skin and skull. Adult female. Locality. (United States): Indian Territory (=OkKahoma) (Adair County), Stilwell. 13 August 1896. Collector. T. Surber. Original number 64. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. S. a. aereus was retained as a valid sub- species by Hall (1981: 72). Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Genus SCAPANUS Pomel, 1848 Scapanus Californicus minusculus Bangs, 1899} Proc. New England Zool. Club, 1: 70, 31 July. = Scapanus latimanus minusculus Bangs, 1899. See Grinnell and Swarth (1912: 133): Holotype. B9189. Skin and skull. Adult female. Locality. (United States): California, El Dorado County, Fyffe. 10 June 1897. Collector. W. W. Price and E. M. Nutting. Original number 15. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. S. l. minusculus was retained as a valid subspecies by Hall (1981: 70). Order CHIROPTERA Blumenbach, 1779 Family PTEROPODIDAE Gray, 1821 Genus AETHALOPS Thomas, 1923 Aethalops aequalis G. M. Allen, 1938b J. Mammal., 19: 497, 14 November. =Aethalops alecto aequalis G. M. Allen, 1938. See Hill (1961: 639). Holotype. MCZ 36582. Skin, skull, and postcranial skeleton. Adult female. Locality. (Malaysia): British North Borneo (=Sa- bah), Mount Kinabalu, Luma Luma, 5,500 ft (1,678 m). 12 July 1937. Collector. J. A. Griswold, Jr., Asiatic Primate Ex- pedition. Original number 510. Condition. Skin, skull, and posteranial skeleton complete. Type Series. 3 paratypes; MCZ 36583, skin, skull and skeleton, female; MCZ 36584, skin and skull, female; MCZ 36586, skin and skull, female. Comments. A. a. aequalis was retained as a valid subspecies by Koopman (1994: 35). Genus EONYCTERIS Dobson, 1873 Eonycteris spelaea glandifera Lawrence, 1939 In Barbour, Lawrence, and Peters, Bull. Mus. Comp. Zool., 86: 38, November. Holotype. MCZ 35159. Skin and skull. Adult male. Locality. Philippines: Luzon, (Nueva Ecija), Rizal, Montalban caves near Manila. 27 February 1937. Collector. B. Lawrence. Original number 253. Condition. Skin and skull complete. Type Series. In the original description, Lawrence TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden implies the existence of several specimens in ad- dition to the holotype; she collected 18 specimens of glandifera during her 1936-37 expedition to the Philippines, all of which are in the MCZ. Comments. Retained as a valid subspecies by Ma- haradatunkamsi and Kitchener (1997: 59). Genus HAPLONYCTERIS Lawrence, 1939 Haplonycteris fischeri Lawrence, 1939 In Barbour, Lawrence, and Peters, Bull. Mus. Comp. Zool., 86: 33, November. Holotype. MCZ 35258. Skin and skull. Adult male. Locality. Philippines: Mindoro (Oriental), Mount Halcon, Bignay. 26 April 1937. Collector. F. S. Rivera. Original number BL 502. Condition. Skin and skull complete. Type Series. Holotype only. Comments. Type species of the genus Haplonyc- teris Lawrence, 1939. Considered a valid species by Koopman (1993: 142) and Nowak (1999: 292). Genus PTEROPUS Erxleben, 1777 Pteropus anetianus aorensis Lawrence, 1945 Proc. New England Zool. Club, 23: 66, 26 March. Holotype. MCZ 42183. Skin and skull. Adult male. Locality. (Vanuatu): New Hebrides, off southwest corner of Espiritu Santo Island, Aore Island. 8 April 1944. Collector. O. L. Austin, Jr. Original number 5. Condition. Skin complete. Skull partial (basioccip- ital missing), right ramus of mandible broken. Type Series. 1 paratype; MCZ 42182, skin and skull, adult male. Comments. Retained as a valid subspecies by Koopman (1994: 25) and Flannery (1995b). Pteropus ariel G. M. Allen, 1908 Bull. Mus. Comp. Zool., 52: 28, July. =Pteropus giganteus ariel G. M. Allen, 1908. See Hill (1958: 5). Holotype. MCZ 10565. Skin and skull. Adult male. Locality. Maldive Islands: Male Atoll. 24 Decem- ber 1901. Collector. H. B. Bigelow, A. Agassiz Expedition. Condition. Right wing of skin damaged. Skull in- tact, with 2 small holes in braincase. Type Series. 1 paratype; MCZ 10566, skin and skull, juvenile female. Comments. P. g. ariel was retained as a valid sub- species by Koopman (1994: 26). 141 Pteropus austini Lawrence, 1945 Proc. New England Zool. Club, 23: 59, 26 March. =Pteropus woodfordi Thomas, 1888. See Sanborn and Beecher (1947: 389). Holotype. MCZ 42166. Skin and skull. Subadult fe- male. Locality. Solomon Islands: Florida Island (Nggela Group). 20 February 1944. Collector. O. L. Austin, Jr. Original number 2. Condition. Skin and skull complete. Type Series. 1 paratype; MCZ 42167, skin and skull, subadult male. Comments. Lawrence referred to the holotype as an adult in the original description, but Sanborn and Beecher (1947: 389) recognized it as a sub- adult. Pteropus rayneri monoensis Lawrence, 1945 Proc. New England Zool. Club., 23: 63, 26 March. Holotype. MCZ 42191. Skin and skull. Adult male. Locality. Solomon Islands: Treasury (Mono) Island. 11 October 1944. Collector. O. L. Austin, Jr. Original number 27. Condition. Skin and skull complete. Type Series. 2 paratypes; MCZ 42192, skin and skull, adult male; MCZ 42193, skin and skull, adult male. Comments. Retained as a valid subspecies by Koopman (1994: 24) and Flannery (1995b: 285). Genus ROUSETTUS Gray, 1821 Rousettus madagascariensis G. Grandidier, 1930b Bull. Acad. Malgache, n. ser., 11: 91 (for 1928). Holotype. MCZ 45432. Alcoholic and skull. Adult male. Locality. Malagasy Republic (Madagascar): (Anta- nanarivo), between Tananarive (=Antananarivo) and Andevoranto, Grand forest de Est, near Be- forona. Collector. Received by G. Grandidier from the Académie Malgache, 1917. Condition. Alcoholic, skull complete. Type Series. Holotype only. Comments. Considered a valid species by Koop- man (1993: 153) and Nowak (1999: 261). In his review of known material of R. madagascariensis, Bergmans (1977: 67) commented in error that the holotype was in the Académie Malgache, Antana- narivo, Madagascar. 142 Family EMBALLONURIDAE Gervais, 1855 Genus PEROPTERYX Peters, 1867 Peropteryx canina phaea G. M. Allen, 1911a Bull. Mus. Comp. Zool., 54: 222, July. =Peropteryx macrotis phaea G. M. Allen, 1911. See Sanborn (1937: 342). Holotype. MCZ 8101. Skin and skull. Adult female. Locality. Grenada: Point Saline(s). 29 August 1910. Collector. G. M. Allen. Original number 15. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. There is a series of paratypes in the MCZ. Comments. P. m. phaea was retained as a valid sub- species by Hall (1981: 82) and Koopman (1994: 47). Genus RHYNCHONYCTERIS Peters, 1867 Rhynchiscus naso priscus G. M. Allen, 1914c Proc. Biol. Soc. Washington, 27: 109, 10 July. =Rhynchonycteris naso (Wied-Neuwied, 1820). See Sanborn (1937: 326). Holotype. MCZ 13208. Skin and skull. Adult. Locality. Mexico: Quintana Roo, Xcopen. 18 Feb- ruary 1912. Collector. J. L. Peters. Original number 13. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; MCZ 13209, skin and skull, adult male; MCZ 14637, alcoholic, adult male: MCZ 14638, alcoholic, adult female. Family NYCTERIDAE Van der Hoeven, 1855 Genus NYCTERIS G. Cuvier and E. Geoffroy, 1795 Nycteris madagascariensis G. Grandidier, 1937 Bull. Mus. Nat. Hist. Paris, 9: 353, 25 November. Holotype. MCZ 45433. Body in alcohol, skull extract- ed. Locality. Madagascar: (Antsiranana), Diego-Suarez (=Antsiranana), Valley of the Rodo, north of Pir kana near the Ankarana, 12°5’—13°0'S, 49°5’E. June 1910. Collector. Grandidier collection. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Condition. Alcoholic, skull complete. Type Series. 1 paratype, MCZ 45434, in alcohol, skull extracted, female. Comments. N. madagascariensis was included in N. macrotis by Koopman (1993: 162) but retained as a valid species by Peterson et al. (1995: 63). Nycteris nana tristis G. M. Allen and Lawrence, 1936 Bull. Mus. Comp. Zool., 79: 47, January. =WNycteris nana (Andersen, 1912). See Hayman and Hill (1971: 19). Holotype. MCZ 31156. Skin and skull. Adult female. Locality. Kenya: (W. Nyanza), Kakamega District, Kaimosi. 13 February 1934. Collector. A. Loveridge. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Family RHINOLOPHIDAE Gray, 1825 Genus HIPPOSIDEROS Gray, 1831 Hipposideros curtus G. M. Allen, 1921 Rev. Zool. Africaine, 9: 194, December. Holotype. MCZ 19305. Body in alcohol. Female. Locality. Cameroons (=Cameroon): (Littoral), Sak- bayeme. 1920. Collector. G. Schwab. Condition. Alcoholic. cleaned (still in alcohol). Type Series. Holotype only. Comments. H. curtus was considered a valid spe- cies by Koopman (1993: 172) and Nowak (1999: 333). extracted but not Skull Hipposideros erigens Lawrence, 1939 In Barbour, Lawrence, and Peters, Bull. Mus. Comp. Zool., 86: 56, November. = Hipposideros bicolor erigens Lawrence, 1939. See Hill (1963: 28). Holotype. MCZ 35197. Skin and skull. Adult male. Locality. Philippines: Mindoro, (Oriental) Tabucala cave near Calapan, northern base of Mount Hal- con. 7 March 1937. Collector. B. Lawrence. Original number 307. Condition. Skin complete. Skull partial (occiput missing), and mandible disarticulated. Type Series. 3 paratypes; MCZ 35195, skin and skull, adult female; MCZ 35196, skin and skull, adult male; MCZ 35198, skin and skull, adult fe- male. Comments. H. b. erigens was retained as a valid subspecies by Koopman (1994: 61). TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Hipposideros turpis Bangs, 1901a Amer. Nat., 35: 561, 31 July. = Hipposideros turpis turpis Bangs, 1901. See Hill (1963: 94). Holotype. MCZ 10003. Skin and skull. Adult female. Locality. (Japan): Ryukyu Islands, southern group of Liu Kiu Islands, Ishigaki Island. 10 May 1899. Collector. 1. Zensaku. Condition. Skin complete. Skull partial (occipital region missing). Type Series. 2 skull, adult female; adult male. Comments. H. turpis was considered a valid species by Koopman (1993: 175) and Nowak (1999: 334). Genus RHINOLOPHUS Lacépéde, 1799 Rhinolophus megaphyllus ignifer G. M. Allen, 1933 J. Mammal., 14: 149, 15 May. = Rhinolophus megaphyllus megaphyllus Gray, 1834. See Koopman (1984: 9). Holotype. MCZ 29078. Skin and skull. Adult male. Locality. Australia: Queensland, Cape York, Coen. 12 June 1932. Collector. P. J. Darlington, Jr., Expedition. Original number 185. Condition. Slani: and skull complete. Mandible dis- articulated. Type Series. 1 paratype; MCZ 29079, skull, adult male. paratypes; MCZ 10002, skin and MCZ 10004, skin and _ skull, Harvard Australian skin and Rhinolophus philippinensis alleni Lawrence, 1939 In Barbour, Lawrence, and Peters, Bull. Mus. Comp. Zool., 86: 46, November. Holotype. MCZ 35097. Skin and skull. Adult female. Locality. Philippines: Mindoro, (Oriental) Tabucala cave near Calapan, northern base of Mount Hal- con. 7 March 1937. Collector. B. Lawrence. Original number 302. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 paratypes; skull, adult female; MCZ adult female. Comments. Retained as a valid subspecies by Koopman (1994: 57). Genus TRIAENOPS Dobson, 1871 Triaenops aurita G. Grandidier, 1912 Bull. Mus. Hist. Nat. Paris, 18: 8, 25 January. Holotype. MCZ 45080. Muminy. Locality. Malagasy Republic (=Madagascar): (An- tsiranana), Diego-Suarez (=Antsiranana). 1910. MCZ 35099, 35098, skin and skin and _ skull, 143 Collector, Dr. Mazieres. Condition. Mummy (dried carcass), complete. Type Series. Holotype only. Comments. Traditionally included in Triaenops fur- culus, as in G. M. Allen (1939: 82) and Koopman (1993: 175), but retained as distinct by Peterson et al. (1995: 81) pending further material from the area of the type locality. Known only from the ho- lotype. Family MORMOOPIDAE de Saussure, 1860 Genus PTERONOTUS Gray, 1838 Chilonycteris parnellii pusillus G. M. Allen, 1917c Proc. Biol. Soc. Washington, 30: 168, 23 October. =Pteronotus parnellii pusillus (G. M. Allen, 1917). See Smith (1972: 67). Holotype. MCZ 16468. Skin and skull. Female. Locality. Dominican Republic: Santo Domingo, Arroyo Salado. 7 March 1916. Collector, J. L. Peters. Original number 227. Condition. Skin complete. Skull partial (left wall of braincase broken, left tympanic bulla missing). Type Series. 2 paratypes; MCZ 16599, female; MCZ 16600, female; both in alcohol. Comments. P. p. pusillus was retained as a valid subspecies by Hall (1981: 92) and Koopman (1994: 71). Chilonycteris torrei G. M. Allen, 1916a Proc. New England Zool. Club, 6: 4, 8 February. =Pteronotus quadridens quadridens (Gundlach, 1840). See Silva-Taboada (1976: 7). Holotype. MCZ 11672. Body in alcohol, skull extract- ed. Adult female. Locality. Cuba: (Guantanama), Baracoa, La Cueva de la Majana. 15 June 1915. Collector. V. J. R. Verrier. Presented to the MCZ by Carlos de la Torre. Condition. Alcoholic, skull complete. Type Series. 2 paratypes; MCZ 11670, male; MCZ 11671, female; both in alcohol. Family PHYLLOSTOMIDAE Gray, 1825 Genus AMETRIDA Gray, 1847 Ametrida minor H. Allen, 1894 Proc. Boston Soc. Nat. Hist., May. =Ametrida centurio Gray, 1847. See Peterson (1965: 5). Holotype. MCZ 11274. Body in alcohol, skull extract- ed. Adult male. 26: 240, 16 144 Locality. Suriname: Paramaribo. Collected some- time between 1832 and 1839. Collector. F. W. Cragin. Condition. Alcoholic, skull complete. Type Series. Holotype only. Comments. For comments on the type locality and date of acquisition, see G. M. Allen (1902a: 88). A. minor actually represents the male specimens of A. centurio. Genus ARTIBEUS Leach, 1821 Artibeus femurvillosum Bangs, 1899k Proc. New England Zool. Club, 1: 73, 24 November. =Artibeus lituratus palmarum J. A. Allen and Chapman, 1897. See Hershkovitz (1949: 445). Holotype. B8314. Skin and skull. Adult male. Locality. Colombia: (La Guajira), La Concepcion, 3,000 ft (915 m). 21 March 1899. Collector. W. W. Brown, Jr. Original number 31. Condition. Skin and skull complete. Type Series. Holotype only. Genus EROPHYLLA Miller, 1906 Erophylla sezekorni syops G. M. Allen, 1917c Proc. Biol. Soc. Washington, 30: 167, 23 October. Holotype. MCZ 13713. Body in alcohol, skull extract- ed. Adult male. Locality. Jamaica: (St. James), Montego Bay. 14 March 1912. Collector. J. A. Cushman. Condition. Alcoholic, skull complete. Type Series. 7 paratypes; MCZ 13709-13712, 13714—13716, all alcoholic. MCZ 13709 and 13712 were sent in exchange to the USNM. Comments. Retained as a valid subspecies by Hall (1981: 171) and Koopman (1994: 79). Genus GLOSSOPHAGA E. Geoffroy, 1818 Glossophaga longirostris Miller, 1898 Proc. Acad. Nat. Sci. Philadelphia, 1898, p- 330, 2 August. = Glossophaga longirostris longirostris Miller, 1898. See Miller (1913: 422). Holotype. B8046. Skin and skull. Adult female. Locality. Colombia: (Magdalena), Santa Marta Mountains, near Santa Marta. 10 February 1898. Collector. W. W. Brown, Jr. Original number 60. Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Condition. Skin complete. Skull partial (left zygo- matic arch missing). Type Series. Holotype only. Comments. G. longirostris was considered a valid species by Koopman (1993: 184) and Nowak (1999: 368). Genus LONCHOPHYLLA Thomas, 1903 Lonchophylla hesperia G. M. Allen, 1908 Bull. Mus. Comp. Zool., 52: 35, July. Holotype. MCZ 7011. Body in alcohol, skull extract- ed. Adult male. Locality. Peru: (Contralmirante Villar), Tumbes, Zorritos. Collector. F. H. Bradley. Condition. Alcoholic, skull complete. Type Series. 2 paratypes; in the YPM; 1034 and 1035; both in alcohol. Comments. Considered a valid species by Koop- man (1993: 181) and Nowak (1999: 372). This rare- ly collected bat is known by only two museum specimens in addition to the type series: USNM 283177 and LSUMZ 14121 (Gardner 1976: 5). Genus PLATYRRHINUS de Saussure, 1860 Vampyrops umbratus Lyon, 1902 Proc. Biol. Soc. Washington, 15: 151, 20 June. =Platyrrhinus umbratus (Lyon, 1902). See Koopman (1993: 191). Holotype. B8180. Skin and skull. Adult male. Locality. Colombia: (LaGuajira), San Miguel. 8 June 1898. Collector. W. W. Brown, Jr. Original number 234. Condition. Skin and skull complete. Type Series. 2 paratypes; B8300, skin, male; B8301, skin, male. Comments. P. umbratus was considered a valid spe- cies by Koopman (1993: 191) and Nowak (1999: 389). Platyrrhinus has priority over the genus name Vampyrops (Gardner and Ferrell 1990: 501-503). Vampyrops zarhinus H. Allen, 1891 Proc. Acad. Nat. Sci. Philadelphia, 1891, p. 400, 22 September. =Platyrrhinus helleri (Peters, 1866). See Hall and Kelson (1959: 131). Holotype. MCZ 3211. Body in alcohol, skull extract- ed. Adult female, pregnant. Locality. Panama: Canal Zone, Obispo. 1872. See comments. Collector. Hassler Expedition. Condition. Alcoholic, skull complete. Type Series. Holotype only. TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden Comments. In the original description, H. Allen re- ported that this specimen had been collected in Brazil by the Thayer expedition. G. M. Allen emended this apparently erroneous locality to Obispo, Panama, in accordance with the accession catalogue of the MCZ (1931: 236-237). In support of Allen’s decision, Rouk and Carter (1972: 4) stat- ed, after examining the holotype of zarhinus, that it is “quite like specimens of [Platyrrhinus] helleri from Mexico and Central America, and unlikely to have come from Brazil.” For the use of the genus Platyrrhinus over Vampyrops, see Gardner and Ferrell (1990: 501-503). Genus VAMPYRODES Thomas, 1900 Vampyrodes major G. M. Allen, 1908 Bull. Mus. Comp. Zool., 52: 38, July. =Vampyrodes caraccioli major G. M. Allen, 1908. See Handley (1966: 766). Holotype. MCZ 6756. Body in alcohol. Adult female. Locality. Panama: San Pablo (now covered by Ga- tun Lake). Date unrecorded. Collector. A. Lesley. Condition. Alcoholic. Type Series. Holotype only. Comments. V. c. major was retained as a valid sub- species by Koopman (1994: 88). Family MOLOSSIDAE Gervais, 1855 Genus MOPS Lesson, 1842 Chaerephon leucostigma G. M. Allen, 1918a Bull. Mus. Comp. Zool., 61: 513, February. =Mops condylurus leucostigma (G. M. Allen, 1918). See Koopman (1994: 141). Holotype. MCZ 16344. Skin and skull. Adult female. Locality. Malagasy Republic (Madagascar): (Anta- nanarivo), Tananarive (=Antananarivo). December 1915. Collector. F. R. Wulsin. Condition. Skin partial (bare spot on ventrum). Skull damaged (right and left zygomatic arch miss- ing; supraoccipital chipped). Type Series. 1 paratype; MCZ 16345, skin and skull, male. Comments. Peterson et al. (1995: 168) used the name Tadarida leucostigma. 145 Mops angolensis orientis G. M. Allen and Loveridge, 1942 Bull. Mus. Comp. Zool., 89: 166, February. =Mops condylurus orientis G. M. Allen and Loveridge, 1942. See Koopman (1994: 141). Holotype. MCZ 38829. Skin and skull. Adult male. Locality. Tanganyika Territory (=Tanzania): Mtwa- ra, Ruvuma River, Kitaya, 300 ft (92 m). 3 April 1939. Collector. A. Loveridge. Condition. Skin and skull complete. Type Series. 9 paratypes; MCZ 38826-38828, 38830-38835, all represented by skin and skull, 4 females and 5 males. Genus OTOMOPS Thomas, 1913 Otomops papuensis Lawrence, 1948 J. Mammal., 29: 413, 31 December. Holotype. MCZ 45769. Body in alcohol, skull extract- ed. Adult female. Locality. Papua New Guinea: Vailala River. Collector. Bought from Ward’s Natural Science Es- tablishment, April 1948. Condition. Alcoholic, skull partial (right zygomatic arch missing). Type Series. Holotype only. Comments. Considered a valid species by Koop- man (1993: 239) and Nowak (1999: 482). Accord- ing to Flannery (1995a: 481), O. papuensis has been collected on only two occasions and, other than the holotype, is known by only 10 specimens; 2 in the BMNH, the remainder in the biological collections of the University of Papua New Guinea. Family VESPERTILIONIDAE Gray, 1821 Genus EPTESICUS Rafinesque, 1820 Eptesicus darlingtoni G. M. Allen, 1933 J. Mammal., 14: 150, 15 May. Holotype. MCZ 29113. Skin and skull. Adult female. Locality. Australia: Queensland, Queensland Na- tional Park, MacPherson Ranges, 3,000 ft (915 m). 10 March 1932. Collector. P. J. Darlington, Jr., Harvard Australian Expedition. Original number 30. Condition. Skin and skull complete. Type Series. 1 paratype; MCZ 29120 (now Queens- land Museum J 5476), skin and skull, adult female. Comments. McKean et al. (1978: 533) and Koop- man (1993: 203) included darlingtoni in Eptesicus pumilus. However, Koopman also used the name Pipistrellus darlingtoni (1994: 116). Hoye (1995: 146 537) considered darlingtoni to be a valid species of Vespadelus, to which he gave full generic rank. Genus HARPIOCEPHALUS Gray, 1842 Harpiocephalus rufulus G. M. Allen, 1913 Proc. Biol. Soc. Washington, 26: 214, 20 December. =Harpiocephalus harpia rufulus G. M. Allen, 1913. See Ellerman and Morrison- Scott (1951: 187). Holotype. MCZ 14206. Skin and skull. Adult male. Locality. Vietnam: Tonkin, Lao-Kai (=Lao Cai). 3 January 1912. Collector. Kobayashi Collection. Original number 14. Condition. Skin complete. Skull partial (parietals broken). Type Series. Holotype only. Comments. H. h. rufulus was retained as a valid subspecies by Koopman (1994: 133). Genus IDIONYCTERIS Anthony, 1923 Corynorhinus phyllotis G. M. Allen, 1916b Bull. Mus. Comp. Zool., 60: 352, April. =Idionycteris phyllotis phyllotis (G. M. Allen, 1916). See Tumlison (1993: 418). Holotype. MCZ 5943. Skin and skull. Adult. Locality. Mexico: San Luis Potosf. 24 March 1878. Collector. E. Palmer. Condition. Skin and skull complete. Type Series. Holotype only. Comments. I. phyllotis was considered a valid spe- cies by Koopman (1993: 205) and Nowak (1999: 457). Genus LAS/URUS Gray, 1838 Atalapha brachyotis J. A. Allen, 1892 Bull. Amer. Mus. Nat. Hist., 4: 47, 25 March. = Lasiurus borealis brachyotis (J. A. Allen, 1892). See Niethammer (1964: 595). Holotype. MCZ 11143. Body in alcohol. Male. Locality. (Ecuador), Galapagos Islands: Chatham Island. 23 June 1891. Collector. G. Baur. Condition. Alcoholic. The specimen was received without a skull. Type Series. Holotype only. Comments. Lasiurus brachyotis has often been ac- corded specific status, as in Nowak (1999: 451). In- cluded in L. borealis as a valid subspecies by Koop- man (1994: 129). Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Genus MYOTIS Kaup, 1829 Myotis abbotti nugax G. M. Allen and Coolidge, 1940 Bull. Mus. Comp. Zool., 87: 137, 31 December. = Myotis muricola nugax G. M. Allen and Coolidge, 1940. See Koopman (1994: 104). Holotype. MCZ 36076. Skin and skull. Adult male. Locality. Malaysia: north Borneo, Sabah, Mount Kinabalu, Bundutuan, 3,500 ft (1,068 m). 25 July 1937. Collector. J. A. Griswold, Jr., Asiatic Primate Ex- pedition. Original number 626. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 16 paratypes, MCZ 36072-36075, 36077—36080, 36082-83, 36085—89, 36091: all rep- resented by skin and skull, 12 females and 4 males. Myotis albicinctus G. M. Allen, 1919b J. Mammal., 1: 2, 28 November. = Myotis lucifugus carissima Thomas, 1904. See Miller and G. M. Allen (1928: 50). Holotype. MCZ 11747. Skin and skull. Adult male. Locality. (United States): California, (Tulare Coun- ty), Mount Whitney, 11,000 ft (3,355 m). 14 July 1915. Collector. G. M. Allen. Original number 1. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype, probably at USNM. Comments. The skull of the holotype, which had been mislaid at the time that albicinctus was de- scribed, has subsequently been found and reunited with its skin. Myotis sodalis Miller and G. M. Allen, 1928 Bull. U.S. Nat. Mus., 144: 130, 25 May. Holotype. MCZ 10988. Skin and skull. Adult female. Locality. (United States): Indiana, (Crawford County), Wyandotte cave. 7 March 1904. Collector. J. O. Sibert. Condition. Skin and skull complete. Mandible dis- articulated. Comments. Considered a valid species by Koop- man (1993: 215) and Nowak (1999: 419). Type Series. Miller and Allen based their descrip- tion on an examination of 443 specimens, described by locality in the original description (1928: 133). 30 paratypes are in the MCZ, the others are in the FMNH, USNM, AMNH, and BMNH. TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Genus NYCTICEIUS Rafinesque, 1819 Nycticeius africanus G. M. Allen, 1911b Bull. Mus. Comp. Zool., 54: 328, December. =Nycticeius schlieffeni albiventer Thomas and Wroughton, 1908. See Hayman and Hill (1971: 36). Holotype. MCZ 8272. Skin and skull. Male. Locality. British oe Africa (=Kenya): Meru Riv- er, effluent of northern Guaso Nyiro (=Ewaso Ngi- ro). 11 August 1909. Collector. G. M. Allen. Original number 113. Condition. Skin and skull complete. Type Series. Holotype only. Genus PIPISTRELLUS Kaup, 1829 Eptesicus phasma G. M. Allen, 1911b Bull. Mus. Comp. Zool., 54: 327, December. = Pipistrellus rendalli phasma (G. M. Allen, 191); See, Koopman (1994: 117): Holotype. MCZ 8279. Skin and skull. Male. Locality. British East Africa (=Kenya): Meru Riv- er, effluent of northern Guaso Nyiro (=Ewaso Ngi- ro). 6 August 1909. Collector. G. M. Allen. Original number 94. Condition. Skin and skull complete. Type Series. There is a small series of paratypes in the MCZ. Scabrifer notius G. M. Allen, 1908 Bull. Mus. Comp. Zool., 52: 46, July. =Pipistrellus capensis notius (G. M. Allen, 1908). See Koopman (1994: 117). Holotype. MCZ 4555. Alcoholic, skull extracted. Adult male. Locality. South Africa: (Western Cape), Cape Town. Collector. Received from E. L. Layard, August 1864. Condition. Alcoholic, skull partial (right and left zygomatic arches missing; supraoccipital chipped). Type Series. Holotype only, Genus PLECOTUS E. Geoffroy, 1813 Plecotus sacrimontis G. M. Allen, 1908 Bull. Mus. Comp. Zool., 52: 50, July. =Plecotus auritus sacrimontis G. M. Allen, 1908. See Ognev (1928: 607). Holotype. MCZ 6932. Body in alcohol. Adult male. Locality. Japan: (Honshu), Mount Fuji. 4 Decem- ber 1906. 147 Collector. A. Owston. Condition. Alcoholic. Type Series. Holotype only. Comments. P. a. sacrimontis was retained as a valid subspecies by Koopman (1994: 110). Genus SCOTOPHILUS Leach, 1821 Scotophilus altilis G. M. Allen, 1914d Bull. Mus. Comp. Zool., 58: 350, July. = Scotophilus leucogaster (Cretzschmar, 1826). See Koopman (1993: 227). Holotype. MCZ 14463. Skin and skull. Adult male. Locality. Sudan: Blue Nile, north of (Er) Roseires, Aradeiba. 22 a ary 1913. Collector. G. Allen, Phillips Sudan Expedition. Original Mehta 73. Condition. Skin and skull complete. Type Series. 3 paratypes; MCZ 14462, skin and skull, male, exchanged to FMNH; and 14610 and 14611, both males in alcohol. Family THYROPTERIDAE Miller, 1907 Genus THYROPTERA Spix, 1823 Thyroptera tricolor albigula G. M. Allen, 1923c Proc. New England Zool. Club, 9: 1, 10 December. = Thyroptera tricolor albiventer (Tomes, 1856). See Dunn (1931: 430). Holotype. MCZ 20143. Body in alcohol, skull extract- ed. Adult female. Locality. Panama: Gutierrez, 25 miles (40.2 km) inland from Chiriquiscito on trail from Chiriqui La- goon, Bocas del Toro to Boquete, Chiriqui. August 1923. Collector. E. R. Dunn and C. B. Duryea. Condition. Alcoholic, skull complete. Type Series. 3 paratypes; MCZ 20144, adult male; MCZ 20145, juvenile; MCZ 20146, juvenile; all in alcohol. Order ARTIODACTYLA Owen, 1848 Family TAYASSUIDAE Palmer, 1897 Genus PECARI Reichenbach, 1835 Tayassu crusnigrum Bangs, 1902b Bull. Mus. Comp. Zool., 39: 20, April. =Pecari tajacu crusnigrum (Bangs, 1902). See Hershkovitz (1951: 567). Holotype. MCZ 10163. Skin and skull. Adult male. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 4,000 ft (1,220 m). 13 April 1901. 148 Collector. W. W. Brown, Jr. Original number 290. Condition. Skin and skull complete. Type Series. 2 paratypes; MCZ 10162, adult fe- male; MCZ 10164, juvenile female. Comments. P. t. crusnigrum was retained as a valid subspecies by Hall (1981: 1080) under the genus Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Locality. China: Kiangsu (=Jiangsu), Kiangyin, 80 miles (129 km) northwest of Shanghai. 7 April 1922. Collector. F. R. Wulsin. Condition. Skin and postcranial skeleton complete. Type Series. Holotype only. Comments. For a discussion of the nomenclature and synonymy of this form, consult van Bree (1973). Because Neomeris asiaorientalis Pilleri and Gihr, 1972 is in fact a replacement name for the preoccupied name Delphinus melas Schlegel, 1841, Dicotyles. Tayassu torvus Bangs, 1898k Proc. Biol. Soc. Washington, 12: 164, 10 August. cs 5 I) eh the holotype of this new name is the same as that =Pecari tajacu torvus (Bangs, 1898). See of Schlegel’s name, RMNH 23079. comments. Family CERVIDAE Goldfuss, 1820 Genus ODOCOILEUS Rafinesque, 1832 Cariacus osceola Bangs, 1896b Proc. Biol. Soc. Washington, 10: 26, 25 February. =Odocoileus virginianus osceola (Bangs, 1896). See Lydekker (1915: 148). Holotype. B2394. Skin and skull. Adult female. Locality. (United States): Florida, Citrus County, Citronelle. 29 December 1893. Collector. F. L. Small. Original number 1107. Condition. Skin and skull complete. Mandible dis- a articulated. Beluga declivis Cope, 1865 Type Series. 4 paratypes; B2391, adult male, Proc. Acad. Nat. Sci. Philadelphia, 17: B2392, adult male, B2393, adult female, B2395, 27S. juvenile male, all represented by skin and skull. =Delphinapterus leucas (Pallas, 1776). Comments. O. v. osceola was retained as a valid Gan ee een (1966: aa ) i baka by Hall (1981: 1096) and Smith (1991: Holotype. MCZ 1195. Skull and postcranial skeleton. Locality. “Arctic Seas” (probably Greenland). Collector. E. K. Kane. Condition. Skull partial (right mandibular ramus missing). Postcranial skeleton complete except for missing left flipper. Right flipper and tail are un- cleaned, with tissue largely intact. Type Series. Holotype only. Holotype. BS038. Skin and skull. Adult male. Locality. Colombia: Magdalena, Santa Marta. 26 January 1898. Collector. W. W. Brown, Jr. Original number 50. Condition. Skin and skull complete. Type Series. Holotype only. Comments. Retained as a valid subspecies by Ca- brera (1961: 319) under the genus Tayassu. Use of the genus Pecari follows Grubb (1993: 380). Family MONODONTIDAE Gray, 1821 Genus DELPHINAPTERUS Lacépede, 1804 Odocoeleus [sic] virginianus louisianae G. M. Allen, 1901 Amer. Nat., 35: 449, 28 June. = Odocoileus virginianus macroura (Rafinesque, 1817). See Miller and Kellogg (1955: 804). Holotype. B9111. Skin and skull. Adult male. Locality. (United States): Louisiana, Morehouse Parish, Mer Rouge. 8 November 1898. Collector. B. V. Lilly. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; B9112, B8622, B8623, all males represented by skin and skull. Comments. Hershkovitz (1966: 111) erroneously stated that the holotype of declivis was deposited in the Academy of Natural Sciences in Philadel- phia, and the holotype of Beluga concreta Cope, 1865 was in the MCZ. The opposite is in fact true; Philadelphia holds the type of B. concreta. Family PHOCOENIDAE Gray, 1825 ens WS CnOCAIEN A Pals Ue Odocoileus americanus borealis Miller, Neomeris asiaeorientalis Pilleri and Gihr, 1900 1972 Bull. New York State Mus. 8: 83, 21 Invest. Cetacea, 4: 126. November. =Neophocaena phocaenoides asiaorientalis (Pilleri and Gihr, 1972). See van Bree (1973: 17). Holotype. MCZ 19998 (but see comments). Skull and postcranial skeleton. Adult male. =Odocoilus virginianus borealis Miller, 1900. See Trouessart (1905: 704). Holotype. B4999. Skin and skull. Adult male. Locality. (United States): Maine, (Hancock Coun- ty), Bucksport. 12 December 1895. TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Collector. A. G. Dorr. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. O. v. borealis was retained as a valid subspecies by Hall (1981: 1092) and Smith (1991: ib) Odocoileus virginianus clavium Barbour and G. M. Allen, 1922 J. Mammal., 3: 73, 9 May. Holotype. MCZ 19120. Skull and head skin. Adult male. Locality. (United States): Florida, (Monroe Coun- ty), Big Pine Key. Winter 1920. Collector. T. Barbour. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 2 paratypes, MCZ 18497, skin and skull, juvenile male; MCZ 18060, skin and skull, juvenile male. Comments. Retained as a valid subspecies by Hall (1981: 1093) and Smith (1991: 1). Genus RANGIFER Hamilton Smith, 1827 Rangifer arcticus caboti G. M. Allen, 1914a Proc. New England Zool. Club, 4: 104, 24 March. = Rangifer tarandus caribou (Gmelin, 1788). See Banfield (1962: 70). Holotype. MCZ 15372. One shed antler. Adult male. Locality. Canada: northeast coast of Labrador, about 30 miles (48.3 km) north of Nachvak. 1909. Collector. O. Bryant. Condition. Single antler; complete. Type Series. Holotype only. Rangifer terraenovae Bangs, 1896f Preliminary Description of the Newfoundland Caribou, Boston, p. 1, 11 November. = Rangifer tarandus caribou (Gmelin, 1788). See Banfield (1962: 70). Holotype. B3778. Skull and head skin. Adult male. Locality. Canada: Newfoundland, Codroy. 8 Sep- tember 1895 Collector. F. Doane. Condition. Skull and head skin complete. Type Series. Bangs remarks that he has “secured a series of this fine caribou,” corresponding to B3779-—B3781 and B5757—B5760. Comments. J. A. Allen published a description of Rangifer terraenovae on 21 November 1896, spec- ifying AMNH 11775, a mounted specimen of a 149 male adult, as the holotype (1896: 233). Bangs’ de- scription of this taxon pre-dates J. A. Allen’s by 10 days and thus has priority. Family BOVIDAE Gray, 1821 Genus DAMALISCUS Sclater and Thomas, 1894 Damaliscus phillipsi Harper, 1939 Proc. Biol. Soc. Washington, 50: 90, 5 June. =Damaliscus pygargus phillipsi Harper, 1939. See comments. Holotype. MCZ 35443. Skull and skin. Adult male. Locality. South Africa: Orange Free State. 23 July 1935. Collector. P. Andreka. Original number 1958d. Condition. Skin complete. Skull partial (most of palate, left maxilla, and left mandibular ramus missing). Type Series. Paratype material consists of MCZ 35444, skin and skull of an adult female as well as the following, which Harper examined in the col- lection of the Academy of Natural Sciences of Phil- adelphia: “a mounted head, a skull, and a set of horns purchased in Kimberley, Cape Province; a mounted head and a set of horns from ‘South Af- rica’; and two skins and skulls from the Zoological Society of Philadelphia.” Comments. Ansell (1972: 55) used the name Dam- aliscus dorcas phillipsi. For the use of pygargus over dorcas, see Rookmaaker (1991: 190). Order CARNIVORA Bowdich, 1821 Family CANIDAE Fischer de Waldheim, 1817 Genus CANIS Linnaeus, 1758 Canis lupus beothucus G. M. Allen and Barbour, 1937 J. Mammal., 18: 230, 14 May. Holotype. MCZ 351. Skull and postcranial skeleton. Adult, probably male. Locality. Canada: Newfoundland. About 1865. Collector. J. M. Nelson. Condition. Skull and postecranial skeleton com- plete. Type Series. 4 paratypes; 348, skull, adult male; 349, skull, adult, probably male; 350, skull, adult female; MCZ 28726, skin. Comments. C. |. beothucus became extinct around 1911. Retained as a valid subspecies by Hall (1981: 930). 150 Pachycyon robustus J. A. Allen, 1885 Mem. Mus. Comp. Zool., 10: 4, December. = Canis familiaris Linnaeus, 1758. See G. M. Allen (1920a: 498). Holotype. MCZ 7091. Postcranial skeleton. Locality. (United States): Virginia, Lee County, Ely Cave. Probably 1875. Collector. N. S. Shaler. Condition. Skeleton partial (right scapula, right hu- merus, right femur, right tibia, pelvis). Type Series. Holotype only. Comments. This specimen is a domesticated dog of Native Americans. C. lupus familiaris is the name now widely used for the domestic dog (Wozencraft 1993: 281). PB. robustus is the type species of Pa- chycyon J. A. Allen, 1885. Pachycyon is a synonym of Canis Linnaeus, 1758, which is commonly over- looked, for example, in Wozencraft (1993) and Mc- Kenna and Bell (1997). Genus CERDOCYON Hamilton Smith, 1839 Cerdocyon thous germanus G. M. Allen, 1923b Proc. Biol. Soc. Washington, 36: 55, 28 March. Holotype. MCZ 19850. Skin and skull. Adult. Locality. Colombia: high savannah of Bogota, 9,000 ft (2,745 m). Colleeror N. Maria. Original number 25. Condition. Skin and skull complete. Type Series. 5 paratypes; MCZ 19849, skin and skull, juvenile; MCZ 20097, skin and skull, juvenile male; 3 specimens from the AMNH are also men- tioned in the description. Comments. Retained as a valid subspecies by Berta (1982: 1). Urocyon aquilus Bangs, 1898h Proc. Biol. Soc. Washington, 12: 93, 30 April. =Cerdocyon thous aquilus (Bangs, 1898). See Langguth (1969: 178). Holotype. BSO001. Skin and skull. Adult male. Locality. Colombia: (Magdalena), Santa Marta Mountains, between 2 (000° and 3,000 ft (610-915 m). 10 February 1898. Collector. W. W. Brown, Jr. Original number 58. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. adult female. Comments. Retained as a valid subspecies by Berta @LOS2 auld)» 1 paratype; BS002, skin and skull, Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Genus UROCYON Baird, 1858 Urocyon cinereoargenteus furvus G. M. Allen and Barbour, 1923 Bull. Mus. Comp. Zool., 65: 266, February. Holotype. MCZ 19774. Skin and skull. Probably fe- male. Locality. Panama: Canal Zone, 3 miles (4.8 km) west of Balboa. April 1922. Collector. T. Barbour and W. S. Brooks. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. Retained as a valid subspecies by Hall (1981: 943) and Fritzell and Haroldson (1982: 1). Urocyon cinereoargenteus ocythous Bangs, 1899h Proc. New England Zool. Club, June. 1: 43, 5 Holotype. B4290. Skin and skull. Adult female. Locality. (United States): Wisconsin, Grant Coun- ty, Platteville. 25 January 1896. Collector. N. E. France. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. Retained as a valid subspecies by Hall (1981: 943) and Fritzell and Haroldson (1982: 1). Genus VULPES Frisch, 1775 Vulpes deletrix Bangs, 1898d Proc. Biol. Soc. Washington, 12: 36, 24 March. = Vulpes vulpes rubricosa Bangs, 1898. See Churcher (1960: 359). Holotype. B6967. Skin and skull. Adult female. Locality. (Canada): Newfoundland, Bay St. George. 24 April 1897. Chllezor E. Doane. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Vulpes pennsylvanica vafra Bangs, 1897f Proc. Biol. Soc. Washington, 11: 53, 16 March. Name preoccupied by Vulpes vafer Leidy, 1869. TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Vulpes pennsylvanica rubricosa Bangs, 1898a Science, n. ser., 7: 271, 25 February. (Replacement name for Vulpes pennsylvanica vafra Bangs, 1897) = Vulpes vulpes rubricosa Bangs, 1898. See Churcher (1960: 359). Holotype. B116. Skin and skull. Adult female. Locality. (Canada): Nova Scotia, Digby. 3 Novem- ber 1893. Collector. O. Bangs. Condition. Skin and skull complete. Type Series. 4 paratypes; B1991, skin and skull, and B2001, skull, both adult males; B1992, skin and skull, and B2002, skull. Comments. V. v. rubricosa was retained as a valid subspecies by Hall (1981: 939). Vulpes fulvus rub- ricatus Miller, 1900 (p. 128) was a misspelling and thus accidental renaming of Vulpes pennsylvanica rubricosa Bangs, 1898. Vulpes rubricosa bangsi Merriam, 1900 Proc. Washington Acad. Sci., 2: 667, 28 December 28. = Vulpes vulpes rubricosa Bangs, 1898. See Churcher (1960: 359). Holotype. B8880. Skin and skull. Juvenile female. Locality. (Canada): Labrador, Lance (=Lanse) au Loup. 2 October 1899. Collector. E. Doane. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype; B8879, skin and_ skull, adult male. Family URSIDAE Fischer de Waldheim, 1817 Genus URSUS Linnaeus, 1758 Ursus (Euarctos) americanus sornborgeri Bangs, 1898) Amer. Nat., 32: 500, July. = Ursus americanus americanus Pallas, 1780. See Bangs (1909: 467). Holotype. B7411. Skull. Adult, probably female. Locality. Canada: Labrador, Okkak (=Okak). Sum- mer 1897. Collector. J. D. Sornborger, obtained “from the Es- kimo.” Condition. Skull complete. Type Series. 2 paratypes; B7412, skull, female; B7413, skull, female. Comments. A skull from Hopedale, Labrador (MCZ 7365), has in the past been erroneously la- beled as the holotype of sornborgeri; B7411, the 151 true holotype of sornborgeri, is now correctly la- beled as such. Family PROCYONIDAE Gray, 1825 Genus PROCYON Storr, 1780 Procyon gloveralleni Nelson and Goldman, 1930 J. Mammal., 11: 453, 11 November. =Procyon lotor (Linnaeus, 1758). See Corbet and Hill (1991: 104). Holotype. MCZ 18591. Skin and skull. Juvenile male. Locality. Barbados. 1920. Collector. F. Watts. Condition. Skin and skull complete Type Series. Holotype only. Comments. Considered a valid species by Wozen- craft (1993: 335) and Nowak (1999: 698) but almost certainly introduced to Barbados in the 17th cen- tury (Helgen and Wilson, in prep.). The last rac- coon on Barbados was seen in 1964, and the pop- ulation is probably extinct. Procyon lotor elucus Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 219, 15 March. Holotype. B3502. Skin and skull. Adult male. Locality. (United States): Florida, Brevard County, Oak Lodge, east peninsula opposite Micco. 15 Feb- ruary 1895. Collector. O. Bangs. Condition. Skin complete. Skull partial (condyle, coronoid, and angular processes of left mandibular ramus broken). Mandible disarticulated. Type Series. There is a series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 968). Procyon maynardi Bangs, 1898g Proc. Biol. Soc. Washington, 12: 92, 30 April. =Procyon lotor (Linneaus, 1758). See Koopman et al. (1957: 164). Holotype. B7750. Skin and skull. Juvenile male. Locality. Bahamas: New Providence Island, Nas- sau. August 1897. Collector. H. L. Claridge. Condition. Skin complete. Skull partial (broken from frontals to occiput). Mandible disarticulated. Type Series. Holotype only. Comments. Considered a valid species by Wozen- craft (1993: 336) and Nowak (1999: 698) but un- doubtedly a recent introduction to New Providence Island (see Olson and Pregill, 1982: 5). 152 Family MUSTELIDAE Fischer de Waldheim, 1817 Genus LONTRA Gray, 1843 Lutra degener Bangs, 1898d Proc. Biol. Soc. Washington, 12: 35, 24 March. =Lontra canadensis canadensis (Schreber, 1776). See van Zyll de Long @lIS72aS1)) Holotype. B6965. Skin and skull. Adult male. Locality. Canada: Newfoundland, Bay St. George. 23 April 1897. Collector. E. Doane. Condition. Skin and skull complete. Type Series. Paratype material consists of B6966, skin and skull of an adult female, mentioned by number in the original description, as well as “two extra skulls,” corresponding to B3755 and B3799, and “a large series of unsexed otter skulls from Newfoundland,” corresponding to MCZ 494—508. Lutra hudsonica vaga Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 224, 15 March. =Lontra canadensis laxatina F. Cuvier, 1823. See van Zyll de Long (1972: 81). Holotype. B5749. Skin and skull. Adult male. Locality. (United States), Florida, Brevard County, Micco. 17 March 1897. Collector. F. R. Hunter. Condition. Skin and skull complete. Type Series. 4 paratypes; B4995, skin and skull, adult female; B4998, skin and skull, adult male; BG6092, skin and skull, adult male; B6093, skin and skull, adult female. Genus MARTES Pinel, 1792 Mustela atrata Bangs, 1897b Amer. Nat., 31: 162, 1 February. =Martes americana atrata (Bangs, 1897). See G. M. Allen (1942: 166) Holotype. B5752. Skin and skull. Adult female. Locality. Canada: Newfoundland, Bay St. George. 29 September 1896. ; Collector. E. Doane. Original number 2. Condition. Skin and skull complete. Type Series. 12 paratypes; B5751, skin and skull, adult female; MCZ 492-93, 509-517, unsexed skulls. Comments. M. a. atrata was retained as a valid sub- species by Hall (1981: 983). Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Mustela brumalis Bangs, 1898) Amer. Nat., 32: 502, July. =Martes americana atrata (Bangs, 1897). See Clark et al. (1987: 1). Holotype. B7417. Skull. Adult, probably male. Locality. Canada: Labrador, Okkak (=Okak). Sum- mer 1897. Collector. J. D. Sornborger, obtained “from the Es- kimo.” Condition. Skull complete. Type Series. 2 paratypes; B7418, skull; B7419, skull; both probably male. Genus MUSTELA Linnaeus, 1758 Mustela cicognanii mortigena Bangs, 1913 Bull. Mus. Comp. Zool., 54: 511, July. = Mustela erminea richardsonii Bonaparte, 1838. See Hall (1951: 110). Holotype. B3745. Skin and skull. Adult male. Locality. Canada: Newfoundland, Bay St. George. 27 September 1895. Collector. E. Doane. Original number 1. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Putorius frenatus neomexicanus Barber and Cockerell, 1898 Proc. Acad. Nat. Sci. Philadelphia, 1898, p- 188, May. = Mustela frenata neomexicana (Barber and Cockerell, 1898). See Miller (1912: 100). Holotype. MCZ 10475. Skin with extra tail, and skull. Adult male. Locality. (United States): New Mexico, (Dona Ana County), Mesilla, shore of Armstrongs’ Lake, 3,800 ft (1,159 m). 1 February 1898. Collector. A. C. Tyson. Original number 58. Condition. Skin complete, but tail poorly prepared. The tail belonging to a discarded topotype is tied to the holotype as an example. Skull complete. Type Series. A topotype taken at the same time as the holotype was partially decomposed and dis- carded; the tail of this specimen is included with MCZ 10475. The original description also refers to “a specimen, without any history, in alcohol ... in the collection of the New Mexico Agricultural Col- lege.” Comments. M. f. neomexicana was retained as a valid subspecies by Hall (1981: 995). TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden Putorius (Arctogale) longicauda oribasus Bangs, 1899m Proc. New England Zool. Club, 1: 81, 27 December. = Mustela frenata oribasus (Bangs, 1899). See Hall (1936: 105). Holotype. B9058. Skin and skull. Adult female. Locality. Canada: British Columbia, source of Ket- tle River, 7,500 ft (2,288 m). 10 September 1898. Collector. A. C. Brooks. Original number 1368. Condition. Skin and skull complete. Type Series. Holotype only. Comments. M. f. oribasus was retained as a valid subspecies by Hall (1981: 998). Putorius (Lutreola) lutensis Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 229, 15 March. = Mustela vison lutensis (Bangs, 1898). See Hollister (1913: 474). Holotype. B7225. Skin and skull. Adult male. Locality. (United States), Florida, St. Johns Coun- ty, salt marsh opposite Matanzas Inlet. 16 February 1897. Collector. O. Bangs. Original number 7. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. M. v. lutensis was retained as a valid subspecies by Hall (1981: 1003) and Lariviére (1999: 1). Putorius (Arctogale) muricus Bangs, 1899} Proc. New England Zool. Club, 1: 71, 31 July. = Mustela erminea muricus (Bangs, 1899). See Hall (1945: 77). Holotype. B9146. Skin and skull. Juvenile male. Locality. (United States): California, El Dorado County, Echo, 7,500 ft (2,288 m). 15 July 1897. Collector. W. W. Price and E. M. Nutting. Original number 266. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. Holotype only. Comments. M. e. muricus was retained as a valid subspecies by Hall (1981: 990) and King (1983: 1). Putorius noveboracensis notius Bangs, 1899) Proc. New England Zool. Club, 1: 53, 9 June. = Mustela frenata noveboracensis (Emmons, 1840). See Hall (1936: 104). Holotype. B2678. Skin and skull. Juvenile male. Locality. (United States): North Carolina, Bun- combe County, Weaverville. 10 July 1892. 153 Collector. J. S. Cairns. Original number 2214. Condition. Skin complete. Skull partial (two frag- ments only, premaxilla—lachrymal). Mandible dis- articulated. Type Series. 2 paratypes; AMNH 1247, adult male; USNM 32239, adult male. Putorius occisor Bangs, 1899i Proc. New England Zool. Club, 1: 54, 9 June. = Mustela frenata occisor (Bangs, 1899). See Hall (1936: 104). Holotype. B9102. Skin and skull. Adult male. Locality. (United States): Maine, Hancock County, Bucksport, near mouth of Penobscot River. 15 Jan- uary 1899. Collector. A. G. Dorr. Condition. Skin complete. Skull slightly damaged (left zygomatic arch broken). Type Series. There is a series of paratypes in the MCZ. Comments. M. f. occisor was retained as a valid subspecies by Hall (1981: 997). Putorius rixosus Bangs, 1896a Proc. Biol. Soc. Washington, 10: 21, 25 February. = Mustela nivalis rixosa (Bangs, 1896). See Reichstein (1958: 169). Holotype. B642. Skin and skull. Adult female. Locality. Canada: Saskatchewan, Osler. 15 July 1893. Collector. W. C. Colt. Original number 79/181. Condition. Skin complete. Skull slightly damaged (left zygomatic arch broken). Type Series. Three specimens other than the ho- lotype are mentioned by number in the original description; MCZ 5532, USNM 4231, probably fe- male, and USNM 13904, probably male. All are unsexed skins. Comments. M. n. rixosa was retained as a valid sub- species by Hall (1981: 993) and Sheffield and King (1994: 1). Putorius vison energumenos Bangs, 1896c Proc. Boston Soc. Nat. Hist., 27: 5, March. = Mustela vison energumenos (Bangs, 1896). See Miller (1912: 101). Holotype. B3555. Skin and skull. Adult male. Locality. Canada, British Columbia, Sumas. 23 September 1895. Collector. A. C. Brooks. Original number 514. Condition. Skin and skull complete. 154 Type Series. | paratype; B3556, skin and skull, ju- venile male. Comments. M. v. energumenos was retained as a valid subspecies by Hall (1981: 1001) and Lariviére (1999: 1). Putorius (Lutreola) vulgivagus Bangs, 1895b Proc. Boston Soc. Nat. Hist., 26: 539, 31 July. = Mustela vison vulgivaga (Bangs, 1895). See Miller (1912: 102). Holotype. B2751. Skin and skull. Adult male. Locality. (United States): Louisiana, Plaquemines Parish, Burbridge. 10 January 1895. Collector. F. L. Small. Original number 1439/54. Condition. Skin and skull complete. Type Series. 10 paratypes; B2752-B2761, all rep- resented by skin and skull, 9 males and 1 female. Comments. M. v. vulgivaga was retained as a valid subspecies by Hall (1981: 1004) and Lariviére (1999: 1). Putorius xanthogenys mundus Bangs, 1899i Proc. New England Zool. Club, 1: 56, 9 June. = Mustela frenata munda (Bangs, 1899). See Hall (1936: 107). Holotype. B5459. Skin and skull. Adult male. Locality. (United States): California, Marin Coun- ty, Point Reyes. 19 June 1896. Collector. C. A. Allen. Original number 931. Condition. Skin complete. Skull slightly damaged (left zygomatic arch broken). Type Series. | paratype, BS8632 (not B8631, men- tioned erroneously in the original description), skin and skull, male. Comments. M. f. munda was retained as a valid subspecies by Hall (1981: 995). Family MEPHITIDAE Bonaparte, 1845 Genus MEPHITIS E. Geoffroy and G. Cuvier, 1795 Mephistis avia Bangs, 1898c Proc. Biol. Soc. Washington, 12: 32, 24 March. =Mephitis mephitis avia Bangs, 1898. See Hall (1936: 65). Holotype. B5747. Skin and skull. Adult male. Locality. (United States): Illinois, Mason County, San Jose. 10 March 1897. Collector. H. H. and C. S. Brimley. Original num- ber 2500. ; Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Condition. Skin and skull complete. Type Series. 1 paratype; B5783, skin and skull, adult male. Comments. M. m. avia was retained as a valid sub- species by Hall (1981: 1019) and Wade-Smith and Verts (1982: 1). Mephitis mephitica elongata Bangs, 1895b Proc. Boston Soc. Nat. Hist., 26: 531, 31 July. = Mephitis mephitis elongata Bangs, 1895. See A. H. Howell (1921: 39). Holotype. B3051. Skin and skull. Adult male. Locality. (United States): Florida, Brevard County, Micco. 5 March 1895. Collector. O. Bangs. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. Retained as a valid subspecies by Hall (1981: 1019) and Wade-Smith and Verts (1982: 1). Mephitis mephitica scrutator Bangs, 1896i Proc. Biol. Soc. Washington, 10: 141, 28 December. = Mephitis mephitis mesomelas Lichtenstein, 1832. See Hall (1936: 66). Holotype. B2889. Skin and skull. Adult male. Locality. (United States): Louisiana, Acadia Parish, Cartville. 25 May 1895. Collector. F. L. Small. Original number 1842. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 1 paratype, B2886, skin and skull, adult female. Mephitis spissigrada Bangs, 1898c Proc. Biol. Soc. Washington, 12: 31, 24 March. = Mephitis mephitis spissigrada Bangs, 1898. See Hall (1936: 67). Holotype. B3699. Skin and skull. Adult female. Locality. Canada: British Columbia, Sumas. 30 September 1895. Collector. A. C. Brooks. Original number 518. Condition. Skin and skull complete. Mandible dis- articulated. Type Series. 3 paratypes; B3700, skin and skull, adult female, and B5548, skin and skull, adult male; B7435, skull, adult male. Comments. M. m. spissigrada was retained as a val- id subspecies by Hall (1981: 1022 and Wade- Smith and Verts (1982: 1). TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden Genus SPILOGALE Gray, 1865 Spilogale ambarvalis Bangs, 1898b Proc. Boston Soc. Nat. Hist., 28: 222, 15 March. = Spilogale putorius ambarvalis Bangs, 1898. See Van Gelder (1953: 255). Holotype. B3481. Skin and skull. Adult male. Locality. (United States): Florida, Brevard County, Oak Lodge, east peninsula opposite Micco. 30 Jan- uary 1895. Collector. O. Bangs. Original number 11. Condition. Skin and skull complete. Type Series. There is a large series of paratypes in the MCZ. Comments. S. p. ambarvalis was retained as a valid subspecies by Hall (1981: 1014) and Kinlaw (1995: 1). Family VIVERRIDAE Gray, 1821 Genus EUPLERES Doyere, 1835 Eupleres major Lavauden, 1929 Compt. Rend. Acad. Sci. Paris, 189: 198, QS) July DOR =Eupleres goudotii major Lavauden, 1929. See Albignac (1973: 23). Syntypes. MCZ 45691: Skin, skull, and poster: anial skeleton. Subadult female. MCZ 45962: Skin, skull, and postcranial skeleton. Subadult male. Locality. Madagascar: (Antsiranana), foot of the Massif Tsaratanna (=Tsaratanana), Upper Sombi- rano Valley, above village of Beangona, 1,500 m. April 1929. Collector. Lavauden. Condition. MCZ 45691: Skin complete, with bald spot on dorsum and tail slightly damaged. Skull and skeleton complete. Mandible eneeculated Teeth removed from skull but present. MCZ 45962: Skin complete. Skull and skeleton complete. Mandible disarticulated. Teeth removed from skull but pres- ent. Type Series. 2 syntypes, described above. Comments. These are the two specimens from G. Grandidier’s personal collection on which Lavau- den based his original description of Ewpleres ma- jor. Albignac (1973: 23) wrote that these type spec- imens were “introuvable [nowhere to be found].” Family HERPESTIDAE Bonaparte, 1845 Genus GALIDICTIS |. Geoffroy, 1839 Galidictis grandidiensis [sic] Wozencraft, 1986 J. Mammal. 67: 561, 8 August. = Galidictis grandidieri Wozencraft, 1986. See Wozencraft (1987: 198). Holotype. MCZ 45983. Skin, skull, and postcranial skeleton. Adult. 155 Locality. Madagascar (no further data available). The locality of the paratype, stored in the AMNH, s “Madagascar, Lac Tsimanampetsotsa, 24°08’ S, 43°46’ E.” Collector. No collection data available. The holo- type is part of the collection of G. Grandidier. Condition. Skin prepared flat; incomplete (ven- trum missing). Skull complete. Type Series. 1 paratype; AMNH 100478, skin and skull, adult male. Comments. G. grandidieri was considered a valid species by Wozencraft (1993: 300) and Nowak (1999: 769). Family FELIDAE Fischer de Waldheim, 1817 Genus LEPTAILURUS Severtzov, 1858 Felis capensis phillipsi G. M. Allen, 1914d Bull. Mus. Comp. Zool., 58: 337, July. =Leptailurus serval phillipsi (G. M. Allen, 1914). See comments. Holotype. MCZ 14908. Skin and _ skeleton. Adult male. Locality. Sudan: Blue Nile, El Garef. 10 January 1913. Collector. J. C. Phillips. Condition. Skin and skull complete. Type Series. Holotype only. Comments. G. M. Allen (1939: 241) used the name Felis serval phillipsi. The use of the genus Lep- tailurus follows Wozencraft (1993: 292). L. s. phil- lipsi was retained as a valid subspecies by Smithers (1975: 7). Genus LYNX Kerr, 1792 Lynx (Cervaria) fasciatus oculeus Bangs, 1899e Proc. New England Zool. Club, 1: 23, 31 March. = Lynx rufus californicus Mearns, 1897. See Grinnell and Dixon (1924: 346). Holotype. B8633. Skin and skull. Adult male. Locality. (United States): California, Marin Coun- ty, Nicasio. 11 December 1898. Collector. C. A. Allen. Original number 981. Condition. Skin and skull complete. Type Series. 1 paratype; B4789, skin and _ skull, adult male. Lynx gigas Bangs, 1897e Proc. Biol. Soc. Washington, 11: 50, 16 March. =Lynx rufus gigas Bangs, 1897. See Peterson and Downing (1952: 11). Holotype. B4951. Skin and skull. Adult male. Locality. Canada: Nova Scotia, 15 miles (24.1 km) back of Bear River. 11 December 1895. 156 Collector. D. R. Ritchie. Condition. Skin and skull complete. Type Series. Holotype only. Comments. L. r. gigas was retained as a valid sub- species by Hall (1981: 1054). Lynx subsolanus Bangs, 1897e Proc. Biol. Soc. Washington, 11: 49, 16 March. =Lynx canadensis subsolanus Bangs, 1897. See Elliot (1901: 296). Holotype. B1190. Skin and skull. Adult male. Locality. Canada: Newfoundland, Codroy. 13 June 1894. Collector. E. Doane. Condition. Skin and skull complete. Type Series. 2 paratypes; B5754, skin and skull, ju- venile female: B3798, skull, adult male. Comments. L. c. subsolanus was retained as a valid subspecies by Hall (1981: 1051). Genus PUMA Jardine, 1834 Felis bangsi Merriam, 1901 Proc. Washington Acad. Sci., 3: 595, 11 December. =Puma concolor bangsi (Merriam, 1901). See comments. Holotype. B8413. Skin and skull. Adult male. Locality. Colombia: (La Guajira), Dibulla. 8 Oc- tober 1899. Collector. W. W. Brown, Jr. Condition. Skin and skull complete. Type Series. 3 paratypes; B8147, skin and skull, adult female, and “two skulls from Peru, in the American Museum of Natural History.” Comments. Nelson and Goldman (1929: 347) used the name Felis concolor bangsi. The use of the ge- nus Puma follows Wozencraft (1993: 296). Re- tained as a valid subspecies by Currier (1983: 1). Felis bangsi costaricensis Merriam, 1901 Proc. Washington Acad. Sci., 3: 596, 11 December. =Puma concolor costaricensis (Merriam, 1901). See comments. Holotype. MCZ 10118. Skin and skull. Adult female. Locality. Panama: Chiriqui, Boquete (=Bajo Bo- quete), 4,000 ft (1,220 m). 22 April 1901. Collector. W. W. Brown, Jr. Original number 337. Condition. Skin and skull complete. Type Series. There is a series of paratypes in the MCZ. Comments. Nelson and Goldman (1929: 347) used the name Felis concolor costaricensis. The use of the genus Puma follows Wozencraft (1993: 296). Retained as a valid subspecies by Currier (1983: 1). Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Felis coryi Bangs, 1899b Proc. Biol. Soc. Washington, 13: 15, 31 January. =Puma concolor coryi (Bangs, 1899). See comments. Holotype. B7742. Skin and skull. Adult male. Locality. (United States): Florida, Brevard County, “wilderness back of Sebastian”. 1 January 1898 Collector. F. R. Hunter. Condition. Skin and skull complete. Type Series. 5 paratypes; B5489, adult female, B5650, adult female, B6992, adult male, B7743, adult female, B7744, juvenile female; all repre- sented by skin and skull. Comments. Nelson and Goldman (1929: 347) used the name Felis concolor coryi; the use of the genus Puma follows Wozencraft (1993: 296). Felis coryi Bangs, 1899 is a replacement name for Felis con- color floridana Cory, 1896 (1896: 109). Retained as a valid subspecies by Currier (1983: 1). Felis improcera Phillips, 1912 Proc. Biol. Soc. Washington, 25: 85, 4 May. =Puma concolor improcera (Phillips, 1912). See comments. Holotype. MCZ 12704. Skin and skull. Adult male. Locality. (Mexico): Lower (= Baja) California, Cal- malli. 3 September 1911. Collector. E. W. Funcke. Original number 10. Condition. Skin and skull complete. Type Series. Holotype only. Comments. Nelson and Goldman (1929: 347) used the name Felis concolor improcera. The use of the genus Puma follows Wozencraft (1993: 296). Re- tained as a valid subspecies by Currier (1983: 1). Order CIMOLESTA McKenna, 1975 Family MANIDAE Gray, 1873 Genus PHATAGINUS Rafinesque, 1821 Phataginus tricuspis mabirae G. M. Allen and Loveridge, 1942 Bull. Mus. Comp. Zool., 89: 178, February. Holotype. MCZ 39417. Skin, skull, and postcranial skeleton. Adult male. Locality. Uganda: (Buganda), Chagwe, Mabira Forest, Mubango. 12 November 1938. Collector. A. Loveridge. Condition. Skin and skull complete. Postcranial skeleton partial (includes atlas, right tibia, and right fibula). Type Series. Holotype only. TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden yz Comments. Retained as a valid subspecies by Meester (1972: 2). REFERENCES AGuayo, C. G. 1950. Observaciones sobre algunos mamiferas Cubanos Extinguidos. Boletin de His- toria Natural de la Sociedad “Felipe Poey,” 1: 121-134. ALBIGNAC, R. 1973. Faune de Madagascar. 36. Mam- miféres carnivores. Paris: O.R.S.T.O.M, 206 pp. ALLEN, G. M. 1901. The Louisiana deer. 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An insular race of cotton rat from the Florida Keys. Journal of Mammalogy, 1: 235-— 236. . 1921. A new horseshoe bat from West Africa. Revue Zoologique Africaine, 9: 193-196. . 1923a. A new shrew from Colombia. Pro- ceedings of the New England Zoological Club, 8: 37-38. . 1923b. The pampa fox of the Bogota savan- na. Proceedings of the Biological Society of Washington, 36: 55-58. . 1923c. A new disc-winged bat from Panama. Proceedings of the New England Zoological Club, 9: 1-2. . 1925. Squirrels collected by the American Museum Asiatic Expeditions. American Museum Novitates, 163: 1—16. . 1927. Murid rodents from the Asiatic expe- ditions. American Museum Novitates, 270: 1-12. . 1928. A new cricetine genus from China. Journal of Mammalogy, 9: 244-246. . 1931. Type specimens of mammals in the Museum of Comparative Zoology. Bulletin of the Museum of Comparative Zoology, 71: 229-289. . 1933. Two new bats from Australia. Journal of Mammalogy, 14: 149-151. . 1936. 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The cotton mouse Peromyscus gos- sypinus. Proceedings of the Biological Society of Washington, 10: Toe 125. 1896f. Preliminary description of the New- foundland caribou. Hosion, Massachusetts: Al- fred Mudge and Son, 2 pp. . 1896g. Preliminary description of a new vole fon Labradors American Naturalist, 30: 1051. 1896h. Some new mammals from Indian Territory and Missouri. Proceedings of the Bio- logical Society of Washington, 10: 135-138. 1896i. The skunks of the genus Mephitis of iste North America. Proceedings of the Bio- logical Society of Washington, 10: 139-144. 1896j. A review of the squirrels of eastern North America. Proceedings of the Biological Society of Washington, 10: 145-167. . 1897a. A new white-footed mouse from Brit- ish Columbia. American Naturalist, 31: 74—75. 1897b. Preliminary description of the New- foundland marten. American Naturalist, 31: 161-162. . 1897c. 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Science (n.s.), 165: 271-272 1898b. The land Mammals of peninsular Florida and the coast region of Georgia. Pro- ceedings of the Boston Society of Natural His- tory, 28: 157-235. ———.. 1898c. Descriptions of two new skunks of the genus Mephitis. ee of the Biological Society of Washington, 12: 31—33. j 1898d. Descriptions of = Newfoundland otter and red fox. Proceedings of the Biological Society of Washington, 12: 35-38. 1898e. The eastern races of the American varying hare with description of a new subspecies from Nova Scotia. Proceedings of the Biological Society of Washington, 12: 77-82. 1898f. Description of a new white-footed mouse from the Mount Baker Range, British Co- lumbia. Proceedings of the Biological Society of Washington, 12: 83-84. . 1898g. A new raccoon from Nassau Island, Bahamas. Proceedings of the Biological Society of Washington, 12: 91—92. 1898h. Description of a new fox from Santa Marta, Colombia. 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Beijing: China Forestry Publishing House, 280 pp. INDEX abaconis Geocapromys, 132 abbotti Myotis, 146 abbreviata Neotoma, 121 aberti Sciurus, 99 abietorum Peromyscus, 125 ablutus Leggada, 118 acadicus Microtus, 115 Neosorex, 138 Zapus, 112 aequalis Aethalops, 140 aereus Scalops, 140 Scalopus, 140 aestuans Guerlinguetus, 106 Sciurus, 106 Aethalops aequalis, 140 alecto, 140 Aethoglis argenteus, 129 hueti, 129 Aethosciurus byatti, 106 laetus, 106 africanus Nycticeius, 147 Afrosoricida, 101 Agouti paca, 130 virgatus, 130 Agoutidae, 130 akka Funisciurus, 104 Akodon apricus, 127 teguina, 127 xerampelinus, 127 alacer Lepus, 134 Sylvilagus, 134 alashanicus Spermophilus, 107 albicinctus Myotis, 146 albigula Thyroptera, 147 albiventer Nycticeius, 147 Thyroptera, 147 alboniger Hylopetes, 105 Pteromys, 105 alecto Aethalops, 140 alfari Microsciurus, 105 alleni Dasymys, 118 Heteromys, 112 Hylomyscus, 118 Lepus, 133 Liomys, 112 Macrotolagus, 133 Melomys, 118 Rhinolophus, 143 aloga Blarina, 137 Alouatta luctuosa, 135 palliata, 135 pigra, 135 trabeata, 135 alpinus Sciuropterus, 104 altilis Scotophilus, 147 amazonicus Nectomys, 121 ambarvalis Spilogale, 155 americana Martes, 152 americanus Euarctos, 151 Lepus, 133, 134 Odocoileus, 148 Ursus, 151 Ametrida centurio, 143 minor, 143 ammodytes Peromyscus, 125 anastasae Peromyscus, 124, 125 Scalops, 140 Scalopus, 140 anchietae Otomys, 120 anchises Hylopetes, 105 Pteromys, 105 anetianus Pteropus, 141 angolensis Mops, 145 Antechinus mayeri, 100 misim, LOO naso, 100 Antillogale marcanoi, 137 aorensis Pteropus, 141 Aplodontia, 103 californica, 103 columbiana, 103 rainieri, 103 rufa, 103 Aplodontidae, 103 Apodemus euxinus, 117 mystacinus, 17, apricus Akodon, 127 Scotinomys, 127 aquaticus Scalopus, 140 aquilis Cerdocyon, 150 Craseomys, 113 Urocyon, 150 aquilonius Fiber, 116 Ondatra, 116 araeum Plagiodonta, 132 araneus Sorex, 138 arcticus Hesperomys, 124 Lepus, 133 Rangifer, 149 Arctogale longicauda, 153 muricus, 153 oribasus, 153 Arctomys avarus, 105 flaviventer, 105 ignavus, 105 arenarius Peromyscus, 126 argentatus Peromyscus, 125 argenteus Aethoglis, 129 Graphiurus, 129 ariel Pteropus, 141 armatus Spermophilus, 107 Artibeus TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden ill femurvillosum, 144 lituratus, 144 palmarum, 144 Artiodactyla, 145 Arvicola breweri, 114 longipilis, 114 riparia, 114 rufidorsum, 114 terraenovae, 114 Arvicolinae, 112 asiaeorientalis Neomeris, 148 Neophocaena, 148 Atalapha brachyotis, 146 ater Suncus, 139 atrata Martes, 152 Mustela, 152 atrodorsalis Thomomys, 111 attwateri Peromyscus, 124 aurita Geogale, 101 Triaenops, 143 auritus Plecotus, 147 aurora Eothenomys, 113 Microtus, 113 austerulus Sigmodon, 127 austini Pteropus, 141 australis Cryptogale, 101 Echymipera, 101 Reithrodontomys, 127 austrinus Geomys, 110 avara Marmota, 105 avarus Arctomys, 105 avia Cryptotis, 138 Mephitis, 154 bactrianus Mus, 121 badius Ictidomys, 108 Spermophilus, 108 bairdii Lepus, 133, 134 172 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Mus, 98 Peromyscus, 98 baliolus Peromyscus, 126 baluensis, Callosciurus, 103 bangsi Felis, 156 Glaucomys, 104 Perognathus, 112 Puma, 156 Sciuropterus, 104 Vulpes, 151 bangsii Lepus, 133 barbouri Otomys, 121 barrerae Octomys, 130 Tympanoctomys, 130 Bathyergidae, 129 bella Neotoma, 121 bellus Peromyscus, 124 Beluga concreta, 145 declivis, 148 beothucus Canis, 149 bicolor Crocidura, 137 Hipposideros, 142 Oecomys, 122 Blarina aloga, 137 brevicauda, 137 compacta, 137 borealis Lasiurus, 146 Odocoileus, 148 Peromyscus, 124 Synaptomys, 116 Tamias, 109 Boromys torrei, 131 borucae Sigmodon, 127 Bovidae, 149 brachyotis Atalapha, 146 Lasiurus, 146 brasiliensis Sylvilagus, 134 brevicauda Blarina, 137 Zygodontomys, 128 brevicaudata Microgale, 102 breweri Arvicola, 114 Microtus, 114 brochus Syntheosciurus, 108 browni Microsciurus, 105 Sciurus, 105 brumalis Martes, 152 Mustela, 152 bufo Leggada, 118 Mus, 118 burrus Proechimys, 131 byatti Aethosciurus, 106 Paraxerus, 106 caboti Rangifer, 149 cacabatus Peromyscus, 124 caecator Castor, 109 caecutiens Sorex, 138 cafer Pedetes, 128 californica Aplodontia, 103 californicus Dipodomys, 111 Lynx, 165 Scapanus, 140 callida Dasyprocta, 130 Callosciurus baluensis, 103 erythraeus, 103 ferrugineus, 103 haemobaphes, 103 medialis, 103 primus, 103 Caluromyidae, 99 Caluromys cicur, 99 lanatus, 99 canadensis Castor, 109 Lontra, 152 Lynx, 156 Peromyscus, 125 Canidae, 149 canina Peropteryx, 142 Canis beothucus, 149 familiaris, 150 lupus, 149, 150 Cansumys canus, 117 canus Cansumys, 117 capensis Felis, 165 Pedetes, 128 Pipistrellus, 146 Capromyidae, 131 Capromys nana, 132 pilorides, 131 relictus, 131 capucinus Cebus, 136 caraccioli Vampyrodes, 145 Cariacus osceola, 148 caribou Rangifer, 149 carissima Myotis, 146 Carnivora, 149 carolinensis Sciurus, 106 carpenteri Hylobates, 136 carrorum Oryzomys, 123 cascadensis Lepus, 134 castaneoventris Sciurus, 103 castanonotus Sciurus, 99 castanotus Sciurus, 99 castanops Cratogeomys, 111 Pappogeomys, 111 Castor caecator, 109 canadensis, 109 Castoridae, 109 cavator Macrogeomys, 110 Orthogeomys, 110 cayennensis Proechimys, 131 Cebidae, 135 Cebus capucinus, 136 curtus, 136 celatus Phenacomys, 116 centurto Ametrida, 143 Cerdocyon aquilus, 150 germanus, 150 thous, 150 Cervaria fasciatus, 155 oculeus, 155 Cervidae, 148 Cetartiodactyla, 97 Chaerophon leucostigma, 145 Cheiromys laniger, 135 madagascariensis, 135 cherriei Zygodontomys, 128 Chilonycteris parnellii, 143 | pusillus, 143 | torrei, 143 ) Chionomys | nivalis, 112 chionopaes Dicrostonyx, 113 chiriquensis Guerlinguetus, 106 Sciurus, 106 Chiroptera, 140 Chlorotalpa tropicalis, 101 chrotorrhinus Microtus, 114 Chrysochloridae, 101 Chrysochloris stuhlmanni, 101 tropicalis, 101 chrysogaster Lemmus, 114 cicognanii Mustela, 152 cicur Caluromys, 99 Philander, 99 Cimolesta, 156 cinereoargenteus Urocyon, 150 cinereus Sorex, 139 Cingulata, 101 Citellus obscurus, 107 siccus, 1O7 Claviglis collaris, 129 soleatus, 129 clavium Odocoileus, 149 Clethrionomys gapperi, 113 proteus, 113 collaris Claviglis, 129 colombiana Dasyprocta, 130 colonus Geomys, 110 coloratus Oryzomys, 123 columbiana Aplodontia, 103 columbianus Geocapromys, 132 compacta Blarina, 137 concolor Felis, 156 Puma, 156 concreta Beluga, 148 condylurus Mops, 145 confucianus Niviventer, 119 consobrinus Sciurotamias, 106 cooperi Synaptomys, 116 Tamias, 109 coryt Felis, 156 Puma, 156 Corynorhinus phyllotis, 146 costaricensis Felis, 156 Puma, 156 cowant Microgale, 102 Craseomys aquilus, 113 Crassus Phenacomys, 116 Cratogeomys castanops, 111 rubellus, 111 creper Reithrodontomys, 127 Cricetinae, 117 Cricetomyinae, 117 cricetulus Saccostomus, 117 crinitus Peromyscus, 125 Crocidura bicolor, 137 fuscomurina, 137 geata, 138 hildegardeae, 138 maurisca, 138 phaios, 138 tephragaster, 137 crusnigrum Pecari, 147 Tayassu, 147 Cryptogale australis, 101 Cryptomys hottentotus, 129 TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden occlusus, 129 whytei, 129 Cryptotis avia, 138 thomasi, 138 cubanus Geocapromys, 132 Solenodon, 137 cumberlandius Geomys, 110 cuppes Ochotona, 133 curtus Cebus, 136 Hipposideros, 142 Damaliscus phillipsi, 149 pygargus, 149 daphaenodon Sorex, 139 darlingtoni Eptesicus, 145 Pipistrellus, 145 Dasogale fontoynonti, 103 Dasymys alleni, 118 incomtus, 118 Dasypodidae, 101 Dasyprocta callida, 130 colombiana, 130 leporina, 130 noblei, 130 nuchalis, 130 punctata, 130 Dasyproctidae, 130 Dasypus hoplites, 101 novemcinctus, 1O1 Dasyuridae, 100 Dasyuromorphia, 100 Daubentonia madagascariensis, 135 Daubentoniidae, 135 davidanus Sciurotamias, 106 decaryi Microgale, 102 declivis Beluga, 148 degener Lutra, 152 delectorum Praomys, 119 deletrix Vulpes, 150 Delphinapterus leucas, 148 Delphinus melas, 148 demidovii Galago, 135 Dendromurinae, 117 desmarestianus Heteromys, 111 devius Oryzomys, 123 diadema Propithecus, 135 Dicrostonyx chionopaes, 113 exsul, 113 groenlandicus, 113 torquatus, 113 Didelphidae, 100 Didelphis marsupialis, 100 particeps, 100 pigra, 100 virginiana, 100 Didelphimorphia, 99 Diplomys labilis, 131 Dipodidae, 112 Dipodomys californicus, 111 ordii, 111 pallidulus, 111 palmeri, 111 Dipodops ordii, 111 palmeri, 111 dispar Sorex, 139 distincta Neotoma, 122 dorcas Damaliscus, 149 dorsalis Tamias, 109 dorsatum Erethizon, 130 Dremomys flavior, 104 pernyi, 104 senex, 104 drouhardi Microgale, 102 douglasii Tamiasciurus, 109 dupreanum Eidolon, 98 Echimyidae, 131 Echymipera australis, 101 kalabu, 100 rufescens, 101 Eidolon dupreanum, 98 sakalava, 98 elegans Spermophilus, 108 elongata Mephitis, 164 elucus Procyon, 151 Emballonuridae, 142 energumenos Mustela, 153 Putorius, 153 eEnixus Microtus, 115 Eonycteris glandifera, 140 spelaea, 140 Eothenomys aurora, 113 eva, 113 melanogaster, 113 mucronatus, 113 Epimys zappeyi, 119 Eptesicus darlingtoni, 145 phasma, 146 pumilus, 145 eremicus Hesperomys, 98 Peromyscus, 98, 126 Erethizon dorsatum, 130 picinus, 130 Erethizontidae, 130 erigens Hipposideros, 142 Erioryzomys monochromos, 128 erminea Mustela, 152, 153 Erophylla sezekorni, 144 syops, 144 erythraeus Callosciurus, 103 Euarctos americanus, 151 sornborgeri, 151 Eulipotyphla, 97 Eupleres goudotii, 155 major, 155 Eutamias minimus, 109 neglectus, 109 euxinus Apodemus, 117 eva Eothenomys, 113 Evotomys proteus, 113 exsputus Sigmodon, 127 173 174 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 exsul Dicrostonyx, 113 extimus Sciurus, 106 familiaris Canis, 150 fasciatus Cervaria, 155 Lynx, 155 fatuus Synaptomys, 116 Felidae, 155 Felis bangsi, 156 capensis, 155 concolor, 156 coryi, 156 costaricensis, 156 floridana, 156 improcera, 156 phillipsi, 155 serval, 155 femurvillosum Artibeus, 144 ferrugineus Callosciurus, 103 fervidus Sigmodon, 128 Fiber aquilonius, 116 obscurus, 116 rivalicius, 116 zibethicus, 116 fischeri Haplonycteris, 141 flavicans Oecomys, 122 Oryzomys, 122 flavidus Isthmomys, 121 Megadontomys, 121 flavior Dremomys, 104 flavipes Tatera, 117 flaviventer Arctomys, 105 flaviventris Marmota, 105 floridana Felis, 156 Neotoma, 122 floridanus Geomys, 110 Sylvilagus, 134 fontigenus Microtus, 115 fontoynonti Dasogale, 103 frenata Mustela, 152, 153, 154 frenatus Putorius, 152 fulvescens Oligoryzomys, 122 fulviventer Marmosa, 99 fumatus Myomys, 119 Praomys, 119 Funisciurus akka, 104 pyrrhopus, 104 victoriae, 104 furculus Triaenops, 143 furvus Sigmodon, 128 Urocyon, 150 fuscipes Neotoma, 98 fuscomurina Crocidura, 137 fusus Peromyscus, 126 Galago demidovii, 135 orinus, 135 Galagoides orinus, 135 Galagonidae, 135 Galidictis grandidiensis, 155 grandidieri, 155 gappert Clethrionomys, 113 geata Crocidura, 138 Myosorex, 138 Geocapromys abaconis, 132 columbianus, 132 cubanus, 132 ingrahami, 132 irrectus, 132 Geogale aurita, 101 orientalis, 101 Geomyidae, 110 Geomys austrinus, 110 colonus, 110 cumberlandius, 110 floridanus, 110 coffi, 110 pinetis, 110 tuza, 110 Gerbillinae, 117 gerbillus Leggada, 119 germanus Cerdocyon, 150 gibbsi Neurotrichus, 140 giganteus Pteropus, 141 gigas Lynx, 155 glaber Heterocephalus, 129 landifera Eonycteris, 140 Glaucomys bangsi, 104 lascivus, 104 makkovikensis, 104 querceti, 105 sabrinus, 104 volans, 104, 105 Glossophaga longirostris, 144 gloveralleni Procyon, 151 Sorex, 138 gofft Geomys, 110 gorgonae Proechimys, 131 gorilla Gorilla, 136 Troglodytes, 136 (ey fo) Gorilla gorilla, 136 gossypinus Hesperomys, 123 Peromyscus, 123, 124, 125 goudotii : Eupleres, 155 Grammomys macmillani, 118 granatensts Sciurus, 106, 107 grandidiensis Galidictis, 155 grandidieri Galidictis, 155 grandis Orthogeomys, 111 Graphiurus argenteus, 129 griseus, 129 hueti, 129 lorraineus, 129 microtis, 129 murinus, 129 schwabi, 129 surdus, 129 griseus Graphiurus, 129 griswoldi Tana, 134 Guerlinguetus aestuans, 106 chiriquensis, 106 guyannensis Proechimys, 131 gymnicus Sciurus, 109 Tamiasciurus, 109 haemobaphes Callosciurus, 103 Sciurus, 103 Haplonycteris fischeri, 141 hardyi Zapus, 112 harpia Harpiocephalus, 146 Harpiocephalus harpia, 146 rufulus, 146 harringtoni Taterillus, 117 helleri Platyrrhinus, 144 Herpestidae, 155 hesperia Lonchophylla, 144 Hesperomys arcticus, 124 eremicus, 98 gossypinus, 123 leucopus, 124 nebrascensis, 124 sonoriensis, 124 Heterocephalus glaber, 129 stygius, 129 Heteromyidae, 111 Heteromys alleni, 112 desmarestianus, 111 repens, 111 hildegardeae Crocidura, 138 Hipposideros bicolor, 142 curtus, 142 erigens, 142 turpis, 143 hirsutus Sigmodon, 128 hispida Suillomeles, 100 hispidus Sigmodon, 127, 128 Hominidae, 138 hoplites Dasypus, 101 hottentotus Cryptomys, 129 hudsonica Lutra, 152 TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden 175 hudsonicus Sciurus, 109 Tamiasciurus, 109 Zapus, 112 hueti Aethoglis, 129 Graphiurus, 129 humulis Reithrodontomys, 127 hyacinthus Neurotrichus, 140 hyleae Proechimys, 131 Hylobates carpenteri, 136 lar, 136 Hylobatidae, 136 Hylomyscus alleni, 118 simus, 118 Hylopetes alboniger, 105 anchises, 105 orinus, 105 phayrei, 105 hypoxanthus Oenomys, 119 Hypudaeus nivicola, 112 Ictidomys badius, 108 tridecemlineatus, 108 Idionycteris phyllotis, 146 ignava Marmota, 105 ignavus Arctomys, 105 ignifer Rhinolophus, 143 illectus Oecomys, 122 Oryzomys, 122 impiger Reithrodontomys, 127 improcera Felis, 156 Puma, 156 incitatus Lepus, 134 Sylvilagus, 134 Tapeti, 134 incomtus Dasymys, 118 Indridae, 135 ingrahami Geocapromys, 132 innuitus Mictomys, 116 Synaptomys, 116 insulanus Peromyscus, 125 insularis Pteropus, 98 irrectus Geocapromys, 132 irroratus Liomys, 112 Isthmomys flavidus, 121 kalabu Echymipera, 100 kempi Tatera, 117 kivuensis Thamnomys, 120 kodiacensis Spermophilus, 99 koreni Microtus, 115 Sorex, 138 labilis Diplomys, 131 Loncheres, 131 lacustris Otomys, 120 ladas Zapus, 112 laetus Aethosciurus, 106 Lagomorpha, 133 lambertoni Nesomys, 120 lanatus Caluromys, 99 laniger Cheiromys, 135 Pteropus, 96 lanigera Pteropus, 98 lar Hylobates, 136 lascivus Glaucomys, 104 Sciuropterus, 104 Lasiurus borealis, 146 brachyotis, 146 latimanus Scapanus, 140 laxatina Lontra, 142 lecontii Reithrodontomys, 127 Leggada ablutus, 118 bufo, 118 gerbillus, 119 Lemmus chrysogaster, 114 paulus, 114 sibiricus, 114 lepida Neotoma, 121 Leporidae, 143 leporina Dasyprocta, 130 Leptailurus phillipsi, 155 serval, 155 Lepus alacer, 134 alleni, 133 americanus, 133, 134 arcticus, 133 bairdi, 133, 134 bangsii, 133 cascadensis, 134 incitatus, 134 palitans, 133 paludicola, 134 struthopus, 133 sylvaticus, 134 transitionalis, 134 leucas Delphinapterus, 148 leucogaster Scotophilus, 147 leucopus Hesperomys, 124 Peromyscus, 126 leucostigma Chaerophon, 145 Mops, 145 levipes Melomys, 118 Liomys alleni, 112 irroratus, 112 Lipotyphla, 136 lituratus Artibeus, 144 Loncheres labilis, 131 Lonchophylla hesperia, 144 longicauda Arctogale, 153 Putorius, 153 longicaudata Microgale, 102 longimembis Perognathus, 112 longipilis Arvicola, 114 longirostris Glossophaga, 144 Lontra canadensis, 152 laxatina, 152 loquax Sciurus, 109 Tamiasciurus, 109 lorraineus Graphiurus, 129 lotor Procyon, 151 louisianae Odocoileus, 148 lucifugus Myotis, 146 luctuosa Alouatta, 135 ludovicianus Sciurus, 106 lupus Canis, 149, 150 lutensis Lutreola, 153 Mustela, 153 Putorius, 153 Lutra degener, 151 hudsonica, 151 vaga, 151 Lutreola lutensis, 153 vulgivagus, 154 Lynx californicus, 155 canadensis, 155, 156 fasciatus, 155 gigas, 155 oculeus, 155 rufus, 155 subsolanus, 156 mabirae Phataginus, 156 macmillani Grammomys, 118 Macrogeomys cavator, 110 pansa, 110 macropygmaeus Sorex, 138 macrotis Nycteris, 142 Peropteryx, 142 Macrotolagus alleni, 133 palitans, 133 macroura Odocoileus, 148 macrurus Sorex, 139 madagascariensis Cheiromys, 135 Daubentonia, 135 Nycteris, 142 Rousettus, 141 major Eupleres, 155 Vampyrodes, 145 makkovikensis 176 Glaucomys, 104 Sciuropterus, 104 maniculatus Peromyscus, 98, 124, 125 Manidae, 156 marcanot Antillogale, 137 Solenodon, 137 Marmosa fulviventer, 99 mitis, 99 robinsoni, 99, 100 Marmosidae, 99 Marmota avara, 105 flaviventris, 105 ignava, 105 monax, 105 marsupialis Didelphis, 100 Martes americana, 152 atrata, 152 brumalis, 152 maurisca Crocidura, 138 mayeri Antechinus, 100 maynardi Procyon, 151 mearnsi Saccostomus, 117 medialis Callosciurus, 103 medioximus Mictomys, 116 Synaptomys, 116 Megadontomys flavidus, 121 megaphyllus Rhinolophus, 143 melanogaster Eothenomys, TELS} melanops Taterillus, 117 melanotis Oryzomys, 123 melanotus Praomys, 119 melas Delphinus, 148 Myoictis, 100 Melomys alleni, 118 levipes, 118 mollis, 118 moncktoni, 118 rubex, 118 stevensi, 118 mephitica Mephitis, 154 Mephitidae, 154 mephitis Mephitis, 154 Mephitis avia, 154 elongata, 154 mephitica, 154 mephitis, 154 mesomelas, 154 scrutator, 154. spissigrada, 154 Mesocapromys nanus, 132 mesomelas Mephitis, 154 mexicana Neotoma, 122 mexicanus Peromyscus, 124 Microgale brevicaudata, 102 cowani, 102 decaryi, 102 drouhardi, 102 longicaudata, 102 parvula, 102 principula, 102 prolixicaudata, 102 pulla, 102 Microsciurus alfari, 105 browni, 105 microtis Graphiurus, 129 Microtus acadicus, 115 aurora, 113 breweri, 114 chrotorrhinus, 114 enixus, 115 fontigenus, 115 koreni, 115 mucronatus, 113 oeconomus, 115 pennsylvanicus, 114, 115 provectus, 115 ravus, 114 shattucki, 115 terraenovae, 114 micrus Nesophontes, 136 Mictomys innuitus, 116 medioximus, 116 MiNntMus Tamias, 99, 109 minor Ametrida, 143 Suncus, 139 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 minusculus Scapanus, 140 miscix Sorex, 139 misim Antechinus, 100 mitis Marmosa, 99 mollipilosus Tamiasciurus, 109 mollis Melomys, 118 Stenomys, 120 Molossidae, 145 monax Marmota, 105 moncktoni Melomys, 118 monochromos Erioryzomys, 128 Oryzomys, 125 Thomasomys, 128 Monodontidae, 148 monoensis Pteropus, 141 Mops angolensis, 145 condylurus, 145 leucostigma, 145 orientis, 145 Mormoopidae, 143 mortigena Mustela, 152 morulus Sciurus, 107 mucronatus Microtus, 113 munda Mustela, 154 mundus Putorius, 154 muricola Myotis, 146 MUPICUS Arctogale, 153 Mustela, 153 Putorius, 153 Muridae, 112 Murinae, 117 MUrinUs Graphiurus, 129 Pseudohydromys, 120 Mus bactrianus, 119 bairdii, 98 bufo, 118 musculus, 119 tantillus, 119 tenellus, 119 musculus Mus, 119 Mustela atrata, 152 brumalis, 152 cicognanii, 152 energumenos, 153 erminea, 152, 153 frenata, 152, 153, 154 lutensis, 153 mortigena, 152 munda, 154 muricus, 153 neomexicana, 152 nivalis, 153 noveboracensis, 153 occisor, 153 oribasus, 153 richardsonii, 152 rixosa, 153 vison, 153, 154 vulgivaga, 154 Mustelidae, 152 Myoictis melas, 100 wallacei, 100 wavicus, 100 Myomys fumatus, 119 Myopus schisticolor, 115 thayeri, 115 Myosorex geata, 138 Myotis abbotti, 146 albicinctus, 146 carissima, 146 lucifugus, 146 muricola, 146 nugax, 146 sodalis, 146 Myoxidae, 129 mystacinus Apodemus, 117 nana Capromys, 132 Nycteris, 142 nanus Mesocapromys, 132 naso Antechinus, 100 Rhynchiscus, 142 natator Oryzomys, 123 navus Oryzomys, 122 nebrascensis Hesperomys, 124 Peromyscus, 124 Nectomys amazonicus, 121 squamipes, 121 neglectus Tamias, 109 Neomeris asiaeorientalis, 148 neomexicana Mustela, 152 neomexicanus Putorius, 152 Neophocaena asiaorientalis, 148 phocaenoides, 148 Neosorex acadicus, 138 palustris, 138 Neotoma abbreviata, 121 bella, 121 distincta, 122 floridana, 122 fuscipes, 98 lepida, 121 mexicana, 122 rubida, 122 nesaeus Sciurus, 107 Nesomyinae, 120 Nesomys lambertoni, 120 Nesophontes micrus, 136 Nesophontidae, 136 Neurotrichus gibbsi, 140 hyacinthinus, 140 niger Sciurus, 106 nigriculus Peromyscus, 125 niobe Stenomys, 120 nitellinus Nyctomys, 122 nivalis Chionomys, 112 Mustela, 153 nivicola Hypudaeus, 112 Niviventer confucianus, 119 noblei Dasyprocta, 130 notius Pipistrellus, 147 Putorius, 153 Scabrifer, 147 noveboracensis Mustela, 153 Putorius, 153 novemcinctus Dasypus, 101 nuchalis Dasyprocta, 130 nugax Myotis, 146 nyasae Steatomys, 117 Nycteridae, 142 Nycteris macrotis, 142 madagascariensis, 142 nana, 142 revoili, 97 tristis, 142 Nycticeius africanus, 147 albiventer, 147 schlieffeni, 147 Nyctomys nitellinus, 122 sumichrasti, 122 obscurus Citellus, 107 Fiber, 115 Ondatra, 115 obsoletus Spermophilus, 99 occidentalis Paramicrogale, 102 occisor Mustela, 153 Putorius, 153 occlusus Cryptomys, 129 Ochotona cupppes, 133 princeps, 133 saxatilis, 133 Ochotonidae, 133 ochraceus Thamnomys, 118 Octodontidae, 130 Octomys barrerae, 130 oculeus Cervaria, 155 Lynx, 155 ocythous Urocyon, 150 Odocoileus americanus, 148 borealis, 148 clavium, 149 louisianae, 146 macroura, 146 osceola, 146 virginianus, 146, 147 Oecomys bicolor, 122 flavicans, 122 illectus, 122 trabeatus, 122 oeconomus TYPE SPECIMENS OF RECENT MAMMALS ® Helgen and McFadden 1 Microtus, 115 Oenomys hypoxanthus, 119 talangae, 119 Oligoryzomys fulvescens, 122 vegetus, 125 Ondatra aquilonius, 116 obscurus, 116 rivalicius, 116 zibethicus, 116 orarius Sciurus, 109 Zapus, 112 ordii Dipodomys, 111 Dipodops, 111 oreas Peromyscus, 126 oribasus Arctogale, 153 Mustela, 153 Putorius, 153 orientalis Geogale, 101 orientis Mops, 145 orinus Galago, 135 Galagoides, 135 Hylopetes, 105 Pteromys, 105 Orthogeomys cavator, 110 grandis, 111 pansa, 110 pluto, 111 Oryzomys carrorum, 123 coloratus, 123 devius, 123 flavicans, 122 fulvescens, 122 illectus, 122 melanotis, 123 monochromos, 128 natator, 123 navus, 122 palustris, 123 rostratus, 123 vegetus, 123 osceola Cariacus, 148 Odocoileus, 146 Otomops papuensis, 145 Otomyinae, 120 Otomys anchietae, 120 barbouri, 121 ~l ~ lacustris, 120 typus, 121 uzungwensis, 121 owent Praomys, 119 paca Agouti, 130 Pachycyon robustus, 150 paitana Tupaia, 134 palitans Lepus, 133 Macrotolagus, 133 palliata Alouatta, 135 pallidulus Dipodomys, 111 pallidus Spermophilus, 99 Tamias, 99 palmarius Peromyscus, 125 palmarum Artibeus, 144 palmeri Dipodomys, 111 Dipodops, 111 paludicola Lepus, 134 Sylvilagus, 134 palustris Neosorex, 138 Oryzomys, 123 Sorex, 138 Sylvilagus, 134 pansa Macrogeomys, 110 Orthogeomys, 110 Pappogeomys castanops, 111 rubellus, 111 papuensis Otomops, 145 Paracapromys, 132 Paramicrogale occidentalis, 102 Paraxerus byatti, 106 vexillarius, 106 parnellii Chilonycteris, 143 Pteronotus, 143 parryi Spermophilus, 99 particeps Didelphis, 100 parvula Microgale, 102 paulus Lemmus, 114 178 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 Pecari crusnigrum, 147 tajacu, 147, 148 torvus, 148 Pedetes cafer, 125 capensis, 128 taborae, 128 Pedetidae, 128 pennsylvanica Vulpes, 150, 151 pennsylvanicus Microtus, 114, 115 Peramelia, 100 pernyt Dremomys, 104 Perognathus bangsi, 112 longimembris, 112 Peromyscus abietorum, 125 ammodytes, 125 anastasae, 124, 125 arenarius, 126 argentatus, 125 attwateri, 124 bairdii, 98 baliolus, 126 bellus, 124 borealis, 124 cacabatus, 124 canadensis, 125 crinitus, 125 eremicus, 98, 126 fusus, 126 gossypinus, 2S 2Ae 125 insulanus, 125 leucopus, 126 maniculatus, 98, 124, 125 mexicanus, 124 nebrascensis, 124 nigriculus, 125 oreas, 126 palmarius, 125 phasma, 126 polionotus, 126 rhoadsi, 126 saturatus, 126 scitulus, 125 subgriseus, 126 texanus, 126 Peropteryx canina, 142 macrotis, 142 phaea, 142 Peroryctidae, 100 perriert Propithecus, 135 personatus Sorex, 139 phaea Peropteryx, 142 phaios Crocidura, 138 phasma Eptesicus, 147 Peromyscus, 126 Pipistrellus, 147 Phataginus mabirae, 156 tricuspis, 156 phayrei Hylopetes, 105 Pteromys, 105 Phenacomys celatus, 116 crassus, 116 ungava, 116 Philander cicur, 99 philippinensis Rhinolophus, 143 phillipsi Damaliscus, 149 Felis, 155 Leptailurus, 155 phocaenoides Neophocaena, 148 Phocoenidae, 148 Phyllostomidae, 143 phyllotis Corynorhinus, 146 Idionycteris, 146 picinus Erethizon, 130 pigra Alouatta, 135 Didelphis, 100 piloroides Capromys, 131 pinetis Geomys, 110 Pipistrellus capensis, 147 notius, 147 phasma, 147 rendalli, 147 Plagiodonta araeum, 132 Platyrrhinus helleri, 144 umbratus, 144 Plecotus auritus, 147 sacrimontis, 147 pluto Orthogeomys, 111 poeyanus Solenodon, 137 polionotus Peromyscus, 126 Praomys delectorum, 119 fumatus, 119 melanotus, 119 oweni, 119 tullbergi, 119 pratensis Steatomys, 117 Primates, 135 primus Callosciurus, 103 princeps Ochotona, 133 principula Microgale, 102 priscus Rhynchiscus, 142 Procyon elucus, 151 gloveralleni, 151 lotor, 151 maynardi, 151 Procyonidae, 151 Proechimys burrus, 131 cayennensis, 131 gorgonae, 131 guyannensis, 131 hyleae, 131 semispinosus, 131 prolixicaudata Microgale, 102 Propithecus diadema, 135 perrieri, 135 proteus Clethrionomys, 113 Evotomys, 113 provectus Microtus, 115 Pseudohydromys murinus, 120 Pteromys alboniger, 105 anchises, 105 orinus, 105 phayrei, 105 Pteronotus parnellii, 143 pusillus, 143 quadridens, 143 Pteropodidae, 140 Pteropus anetianus, 141 aorensis, 141 ariel, 141 austini, 141 giganteus, 141 insularis, 98 laniger, 98 lanigera, 98 monoensis, 141 rayneri, 141 woodfordi, 141 pulla Microgale, 102 Puma bangsi, 156 concolor, 156 coryi, 156 costaricensis, 156 improcera, 156 punctata Dasyprocta, 130 pusillus Chilonycteris, 143 Pteronotus, 143 putorius Spilogale, 155 Putorius energumenos, 153 frenatus, 152 longicauda, 153 lutensis, 153 mundus, 154 muricus, 153 neomexicanus, 153 notius, 153 noveboracensis, 153 occisor, 153 oribasus, 153 rixosus, 153 vison, 153 vulgivagus, 154 xanthogenys, 154 pygargus Damaliscus, 149 pyrrhopus Funisciurus, 104 quadridens Pteronotus, 143 quadrivittatus Tamias, 99, 109 querceti Glaucomys, 105 Sciuropterus, 105 rainieri Aplodontia, 103 Rangifer arcticus, 149 caboti, 149 caribou, 149 tarandus, 149 terraenovae, 149 ravus Microtus, 114 raynert Pteropus, 141 Reithrodontomys australis, 127 creper, 127 TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden 179 humulis, 127 impiger, 127 lecontii, 127 sumichrasti, 127 vulcanius, 127 relictus Capromys, 131 rendalli Pipistrellus, 147 repens Heteromys, 111 revoili Nycteris, 97 Rhinolophidae, 142 Rhinolophus alleni, 143 ignifer, 143 megaphyllus, 143 philippinensis, 143 rhoadsi Peromyscus, 126 Rhynchiscus naso, 142 priscus, 142 Rhynchonycteris naso, 142 richardsonii Mustela, 152 riparia Arvicola, 114 rivalicius Fiber, 116 Ondatra, 116 rixosa Mustela, 153 rixOSUS Putorius, 153 robinsoni Marmosa, 99, 100 roboratus Sorex, 139 robustus Pachycyon, 150 Rodentia, 103 rostratus Oryzomys, 123 Rousettus madagascariensis, 141 rubellus Cratogeomys, 111 Pappogeomys, 111 rubex Melomys, 118 rubida Neotoma, 122 rubricatus Vulpes, 151 rubricosa Vulpes, 150, 151 rufa Aplodontia, 103 rufescens Echymipera, 101 rufidorsum Arvicola, 114 rufulus Harpiocephalus, 145 rufus Lynx, 155 sabrinus Glaucomys, 104 Sciuropterus, 104 Saccostomus cricetulus, 117 mearnsi, 117 sacrimontis Plecotus, 147 sakalava Eidolon, 98 saltuensis Sciurus, 107 sanctaemartae Sigmodon, 128 sanguinidens Sorex, 139 saturatus Peromyscus, 126 saxatilis Ochotona, 133 Scabrifer notius, 147 Scalops aereus, 140 anastasae, 140 texanus, 140 Scalopus aereus, 140 anastasae, 140 aquaticus, 140 Scandentia, 134 Scapanus californicus, 140 latimanus, 140 minusculus, 140 schisticolor Myopus, 115 schlieffeni Nycticeius, 147 schwabi Graphiurus, 129 scitulus Peromyscus, 125 Sciuridae, 103 Sciuropterus alpinus, 104 bangsi, 104 lascivus, 104 makkovikensis, 104 querceti, 105 sabrinus, 104 silus, 104 volans, 104, 105 Sciurotamias consobrinus, 106 davidianus, 106 thayeri, 106 Sciurus aberti, 99 aestuans, 106 browni, 105 carolinensis, 106 castaneoventris, 103 castanonotus, 99 castanotus, 99 chiriquensis, 106 extimus, 106 granatensis, 106, 107 gymnicus, 109 haemobaphes, 103 hudsonicus, 109 loquax, 109 ludovicianus, 106 morulus, 107 nesaeus, LOT niger, 106 orarius, 109 saltuensis, 107 variabilis, 107 vicinus, 106 vulpinus, 106 yucatanensis, 106 Scotinomys apricus, 127 teguina, 127 xerampelinus, IAG Scotophilus altilis, 147 leucogaster, 147 scrutator Mephitis, 154 semispinosus Proechimys, 131 senex Dremomys, 104 seorsus Zygodontomys, 128 serval Felis, 155 Leptailurus, 155 Setifer setosus, 103 setosus Setifer, 103 sezekorni Erophylla, 144 shattucki Microtus, 115 sibiricus Lemmus, 114 SICCUS Citellus, 107 Sigmodon austerulus, 127 borucae, 127 exsputus, 127 fervidus, 128 furvus, 128 hirsutus, 128 hispidus, 127, 128 sanctaemartae, 128 spadicipygus, 128 Sigmodontinae, 121 silus Sciuropterus, 104 simus Hylomyscus, 118 sodalis Myotis, 146 soleatus Claviglis, 129 Solenodon cubanus, 137 marcanoi, 137 poeyanus, 137 Solenodontidae, 137 sonoriensis Hesperomys, 124 Sorex araneus, 138 caecutiens, 138 cinereus, 139 daphaenodon, 139 dispar, 139 gloveralleni, 138 koreni, 138 macropygmaeus, 138 macrurus, 139 miscix, 139 palustris, 138 personatus, 139 roboratus, 139 sanguinidens, 139 tundrensis, 138 ultimus, 138 vir, 139 Soricidae, 137 sornborgeri Euarctos, 151 Ursus, 151 sOoror Tatera, 117 spadicipygus Sigmodon, 128 spelaea Eonycteris, 140 Spermophilus alashanicus, 107 armatus, 107 badius, 108 elegans, 108 kodiacensis, 99 obsoletus, 99 pallidus, 99 parryi, 99 180 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 spilosoma, 99 texensis, 108 tridecemlineatus, 99, 108 Spilogale ambarvalis, 155 putorius, 155 spissigrada Mephitis, 154 squamipes Nectomys, 121 Steatomys nyasae, 117 pratensis, 117 Stenomys mollis, 120 niobe, 120 stevensi, 120 verecundus, 120 stevensi Melomys, 118 Stenomys, 120 striatus Tamias, 107 struthopus Lepus, 133 stuhlmanni Chrysochloris, 101 stygius Heterocephalus, 129 subgriseus Peromyscus, 126 subsolanus Lynx, 156 Suillomeles hispida, 100 sumichrasti Nyctomys, 122 Reithrodontomys, 127 Suncus ater, 139 minor, 139 varilla, 139 surdus Graphiurus, 129 sylvaticus Lepus, 134 Sylvilagus alacer, 134 brasiliensis, 134 floridanus, 134 incitatus, 134 paludicola, 134 palustris, 134 transitionalis, 134 Synaptomys ~ borealis, 116 cooperi, 116 fatuus, 116 innuitus, 116 medioximus, 116 Syntheosciurus brochus, 108 syops Erophylla, 144 taborae Pedetes, 128 tajacu Pecari, 147, 148 talangae Oenomys, 119 Talpidae, 140 Tamias borealis, 109 cooperi, 108 dorsalis, 108 minimus, 99, 109 neglectus, 109 pallidus, 99 quadrivittatus, 99 striatus, 109 townsendii, 108 venustus, LOY Tamiasciurus douglasii, 109 gymnicus, 109 hudsonicus, 109 loquax, 109 mollipilosus, 109 Tana griswoldi, 134 tana, 134 tana Tana, 134 Tupaia, 134 tantillus Mus, 119 Tapeti incitatus, 134 tarandus Rangifer, 149 Tatera flavipes, 117 kempi, 117 soror, 117 valida, 117 Taterillus harringtoni, 117 melanops, 117 Tayassu crusnigrum, 147 torvus, 148 Tayassuidae, 147 teguina Akodon, 127 Scotinomys, 127 tenellus Mus, 119 Tenrecidae, 101 tephragaster Crocidura, 137 terraenovae Arvicola, 114 Microtus, 114 Rangifer, 149 texanus Peromyscus, 126 Scalops, 140 texensis Spermophilus, 108 Thamnomys kivuensis, 120 ochraceus, 118 venustus, 120 thayeri Myopus, 115 Sciurotamias, 106 thomasi Cryptotis, 138 Thomasomys monochromos, 128 Thomomys atrodorsalis, 111 umbrinus, 111 thous Cerdocyon, 150 Thyroptera albigula, 147 albiventer, 147 tricolor, 147 Thyropteridae, 147 torquatus Dicrostonyx, 113 torrei Boromys, 131 Chilonycteris, 143 torvus Pecari, 148 Tayassu, 148 townsendii Tamias, 109 trabeatus Alouatta, 135 Oecomys, 122 transitionalis Lepus, 134 Sylvilagus, 134 Triaenops aurita, 143 furculus, 143 tricolor Thyroptera, 147 tricuspis Phataginus, 156 tridecemlineatus Spermophilus, 99, 108 trinotatus Zapus, 112 tristis Nycteris, 142 Troglodytes gorilla, 136 tropicalis Chlorotalpa, 101 Chrysochloris, 101 tullbergi Praomys, 119 tundrensis Sorex, 138 Tupaia paitana, 134 tana, 134 Tupaiidae, 134 turpis Hipposideros, 143 tuza Geomys, 110 Tympanoctomys barrerae, 130 typus Otomys, 121 ultimus Sorex, 138 umbratus Platyrrhinus, 144 Vampyrops, 144 umbrinus Thomomys, 111 ungava Phenacomys, 116 Urocyon aquilus, 150 cinereoargenteus, 150 furvus, 150 ocythous, 150 Ursidae, 151 Ursus americanus, 151 sornborgeri, 151 UZUNZWENSIS Otomys, 121 vafra Vulpes, 150, 151 vaga Lutra, 152 valida Tatera, 117 Vampyrodes caraccioli, 145 major, 145 Vampyrops umbratus, 144 zarhinus, 144 variabilis Sciurus, 107 varilla Suncus, 139 vegetus Oligoryzomys, 123 Oryzomys, 123 venustus Tamias, 109 Thamnomys, 120 verecundus Stenomys, 120 Vespertilionidae, 145 vexillarius Paraxerus, 106 vicinus Sciurus, 106 victoriae Funisciurus, 104 vir Sorex, 139 virgatus Agouti, 130 virginiana Didelphis, 100 virginianus Odocoileus, 148, 149 vison Mustela, 153, 154 Putorius, 153 Viverridae, 155 volans Glaucomys, 104, 105 Sciuropterus, 104, 105 vulcanius Reithrodontomys, 127 vulgivaga Mustela, 154 vulgivagus Lutreola, 154 Putorius, 154 vulpes Vulpes, 150, 151 Vulpes bangsi, 151 deletrix, 150 pennsylvanica, 150, 151 rubricatus, 151 rubricosa, 150, 151 vafra, 150, 151 vulpes, 150, 151 vulpinus Sciurus, 106 wallacei Myoictis, 100 wavicus Myoictis, 100 whytei Cryptomys, 129 woodfordi Pteropus, 141 xanthogenys Putorius, 154 xerampelinus Akodon, 127 Scotinomys, 127 yucatanensis Sciurus, 106 TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden zappeyt Epimys, 119 Zapus acadicus, 112 hardyi, 112 hudsonicus, 112 ladas, 112 orarius, 112 trinotatus, 112 zarhinus Vampyrops, 144 zibethicus Fiber, 116 Ondatra, 116 Zygodontomys brevicauda, 128 cherriei, 128 seorsus, 128 181 oe j ee SM 4 5) lie (mi e a i = i = - 2 : } - aie ae E > = 4 A = : Te SS = « 7 : el te en 7 en Se -- : eat - ; 2 ne - oN : : : iy 7 _ : = —— a : - = o\: oa a oa o = pre 5 ; Lis a ei s-s i a — hee ia moe Uy é US ISSN 0027-4100) _ A New Species of Lizard Related to _ Stenocercus caducus (Cope) (Squamata: Iguanidae) From Peru and Bolivia, With a Key to the “Ophryoessoides Group” JOHN E. CADLE VOLUME 157, NUMBER 3 AMBRIDGE, MASSACHUSETTS, U.S.A. 14 November 2001 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952— BULLETIN 1863— Memorrs 1865-1938 JouNsoni4, Department of Mollusks, 1941-1974 OccASIONAL PAPERS ON Mo tusks, 1945— SPECIAL 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, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4, Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Emest 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 of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the 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 of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1947. (Complete sets only.) Proceedings of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock, © The President and Fellows of Harvard College 2001. A NEW SPECIES OF LIZARD RELATED TO STENOCERCUS CADUCUS (COPE) (SQUAMATA: IGUANIDAE) FROM PERU AND BOLIVIA, WITH A KEY TO THE “OPHRYOESSOIDES GROUP” JOHN E. CADLE' CONTENTS ENTERS Sa OPT ee tay ee 183 [Reganrrayeran’ 2 oki eicteie el ae ee es 183 Mien @ GLU CHOI yen tek PLUM Pes REE TUE a Re 184 Materials and Methods 000 184 Description of a New Species of Stenocersus 187 Stenocercus prionotus new species —.... 187 Diagnosis and Comparisons 2 191 BBY SS CIs tol Opes ee eee ee as sees oe 193 Distribution Patterns in Stenocercus prionotus ces OE EE aed ae ad oe Lie ee 197 Patterns of Sympatry and Geographic Variation in Stenocercus prionotus, and the Need for Additional Fieldwork 200 Natural History of Stenocercus prionotus 205 Comparison of Stenocercus prionotus with S. COCULCUS Se eke MERE A Ni ee 2 ee Nea RE 206 Distributions of Stenocercus prionotus and S. caducus in Eastern Bolivia ___.................. 210 Is the Distribution of Stenocercus aculeatus GHGHTONIVGI Ci a5 Reet pee 270s Meee cee ee ee 212 Key to Species of the “Ophryoessoides group” Git SHOMOCEROUS Did} Ac noyylecearncints: 3 ae ee 215 Mppendix: Specimens Examined 0). 216 ihiterature Cited 228. 2 218 ABSTRACT. A new species of iguanid lizard, Steno- cercus prionotus, is described Gorn eastern Peru and Bolivia (known range from San Martin Department, Peru, to northern La Paz and El Beni departments, Bolivia). Most localities are in the Andean foothills and immediately adjacent lowlands. Stenocercus prionotus is similar to several other species of Sten- ocercus with large posterior head scales, an enlarged row of supraoculars, and keeled ventral scales. These similar species are referred to as the “Ophryoessoides group” without implying that it is a monophyletic as- semblage. Based on their common possession of a ' Department of Herpetology, Chicago Zoological Society, 3300 Golf Road, Brookfield, iiliots 60513. Research Associate, Department of Herpetology, Museum of Comparative Zoology. Bull. Mus. Comp. Zool., unique scaly flap concealing a portion of the posthu- meral mite pocket, the new species is apparently closely related to S. caducus (Cope), which is known from central Bolivia south to northern Argentina and Paraguay. Stenocercus prionotus is distinguished from S. caducus by having a more prominent vertebral crest and a pattern of alternating light and dark bars on the throat (rarely observed in ct caducus, which usually has light throat spots). These two species also occupy different physiographic regions (western Am- azonian rainforest for S. prionotus; chaco for S. cad- ucus). Populations of S. prionotus from northern Peru have a higher vertebral crest than those in southern Peru and Bolivia. Northern populations are also broadly sympatric with two other species of the “Ophryoessoides group,” S. aculeatus and S. fimbria- tus. However, in southern Peru S. prionotus is not known to be sympatric with other species of that group. I postulate that the higher vertebral crest in northern populations of S. prionotus functions as a species recognition signal in the multispecies assem- blages. A key to species of the Ophryoessoides group” is provided and distributions of the species in Peru and Bolivia are summarized. RESUMEN. Se describe una nueva especie de lagar- tija iguanida, Stenocercus prionotus, del Pert Orien- tal y ob Bolivia. Se conoce la nueva especie desde el departamento de San Martin, Pert, hasta el norte de los departamentos La Paz y El Beni en Bolivia. La mayoria de las localidades se encuentran en las estri- baciones andinas y adyacentes tierras bajas. Stenocer- cus prionotus es similar a varias otras especies de Stenocercus con grandes escamas sobre el posterior de la cabeza, una fila amplia de supraoculares, y es- camas ventrales quilladas. Se refiere estas especies como el “grupo Ophryoessoides,” sin implicar su monophyletismo. Basada en su posesion de un l6bulo escamoso tinico que oculta una porcidén del bolsillo antehumeral, se considera la nueva especie cercana- mente relacionada a Stenocercus caducus (Cope), que se conoce desde Bolivia central hasta el norte de la 157(3): 183-222, November, 2001 183 154 Argentina y del Paraguay. Stenocercus prionotus se distingue de S. caducus al tener una cresta vertebral mas prominente y un patron de alternativas barras claras y oscuras sobre la garganta (un patron rara- mente observado en S. caducus, que usualmente ti- ene manchas claras sobre la garganta). Las poblaciones de Stenocercus prionotus del norte del Pert tienen una cresta vertebral mas alta que la cresta en poblaciones del sur de Perti y Bolivia. Las poblaciones del norte de Pert también son amplia- mente simpatricas con dos otras especies del “grupo Ophryoessoides,” S. aculeatus y S. fimbriatus. Sin em- bargo, en el sur del Peru y Bolivia no se conocen lo- calidades donde se encuentra S. prionotus simpatrica con otras especies del grupo. Postulo que la cresta ver- tebral mas alto en las poblaciones nortefias de S. prion- otus functiona como un sefial para reconocimiento de especies en comunidades donde hay varias especies simpatricas. Se provee una clave para las especies del “grupo Ophryoessoides” y se resumen las distribu- ciones de las especies Peruanas y Bolivianas. INTRODUCTION Stenocercus sensu Frost (1992), includ- ing the nominal genera Ophryoessoides and Proctotretus, isa “moderately diverse assem- blage of South American iguanid lizards (eos Macey et al., 1997; Soule et al., 1998) with about 50 species currently rec- ognized. Most of the species are in the An- des and adjacent lowlands of Colombia, Ec- uador, and Peru, but a few are primarily Amazonian or have distributions in the physiographically diverse terrain south of the Amazon basin. Although new species of Stenocercus continue to be discovered in the field, others have been known from old collections and are only now being de- scribed (e.g., Cadle, 1991, 1998; Avila-Pi- res, 1995). In this category is a new species from the lowlands of eastern Peru and Bo- livia that has been referred erroneously to the names aculeatus O'Shaughnessy (1879) or caducus Cope (1862) in previous litera- ture, and associated with the genera Lei- ocephalus or Ophryoessoides ‘before the current understanding of these names came into use (see Etheridge, 1966; Frost, 1992). Rodriguez and Gadlet (1990) left the new species cnameless in a checklist pend- ing resolution of its status. The new species is ‘apparently closely related to Stenocercus caducus (Cope) and is described herein. Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 MATERIALS AND METHODS Frost (1992; see especially footnote 5) and Cadle (1991) discussed reasons for re- ferring new species such as the one de- scribed here to Stenocercus Duméril and Bibron sensu lato (including Ophryoesso- ides Duméril and Proctotretus Duméril and Bibron). Externally, the new species is similar to those species that Fritts (1974) placed in Ophryoessoides, that is, those species with keeled ventral scales, large posterior head scales (usually including well-differentiated interparietal, parietals, postparietals, and occipitals), and one moderately to greatly enlarged supraocular row. In addition to the new species de- scribed here. species included in the “Ophryoessoides group” are aculeatus O'Shaughnessy, 1879; caducus Cope, 1862; dumerilii Steindachner, 1867; ery- throgaster Hallowell, 1856; fimbriatus Avi- la-Pires, 1995: huancabambae Cadle, 1991: iridescens Giinther, 1859; limitaris Cadle, 1998; scapularis Boulenger, 1901; tricristatus Duméril, 1851; ee two un- described species noted later in this paper under Key to Species of the “Ophryoesso- ides group” of Stenocercus. I use the term “Ophryoessoides group” as a convenience to refer to this group of phenotypically similar species without implication as to its status as a monophyletic or nonmonophy- letic assemblage within Stenocercus. General descriptive protocols follow Ca- dle (1991), who defined terminology of the scales, neck folds, and mite pockets used herein, based in part on Frost (1992). Bi- lateral scale counts (e.g., subdigitals) were made only on one side (the left, unless it was damaged), except for the holotype, for which both left and right counts were re- corded (1, r). A summary of selected scu- tellational and qualitative characters for the new species and similar species from eastern Peru and Bolivia is presented in Mable ws All measurements are in millimeters. The abbreviation SVL refers to the head— body length, from snout to vent. 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IG=N (Assouysneys 0) SNYDAINID SNILIDOUIYS q addy, dey ou - od4y, (=) 16 (SOG) FL (8) OLO'O = 990 19°0-¥9'0 Pal a=ataeG 9S-GE 690 + T9T SI-F1 (9 10 G Ajo.1e.1) P P Ajoret “(%LT) L (%6G) S “(%0F) 9 66S + L0G Go-LI Oo se My GS-LE SSL + SEF TS-66 q addy, dey Ayeos tpiM F odAy, (6FS) £6 2( +861) GL (13) SIO';O + OL'0 €Z'0-69'0 69 L + 0'9G OS-EE OTT + OLT 1é-ST (P Apote.t) 9-G (%GP) 9 (%GG) L (OF) IT + OST €Z-91 (IF) 69'S + BSE SF-08 (GAO Fe GIS FE-PE cp = N (adoy) S71 q ody, dey Apeos uM F odAy, (6GE) 6 (ESE) 68 (O€) 100 + GL'0 VL0-69 0 VIE + 996 TE VG OT + SSI IG-9I (2 Ajeet) Q A][BUOISed30 *G 8 Aertel “(% TT) S “(%GS) 9 “(%09) L 693 + GST 6G-SI 69'S + SFE 68-16 O'S ¥ SOF S95 coc =>N sotoods mou SNIAIDIOUITS yoypod pesoutoysog yoyood eroumyysog So[RULO Y sole (suey 7eI0}) "TAS “(UIUL) OZIS UINUIIXe (YIBuUsy [eIOV/[LEL, s[PUSIpqns 90} yVNoy s[eqsip -qns 1osuy YyINo4 savjnooridng s[eSRULLOyUT sopeos rBpns) Sol[BOS [PIQoqloA, sopeos ApoqpryN ‘(¢LONAISIG SALVATNOV SQOUAOONALS JO NOLLAIIHISIG: AHL ST AAS) SHALOVAVHO AALLVLLLNVAO ANV AALLVLITVAO HINOS NI YAAATG HOTHAA “SNAWIOddS NVIANYAd GNV NVINOGVNOW HLOA AGN IONI SALVATAOV “§ LOA VLVC (1661) HIGVD WOW AYV AVINVEAVONVOH ‘§ HOA VIVE (ATIVOILLHHLNAYVd GHAddV N) SAYNASVAW TVACIAIGNI HOA GAIMVA ATHL AWAHAA LdAOXA NWATOO HOVA AO dOL AHL LY NAAID AV (N) SHZIS ATANVS ‘NAAIO SI “SNVAW NVHL YAHLVY “ALVLS YALOVUVHO UVINOLLYVd V HLIM SNAWIONdS AO ADVINAOUAd YO AGOW AHL (SHUVTNOOVUdNS CNV STVSVNYALNI “D°H) SHHLOVUVHO HINOS HON “GS T + NVAW AP GAMOTION ADNVY SV NAAID AUV SYALOVYVHO AALLVLILNVAQ ‘VIAIIOG GNV NYAd NYALSVA NI (FLET “SLLIYA ASNas «SAAIOSSAOKUHAQ,,) SATVOS IWYLNAA GATAAN HLIM SQOUAOONALS AO SAIOUdS AHL AO NOSINVdNOD ‘T ATIAVL 186 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 S. prionotus S. aculeatus S. fimbriatus S. scapularis S. caducus 200 400 600 800 1000km Figure 1. Distribution of species of Stenocercus emphasized in this paper (western South America, Ecuador to Paraguay and northern Argentina). Open symbols for S. fimbriatus are literature records from Avila-Pires (1995); locality for S. caducus in northern Argentina is the southernmost locality in Argentina reported by Cei (1993). Otherwise, all records are based on spec- imens examined. Numbered localities are documented or suspected cases of sympatry referred to in the text and noted in the Appendix: 1, Pampa Hermosa; 2, Pampa Seca, Rio Mixiollo; 3, Tingo Maria; 4, Manu National Park. Upper left quadrant outlined with dotted line is the area shown in greater detail in Figure 2. figurations of neck folds and mite pockets vary considerably among species of Sten- ocercus and are useful in distinguishing species. The most important qualitative characteristics of these features used here- in are the following, which are discussed more fully by Grails (1991): Neck and Body Folds and Crests. In contrast to many species of Stenocercus, neck folds are usually absent or weakly de- soides group’ veloped in the “Ophryoessoides group.” When present, they are better character- ized as crests rather than folds because they are usually indicated by strongly keeled rows of scales instead of actual folds of skin. The position of such crests corresponds to the position of folds seen in other species of Stenocercus, but only two are commonly seen in the “Ophryoes- - an antehumeral crest, which NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle is a more or less vertical row of strongly keeled scales immediately anterior to the forelimb insertion, and usually highlighted with white scales; and a supra-auricular crest, a strongly keeled horizontal row of scales extending from the posterior tem- poral region to the shoulder region. All species of the “Ophryoessoides group” have a distinct vertebral crest formed by the strongly keeled, and often projecting, scales of the vertebral row. In addition, some of these species have a more or less prominent dorsolateral crest formed by a row of strongly keeled scales separating the flanks from the dorsum proper. The dorsolateral crest varies in length. In some species it is exceptionally long, extending from the proximal portion of the tail to fie neck region, where it is confluent with the supra-auricular crest. In other species it is present only anteriorly or posteriorly. Posthumeral (Axillary) and Postfemoral Mite Pockets. Type 1—pocket absent; no skin modification. Type 2—rudimentary pocket manifested by skin modification, such as bare skin, a series of wrinkles, or a shallow depression lined with scales dif- ferent from surrounding body scales. Type 3—similar to Type 2, but with an over- hanging fold of skin or a thickened border. Type 4—a deep pocket, usually with a broad circular opening, whose depth is greater than half the diameter of its open- ing. Type 5—a deep pocket with a narrow, slit-like opening and a depth greater than half the diameter of its opening. In two species discussed herein the posthumeral pocket is partially concealed by a scaly flap of skin, which I term a posthumeral or ax- illary flap. This structure is described more fully later. Angulate temporal scales are distinctly enlarged, keeled scales posterior to, and in line with, the superciliary scales. When present, they form a distinct border be- tween the posterior head scales and the lateral temporal scales, and they are mor- phologically distinguishable from these se- ries (Cadle, 1991: 6-7; see Fig. 4). Angu- late temporal scales are equivalent to su- 187 pesto as used in some literature (e.g., Avila-Pires, 1995). In several species of eae the angulate temporals are not only keeled but they bear a projecting bladelike vane from the keels; in such cas- es I refer to the scales as “projecting.” Coordinates for localities were obtained from the ornithological gazetteers of the Neotropics (Stephens and Traylor, 1983; Paynter LIS loo 2s M9931 997,), and from Lamas (1976), Morales and Mc- Diarmid (1996), Schulenberg and Awbrey (1997b), and Peruvian department maps produced by the Instituto Geografico Na- cional, Lima. I also consulted fhe on-line versions of the Peru and Bolivia gazetteers of the U.S. Board on Geographic Names at the GEOnet® Names Server: http:// 164.214.2.59/ens/html/index.html. Brack- eted data in localities are inferences from these or other cited sources. Distributions of the new species and others emphasized in this paper are given in Figures 1 and 2. Institutional abbreviations are given at the beginning of the Appendix. DESCRIPTION OF A NEW SPECIES OF STENOCERCUS Stenocercus prionotus? new species Figures 3-7, Figure 12; Table 1 Liocephalus caducus (Cope, 1862): Boulenger (1898), specimen from “Barraca, Rio Madidi” [Bolivia] (BMNH 98.6.9.4) and probably two other northern Bolivian localities discussed in the text. Ophryoessoides caducus (Cope, 1862): Fugler (1989: 63), specimens from San Marcos Ranch, El Beni Department, Bolivia, including ROM 12815. Ophryoessoides aculeatus (O'Shaughnessy, 1879): Fugler (1983, 1986, 1989), specimens from Tumi Chucua, El] Beni Department, Bolivia (USNM par- atypes). Ophryoessoides sp.: Rodriguez and Cadle (1990), specimen from Cocha Cashu, Manu National Park, Madre de Dios Department, Peru (MCZ 150243). * Stenocercus prionotus was recognized as new by R. Etheridge, P. E. Vanzolini, and E. E. Williams many years ago. Vanzolini and Williams applied the unpublished name Stenocercus dorsatus to labels of many specimens in various collections. 188 Bulletin Museum of Comparative Zoology, Woll, ISS INO. 3 . prionotus . aculeatus . fimbriatus . scapularis . huancabambae . limitaris . Iridescens me SN Zul UN kilometers \ al NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle 189 Figure 3. Dorsal and ventral views of the holotype of Stenocercus prionotus (USNM 193683). Approximately <0.87. — Figure 2. Northern Peru and Ecuador (see Fig. 1) showing distributions of species in the “Ophryoessoides group.” Numbered localities are documented or suspected cases of sympatry referred to in the text and the Appendix (see Fig. 1 for names). Open circle at 06°S, 77°W is the type locality for Stenocercus aculeatus (Moyabamba, San Martin Department). All other localities are based on specimens examined; see Cadle (1991, 1998) for S. iridescens, S. limitaris, and S. huancabambae. The known distributions of all species are indicated by the localities plotted; however, the distribution of S. iridescens continues farther north in western Ecuador than the area covered by the map. 190 Holotype (Figs. 3-5). United States National Museum of Natural History (USNM) 193683 (field number WCS 2421). PERU: Drepro. HuANUCo: Jardin Botdnico de la Universidad Agraria de la Selva, Tingo Maria, vicinity of Rio Hual- laga, 670 m elevation [09°18'S, 75°59'W]. Adult male collected 29 June 1966 by Wade C. Sherbrooke. Paratypes from the Vicinity of the Type Locality. PERU: DEpTO. HUANUCO: Ca. 2 mi. by trail W. Tingo Maria, west bank of Rio Huallaga in vicinity of confluence with Rio Monz6n, 670 m elevation (9 October 1966, W. C. Sherbrooke), USNM 193685. Universidad Agraria de la Selva, Tingo Maria, Rio Huallaga, 670 m elevation (18 August 1967, unknown collector for W. C. Sherbrooke), USNM 193686. Vicinity of Cueva de las Lechuzas, ca. 3 mi. SW Tingo Maria, Rio Monz6n, ca. 700 m elevation (17 April 1968, Vito Yaringano for W. C. Sherbrooke), USNM_ 193687. Picuriacu, ca. 2 mi. NW Tingo Maria, Rio Huallaga (20 April 1968, W. C. Sherbrooke), USNM 193688. Other Paratypes. PERU: DEpPpTo. HuANuco: Buena Vista, Valley of the Chimchao [= Rio Chinchao] [approximate- ly (9°35, 7a 52 W)| (l=15> September 1923, E[dmund] Heller), FMNH 5582- 83. Hacienda Pampayacu [09°33’S, 75°54'W] (17 July-16 August 1936, Dr. Snonge), MCZ 43758-59, 43761-62. Rio Llullapichis, 4-5 km upstream from Rio Pachitea, 200 m elevation [09°37’S, 74°55'W] (January 1969, Hans W. Koep- cke), KU 179058. [DEPToO. LORETO]: Elastern| Peru, Pampa Hermosa, near mouth of Rio Cushabatay, Rio Ucayali Val- ley, 500 ft. [152 m] [07°12'S, 75°17'W] (date unknown, H|arvey] Bassler), AMNH 56760-64. DEPTO. MADRE DE DIos: Co- cha Cashu Biological Station, Manu Na- tional Park [11°51’S, 71°19’W] (July 1975, John W. Fitzpatrick), MCZ 150243. Ex- plorer’s Inn, Tambopata Reserve, ca. 30 km (straight line) SSW Puerto Maldonado, 280 m [12°50'S, 69°17’W] (2 September— 7 October 1983, native collectors), USNM Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 247468-69, 247680; (23 May 1986, Vic- tor R. Morales), USNM 269022. DEPpTo. Puno: Prov. Sandia: Tambopata, San Juan (dell Oro], 1520-m or S000: fe 4co7s: 69°08'W |? (22 November—20 December 1950, Hilda H. Heller), FMNH 64788-— 92, 64794-811. [DEPTO. SAN MARTIN]: Juanjui [07°11'S, 76°45’W] (collector and date unknown), MCZ 121233. Tarapoto, B.Orm: lO6280%Ss (6125.1 (25 July 1984, Rainer Schulte), KU 212629. BOLIVIA: DEpTo. BENI: Provincia Va- cadiez, Tumi Chucua [176 m; 11°08’S, 66°10'’W] (23 October—18 November 1981, Charles M. Fugler), USNM 280246-51. Puerto Cruzeiro, San Marcos Ranch at confluence of Rios Isiboro and Ichoa [15°17'S, 65°45'W] (10 February 1977, is Lovisek), ROM 12815. [DEPTo. La Paz]: Barraca, Rio Madidi [12°35’S, 67°02'W] (1893, Luigi Balzan), BMNH 98.6.9.4. Distribution (Fig. 1). Stenocercus prion- otus is known from the lowlands and An- dean foothills of eastern Peru and adjacent Bolivia (San Martin department, Peru, south to the Rio Beni in northern Bolivia). The known elevational range is 176—1,520 m, but the highest recorded elevation is more than twice the elevation of the next lower one. Most localities in Peru are ad- jacent to or near the Andean foothills, and several localities in La Paz and El Beni departments, Bolivia, are unconfirmed. No specimens are known from a broad geo- graphic hiatus between central and south- em Peru (Fig. 1). See Distribution Pat- terns in Stenocercus prionotus for further discussion. Etymology. The epithet prionotus is de- 3 Hilda Heller’s notes on this collection in the Karl P. Schmidt archives of the FMNH describe San Juan as the site of an agricultural station on “the left side of the Rio Tambopata at 5000 feet.” With some hes- itancy I identify Heller’s locality as the town known as San Juan del Oro, which is on the left bank of the Rio Tambopata at approximately the elevation given by Heller. I have located only two other places named San Juan in Puno Department, but neither is on the Rio Tambopata. NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ¢ Cadle rived from the Greek adjective prionotos meaning jagged or serrate. The name re- fers to the strongly serrate vertebral crest of Stenocercus prionotus, which is the most obvious character distinguishing this species from its apparent closest relative, S. caducus. Data on the Holotype. Adult male, hem- ipenes partially everted. SVL, 83 mm. Tail length, 201 mm. Total length, 284 mm. Tail/total length, 0.71. Vertebral scales be- tween the occipital and the posterior mar- gin of the hind limb, 31. Midbody scales, 42. Gular scales between the ears, 16. In- ternasals, 6. Subdigital scales on fourth fin- gers and toes, respectively, 18-18, 25-25. Color pattern well preserved: top of head brown with narrow dark brown interorbital bar extending laterally onto supraoculars; dorsum brown with narrow blackish chev- rons middorsally (1 on neck, 1 above fore- limbs, 2 others anterior to midbody; pos- terior chevrons poorly defined); dark brown scapular blotch bordered anteriorly by white antehumeral stripe; ill-defined grayish dorsolateral streaks between ear and anterior body; throat grayish with poorly defined oblique light grayish stripes; venter brown without distinct pat- tern. Definition. A species of Stenocercus characterized by the following features: (1) Dorsal head scales subimbricate and strongly keeled to multicarinate; temporals keeled, imbricate or subimbricate. (2) Pos- terior head scales larger than anterior ones, with distinct interparietal, a pair of parietals, a pair of postparietals, and a large median occipital (often surrounded by several small irregular scales). (3) In- ternasals usually 7, but pattern irregular and may be 5 or 6. (4) One row of supra- oculars distinctly enlarged. (5) One canthal on each side between the superciliaries and the lateralmost internasal. (6) A pair of strongly keeled angulate temporals in line on each side (rarely, 3 angulate tem- porals are present), each with a low pro- jecting blade; partially or completely sep- arated from enlarged posterior head scales OM by a single row of small scales. (7) Anterior and posterior gular scales strongly keeled. (8) Parietal eye distinct. (9) Neck folds ab- sent; a vertical, strongly keeled row of scales in the antehumeral region and oc- casionally a much less distinct raised series in the supra-auricular region. (10) Dorsal and ventral body scales imbricate, mucro- nate, strongly keeled; dorsal scales at mid- body 36-48. (11) Vertebral row continu- ous, bearing a strongly projecting serrate crest in adults; a dorsolateral crest present on posterior body and the base of the tail. (12) Deep posthumeral pocket (Type 4) partially concealed by a scaly posthumeral flap originating on its anteroventral bor- der; postfemoral pocket absent (Type 1). (13) Scales of posterior thigh imbricate, keeled. (14) Tail strongly compressed in adults, anteriorly with low vertebral and dorsolateral crests continuous with those of the body. (15) Dorsal coloration of males in preservative (Figs. 3, 6) brown with or without distinct chevrons; a dis- tinct white vertical antehumeral stripe ex- tending ventrally to the proximal ventral surface of forelimb; a large dark scapular blotch; in well-preserved specimens the throat bears oblique alternating dark and light stripes (see Description); females similar but pattern elements often more subdued. DIAGNOSIS AND COMPARISONS In having enlarged posterior head scales, an enlarged row of supraoculars, and strongly keeled ventral scales, Steno- cercus prionotus is like other species in the “Ophryoessoides group” of Stenocercus. These are the species most likely to be confused with S. prionotus. Five other de- scribed species of the “Ophryoessoides group occur in eastern Peru or Bolivia: S. aculeatus (O’Shaughnessey, 1879); S. cad- ucus (Cope, 1862); S. fumbriatus Avila-Pi- res, 1995: S. huwancabambae Cadle, 1991: and S. scapularis (Boulenger, 1901). An undescribed species occurs in the Rio Maranon valley of eastern Peru (see Key to Species of the “Ophryoessoides Group” 192 of Stenocercus). Stenocercus prionotus and S. caducus are unique among known species of Stenocercus (perhaps unique within iguanids) in having deep posthu- meral mite pockets (Type 4) that are par- tially concealed anteroventrally by a scaly flap, which may be termed a posthumeral or axillary flap (Fig. 5). Stenocercus prion- otus and S. caducus are compared in greater detail below, but S. prionotus is distinguished from S$. caducus (character- istics in parentheses) by: (1) a strongly pro- jecting, serrate vertebral crest (low and scarcely projecting); (2) 2 (usually) or 3 en- larged, strongly keeled, projecting angu- late temporal scales on each side (scales not greatly enlarged, less projecting); and (3) a gular pattern consisting, when evi- dent, of oblique alternating dark and light lines or bars, or oblique light lines on a dark ground color (usually light spots on a darker ground color, unicolor, or [rarely] a pattern similar to that of S. prionotus). Readily determined characters distin- guishing Stenocercus prionotus from the other four species of the “Ophryoessoides group” known from eastern Peru and Bo- livia include the extent of keeling on dorsal head and body scales, relative develop- ment of the postfemoral pockets, and the number of midbody scale rows (Table 1 and key presented later herein). Stenocer- cus fimbriatus and S. aculeatus are known to be sympatric with S. prionotus at several localities in eastern Peru. In addition to having a posthumeral flap (absent in S. fimbriatus and S. aculeatus), S. prionotus is distinguished from S. fumbriatus (char- acteristics in parentheses; see Avila-Pires, 1995) in having strongly keeled dorsal scales in adults (smooth or weakly keeled), a dorsolateral crest prominent only on the posterior body (prominent anteriorly and continuous with antehumeral and supra- auricular folds or crests), and in lacking “fimbriate” scales on the posterior distal portion of the thigh (present). Stenocercus prionotus is distinguished from S. aculea- tus (characteristics in parentheses) in lack- ing a postfemoral pocket (moderate to Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 deep); in having strongly keeled, often multicarinate, head scales (smooth or weakly striated in adults, wrinkled in ju- veniles); 5-7 internasals (4-5); and only moderately enlarged supraoculars, usually 5—6 supraoculars across the widest part of the orbit (greatly enlarged, usually 4 across the orbit). Stenocercus prionotus differs from S. scapularis (characteristics in parentheses) in lacking squarish or rectangular project- ing superciliary scales (present) and in having fewer than 50 midbody dorsal scale rows (59-70 rows). Stenocercus prionotus differs from S. huancabambae (characters in parentheses) in lacking a postfemoral pocket (deep, Type 5) and in having prom- inent dorsolateral crests on the posterior body (weak, restricted to anterior body when present). ‘Two species of Stenocercus from west- ern Ecuador and Peru, S. iridescens and S. limitaris, have enlarged posterior head scales and supraoculars. In contrast to Stenocercus prionotus, S. iridescens has smooth head plates, 2 canthals, a poorly developed posthumeral pocket (Type 1 or 2), and lacks keeled angulate temporals and dorsolateral crests on the body (see Cadle, 1991, fig. 10). Stenocercus limitaris has a deep postfemoral pocket (Type 5), 2 canthals, a single strongly keeled (but non- projecting) angulate temporal, and lacks dorsolateral crests. Other non-Peruvian species of the “Ophryoessoides group” can be distin- guished from Stenocercus prionotus by features in the key presented later and other superficial characters, such as the presence of 2 canthals and 4 internasals (usually) in S. erythrogaster (1 and 5—7 in S. prionotus), and enlarged, projecting py- ramidal or conical postsuperciliary scales in S. dumerilii and S. tricristatus (Avila- Pires, 1995). Other species of Stenocercus are distin- guished from S. prionotus by a combina- tion of features such as smaller head plates, smooth ventrals, and absence of dorsolateral crests. Most species of Steno- NEW SPECIES OF STENOCERCUS cercus except the “Ophryoessoides group” have smooth or weakly keeled head plates. Other characters, such as the number of dorsal scale rows, morphology of the pos- thumeral and postfemoral mite pockets, extent of sexual dimorphism, and degree of differentiation of the vertebral scale row and crest also aid in distinguishing the spe- cies (see descriptions and discussion in Fritts, 1974; Cadle, 1991, 1998). Apart from Stenocercus fimbriatus, S. scapularis, and S. aculeatus, two other spe- cies of Stenocercus are known from local- ities close to or sympatric with known pop- ulations of S. prionotus: S. crassicaudatus and S. roseiventris. These species are dis- tinguished from S. prionotus by lacking prominent serrate vertebral crests (low crests may be present), having smooth ventral scales, and having pr ominently spi- nose tails with the spines arranged in dis- tinct whorls. DESCRIPTION Head (Fig. 4). Dorsal head scales sub- imbricate (a tendency to be more juxta- posed posteriorly); str ‘ongly keeled to mul- ticarinate or wrinkled. Rostral in contact with first supralabial, first lorilabials, and a series of postrostrals. Usually 7 elongate, strongly keeled internasals between the nasals dorsally; however, the anterior dor- sal head scales are very irregular and oc- casionally only 5 or 6 internasals are pre- sent. One canthal scale between the an- terior superciliary and the lateralmost in- ternasal, separated from the nasals by tiny postnasals. Canthus very strongly angled. Nostril in posterior portion of an elongate nasal scale, which may contact the rostral scale anteriorly or be separated from it by small postrostrals. Four or 5 strongly over- lapping, elongate anterior superciliaries followed by 2 or 3 shorter posterior su- perciliaries slightly overlapping in the re- verse direction (but more or less in a straight line). One supraocular row mod- erately enlarged, 2 mediocentral scales much larger than the others. Five or 6 scales across the supraocular area at its S FROM PERU AND BOLIVIA ¢ Cadle 193 widest part. Interparietal distinct and elon- gate, diamond-shaped or pentagonal (apex posteriorly). P Parietal eye Tobie. A pair of parietals in contact behind the interpari- etal, flanked posterolaterally by a postpar- ietal on each side. Postparietals separated medially by a single median transversely elongate occipital: ‘occasionally 1 or 2 small coales are intercalated at the juncture of the parietal, postparietal, and/or occipital (ee) buen 4) Lateral temporal scales strongly keeled, imbricate to subimbricate; separated from posterior dorsal head scales on each side by 2 (occasionally 3) elongate, strongly keeled angulate temporal scales bearing a low pr ojecting vane. Keels of adjacent an- gulate temporals aligned. Posterior angu- late temporals separated from postparie- tals by 2 or 3 small scales in a longitudinal row. Anterior angulate temporals may con- tact postparietal and one other larger pos- terior head scale or be separated from them by small scales. Anterior border of ear weakly denticulated; posterior border rounded, bordered with keeled imbricate scales. Anterior and posterior gulars strongly keeled. Mental smooth, in contact with first pair of postmentals and first pair of infralabials. Enlarged postmentals 3 or 4 on each side, only the first pair in contact medially. Neck and Body. Dorsal and lateral scales of neck and body imbricate, mucro- nate, strongly keeled. Vertebral row pro- duced into a prominent projecting serrate crest in adults of both sexes that is contin- uous from the nuchal region to the base of the tail, gradually disappearing on the anterior “4 to % of tail. Dorsolateral crest (a raised, strongly keeled row of scales) on posterior % of body, continuing onto base of tail. The dorsolateral crest occasionally appears very indistinctly farther anteriorly on the body, but only on the posterior body does it sharply delimit the dorsolat- eral (paradorsal) scales from the flank scales. Three rows of scales between dor- solateral and vertebral crests at anterior 194 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 | GBB ERS a E( =v Ne ce ER CS SEES hen ES Las es Figure 4. Head scales of the holotype of Stenocercus prionotus (USNM 193683) in dorsal and lateral views. Scale bars = 3 mm. To facilitate coordination with the text, the following scales are indicated on one side (interparietal and occipital are median scales): A, angulate temporal; |, interparietal; O, occipital; P, parietal; PP, postparietal. NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle edge of dorsolateral crest (i.e., just anterior to the pelvic region), 2 rows posteriorly (dorsal to hindlimbs); scales between the crests strongly imbricate and keeled only on posterior part of scale. Flank scales mostly fully keeled (sometimes only the posterior part of each scale), imbricate, mucronate, slightly smaller than dorsolat- eral scales. Ventral body scales mucronate, strongly keeled (keels running the length of the scales). Ventrals approximately the same size as the dorsolateral scales, larger than flank scales. Neck Folds. Distinct neck folds absent. Poorly developed antehumeral crest pre- sent. Tail. Tail strongly compressed, anteriorly bearing low projecting vertebral and dor- solateral crests continuous with those of the body. Dorsal scales moderately keeled, ventral scales strongly keeled. Limbs. Dorsal and ventral scales of fore- limbs, hindlimbs, and posterior thigh strongly keeled, unicarinate, mucronate; some scales of the shank larger than any thigh scales. Supradigitals and subdigitals unicarinate. Palmar scales strongly multi- carinate. Plantar scales strongly unicari- nate. Posthumeral and Postfemoral Mite Pockets. Posthumeral mite pocket a deep cavity (Type 4) with a prominent axillary flap concealing the anteroventral aspect (Fig. 5). Postfemoral pocket absent (Type ib) The flap associated with the posthumer- al pocket projects from the anteroventral and ventral edges of the pocket. Antero- dorsally, a similar but much smaller flap is present in some specimens (e.g., Fig. 5). The posthumeral flap consists of a fleshy ridge covered anteriorly and _ posteriorly (or externally and internally when the flap is lying flat against the body) by keeled im- bricate scales. Externally, usually 3 or 4 larger scales cover the flap ventrally and a series of much smaller scales is present dorsally. One or 2 of the larger scales are sometimes highlighted with white. When appressed against the body (i.e., against 195 Figure 5. Posthumeral (axillary) flap of Stenocercus priono- tus (USNM 193683, holotype). Anterior to the right. The broad oval on the right is the deflected forelimb and the posthumeral mite pocket is the heavily stippled cavity deep to the flap. The posterior border of the axillary flap is marked by the scales with heavily outlined posterior borders and it extends anteriorly to the ventral part of the forelimb. A smaller dorsal flap is also present in this specimen (small patch of heavily outlined scales on anterodorsal edge of the pocket; see text). Scale bar = 1 mm. the opening of the posthumeral pocket) the flap conceals approximately the ventral % to % of the vertical dimension of the pocket. The flap is equally prominent in adults of both sexes and is proportionally as well developed in subadults (including hatchlings) as in adults. Size and Proportions. Largest male (USNM 193683) 89 mm SVL, 323 mm to- tal length (sample size of males with SVL =70 mm = 13). Largest female (KU 212629) 93 mm SVL, 329 mm total length (sample size of females with SVL =70 mm = 16). Tail relatively long, 69-74% of total length in adults (67-71% in juveniles). Coloration and Pattern of Adult Males in Life. The following color descriptions are paraphrased from the field notes of Wade C. Sherbrooke. USNM 193683 (ho- lotype): Lower half of side between limbs is lavender- brown. This color extends from both sides across the belly approximately % of the way on each side, leaving a tan-brown central strip down the belly. General base color of the body is brown, darkest 196 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 Figure 6. Lateral view of Stenocercus prionotus from northern Peru (USNM 193685; male, snout-vent length 76 mm). Note the high vertebral crest and the following pattern elements: dark subocular bar, dark blotches on the dorsal and lateral surfaces of the neck, pale antehumeral bar, indistinct pale dorsolateral stripe, and indistinct oblique bars on the trunk. near the hind legs and tan just in from of the fore- legs. A distinct white line runs dorsally from the top of each foreleg to three quarters of the way to the dorsal crest; it runs through a large black patch just above the forelimb. Head markings consist of a dark brown line running between the eyes on the top of the head; this continues through the eye to broaden slightly at the rear portion of the jaw. The gular area is streaked by several light cream lines. Very slight lavender tinge to all a body behind forelimbs. [Sketch in notes shows black middorsal patches that don’t extend to flanks]. USNM 193685 (WCS 2543): This specimen closely resembles [USNM 193683] in color, with one exception. There are several green spots on... the right dorsal surface between the limbs and on the dorsal portions of the tail behind the hind limbs and the dorsal tibio-fibula portion of the right hind leg. Coloration of Adult Females in Life. Un- known. Coloration in Preservative (Figs. 3, 6, 7). Specimens of Stenocercus prionotus vary greatly in coloration due mainly to varia- tion in initial preservation and the length of time in preservative. Poorly preserved specimens may be more or less uniform brown all over, although obscure pattern elements are usually present. Well pre- served specimens are brown dorsally with darker brown to blackish chevrons mid- dorsally. One chevron dorsal to each pair of limbs and one on the neck are usually evident, and these are darker than others that may be present. Three to five mid- dorsal chevrons are between the limbs. Dark spots or an additional chevron are often present on the dorsal neck and usu- ally on the base of the tail. The light an- tehumeral/humeral line is universally pre- sent and evidence of the dark shoulder patch is usually present (often very prom- inent). Flanks usually unicolor and some- what darker than the dorsum between the dorsolateral crests; however, some speci- mens (e.g., KU 179058, USNM 280246) have distinct dirty white vertical bars or chevrons on the flanks (five between the limbs), and such bars appear occasionally, but more obscurely in other specimens (see Fig. 6). The dorsolateral crest is often highlighted for a variable length with a dis- tinct or indistinct light line, giving the im- pression of a light dorsolateral stripe. Forelimbs more or less unicolor brown or with obscure pattern; hindlimbs brown with darker brown bands. Dark subocular bar distinct. Top of head often with an ob- scure or distinct dark brown interocular bar. Oblique bars on throat (Fig. 7) often visible but throat may be unicolor or have an obscure pattern. Venter of most speci- mens unicolor, dirty white, gray, or beige; however, some specimens (e.g., USNM 280246) have a series of irregular longi- tudinal dark brown streaks. Scale Counts and Qualitative Features (Table 1). Stenocercus prionotus has rela- tively low midbody, vertebral, and gular scale counts. The scales are relatively large and strongly keeled over most of the body. NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle Figure 7. Gular patterns in Stenocercus prionotus. Top, typ- ical throat pattern (oblique view) consisting of light and dark stripes that extend medially to the midline (USNM 193687). Bottom, specimen in which the ground color is lighter and therefore the contrasting pale stripes are less distinct (USNM 193685). Sexual Dimorphism. Stenocercus prion- otus does not exhibit strong sexual dimor- phism. Males and females attain approxi- mately the same size and have the same general pattern, but whether the colora- tion in life reported above for adult males pertains to females as well is unknown. The vertebral crest is only slightly more developed in males than in females of the same population but this character shows strong clinal variation (northern popula- tions with higher crests; further discussed below). Other characters that sometimes vary between the sexes in Stenocercus show little variation in S. priontotus. Nei- U7 ther standard meristic counts (Table 1) nor the relative development of the posthu- meral and postfemoral pockets (Types 4 and 1, respectively, in adults of both sexes and in subadults) show obvious sexual di- morphism. DISTRIBUTION PATTERNS IN STENOCERCUS PRIONOTUS Absence of Stenocercus prionotus from Lowland Localities in Eastern Peru. Sten- ocercus prionotus is widespread in the lowlands along the Andean front from northern Peru to northern Bolivia (Fig. 1). However, all Peruvian localities are close to the Andean foothills and south of the broad extension of the Cordillera Oriental separating the great bend of the Rio Mar- afion from upper reaches of the Rio Hual- laga (Fig. 2). The absence of specimens in comprehensive collections from the Iqui- tos region (Dixon and Soini, 1986), Balta (Ucayali Department; specimens at LSUMNS and University of Arizona), and Cuzco Amazonico (Madre de Dios De- partment; Duellman and Salas, 1991) sug- gest that S. prionotus may be absent from the lowlands distant from the Andean foot- hills, at least in Peru. Similarly, collections from northern Loreto Department (Duell- man and Mendelson, 1995) and northern Amazonas Department (J. E. Cadle and R. W. McDiarmid, unpublished data from the Rio Cenepa and Rio Santiago) suggest that S. prionotus does not occur north of the Rio Maranon. However, these sites have been sampled unevenly. For example, species accumula- tion curves for lizards at Cuzco Amazonico (Duellman and Koechlin, 1991) reached an asymptote after about 15 person-weeks of effort, whereas only 5 person-weeks were expended in northern Loreto and the species accumulation curve for the total herpetofauna showed no asymptote (Duellman and Mendelson, 1995; data not presented for lizards only). Quantitative data are not available for the other sites, but large collections are available for the Iquitos region, including more than 1,000 198 TABLE 2 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 PATTERNS OF PRESUMED SYMPATRY OF STENOCERCUS IN EASTERN PERU. ONLY SPECIES OF THE “OPHRYOESSOIDES GROUP” AND THE SUPERFICIALLY SIMILAR SPECIES, S. ROSEIVENTRIS, ARE LISTED.! LOCALI- TIES ARE LISTED ROUGHLY NORTH TO SOUTH; FOR PRECISE LOCALITIES SEE THE APPENDIX AND LOCALITIES FOR THE TYPE SERIES OF S. PRIONOTUS.. Iquitos San Martin/ Pampa i Tingo Cuzco Explorer's Region W. Loreto? Hermosa Mishollo Maria Manu? Balta Amazonico Inn S. prionotus x x x x x S. aculeatus x S. fimbriatus x x x x x S. roseiventris x x x x '' The only other species of Stenocercus known from primarily the lowlands and lower Andean foothills in this region is S. crassicaudatus, which lacks a projecting vertebral crest and has a very spiny tail. > The distributions of S. prionotus and S. aculeatus overlap both altitudinally and latitudinally in northern Peru but they have not been taken together at the same locality. See Figure 2 for localities. >In Manu National Park, S. fimbriatus and S. roseiventris are sympatric at Pakitza, whereas S. prionotus is known only from Cocha Cashu. lizards obtained by Dixon and Soini (1986) and additional collections from the region made by Harvey Bassler and deposited in the AMNH. Similarly large collections re- sulted from the efforts of Cadle and McDiarmid in Amazonas (specimens in the MVZ and USNM). Thus, barring ar- tifacts introduced by the difficulties of col- lecting cryptic rainforest lizards, S. prion- otus seems to be absent from these sites. Documenting and explaining patterns of absence is always difficult, but the appar- ent absence of Stenocercus prionotus from the lowlands distant from the Andes in Peru is not due to failure to collect Sten- ocercus at these localities because at least one other species of Stenocercus is known from each (Table 2). However, no lowland locality is known in which more than two species of the “Ophryoessoides group” of Stenocercus are sympatric. Of interest in this connection is that S. prionotus has not been taken in the upper reaches of the Rio Perene and its tributaries (Junin Depart- ment), although it is known from north and south of tat region. Numerous spec- imens of Stonocerels of at least five spe- cies (S. boettgeri, S. crassicaudatus, S. for- mosus, S. scapularis, and S. variabilis*) * Based on examination of two of the three synty- pes of Stenocercus variabilis Boulenger (BMNH 1946.8.11.89, 1946.8.11.91) I concur with Fritts (1974: 66) that Boulenger (1901: 553) erred in as- have been collected along an elevational cribing these specimens to “Palca, Bolivia” rather than Palca (Junin Department), Peru. One specimen I collected near Palca, Peru (MCZ 178166) is nearly identical with BMNH 1946.8.11.91 in scale counts, pattern, and qualitative characters (both of these specimens differ considerably from BMNH 1946.8.11.89 in color pattern). However, the attri- bution of the specimens to Peru is not without some equivocation. The types were collected by P. O. Si- mons (see Cadle, 1998: footnote 6), who collected in the vicinity of Palca, Peru, in March and April, 1900, although that specific locality is not listed in his pub- lished itinerary (Chubb, 1919). However, Simons’s itinerary places him at “Palca, 18 miles E of La Paz” [Bolivia] on 9 November 1900 (Chubb, 1919: 5). Si- mons'’s field tags attached to the syntypes record sim- ply “Palca 3000 m,” which is close to the elevation of Palea, Peru (2,740 m), but not that in Bolivia (4,600 m). Adding to the confusion are entries in the BMNH registries for the syntypes, concerning which Colin J. McCarthy provided the following comments via e-mail: Boulenger originally wrote “Palca Peru 3000m” but later struck through Peru and wrote “Bolivia.” There are two sheets of notes (presumably from Simons) stuck in at this page of the register about the localities in the batch. With regard to Palca he has written “Palca, just S. of La Paz, Bolivia.” I assume it was that information that caused Boulen- ger to alter his original entry! Thus, the confusion may have originated with the note that Simons provided subsequent to cataloguing of the collection at the BMNH. In any case, no spec- imens resembling S. variabilis are definitely known from Bolivia and the only species of the genus defi- nitely known to occur above 4,000 m is Stenocercus chrysopygus from northern Peru. NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA * Cadle transect along this well-traveled route (Fritts, 1974; Cadle, 1991 and unpublished data). Thus, analysis of circumstantial dis- tributional data suggests that the distribu- tion of S. prionotus in the lowlands of east- ern Peru may be influenced by the num- ber of sympatric species of Stenocercus. At all localities from which S. prionotus has been taken, only one other species of Sten- ocercus is known (Table 2). These patterns of sympatry are discussed in the next sec- tion with reference to patterns of geo- graphic variation in S. prionotus. In contrast, the apparent restriction of Stenocercus prionotus to the Andean foot- hills and immediately adjacent lowlands in Peru does not seem related to the distri- bution of any major habitat type or phys- iographic region. Most of the extensively sampled localities (e.g., Iquitos and Cuzco Amazonico) include a variety of lowland habitats characteristic of western Amazon- ia. Stenocercus prionotus is known from both floodplain forests (Cocha Cashu) and more upland forests on river terraces in southeastern Peru (Explorer's Inn); Foster (1990) and Dallmeier et al. (1996) de- scribed these floristic communities in southeastern Peru. Thus, the apparent ab- sence of S. prionotus at the localities dis- cussed above is not due to some simple relation to local habitat availability. For ex- ample, it is unclear why S. prionotus was not obtained at Pakitza (Morales and McDiarmid, 1996), even though it occurs at nearby Cocha Cashu. On a broader geo- graphic scale, the restriction of S. priono- tus to lowlands and foothills adjacent to the Andes may be related to the present or historical influence of the Andes on the climate and vegetation (rainfall, tempera- ture, and major soil types) of neighboring regions. A curious and unexplained hiatus in the distribution of Stenocercus prionotus oc- curs between the vicinity of Tingo Maria— Rio Llullapichis (Hudénuco Department) and Cocha Cashu in Manu National Park (Madre de Dios Department), a gap of some 600 km that includes the entire up- gg per reaches of the Rio Ucayali-Urubam- ba—Ene system. Additionally, populations north and south of this gap differ in some qualitative and quantitative characters (see Patterns of Sympatry and Geographic Var- iation in Stenocercus prionotus, and the Need for Additional Fieldwork). Scattered collections (e.g., maps in Fritts, 1974; Avi- la-Pires, 1995), but no comprehensive her- petofaunal surveys or collections, are avail- able from this vast region. Thus, whether the geographic hiatus is real or a sampling artifact cannot be discerned. Efforts to re- solve this issue need to be made. Unconfirmed Bolivian Localities for Stenocercus prionotus. Because of the previous confusion of Stenocercus priono- tus with S. caducus, specimens perhaps re- ferable to S. prionotus from several local- ities in Bolivia are unconfirmed. Boulen- ger (1898) reported specimens of “Lioce- phalus caducus, Cope” in the Museo Civico Storia Naturale Giacomo Doria in Genoa collected by Luigi Balzan from four localities (Balzan, 1931). I have not at- tempted to verify the existence of these specimens. However, one of Balzan’s spec- imens from “Barraca, Rio Madidi” was ex- changed to the BMNH (now BMNH 98.6.94), and is confirmed as S. prionotus. The “Leiocephalus caducus” specimens from the other three Balzan localities are outside the known distribution of S. cad- ucus but are close to other known localities for S. prionotus in northern Bolivia (Fig. 1). I suspect these are S. prionotus based on geographic location. The localities are, as listed by Boulenger (1898) (see Fig. 1), (1) “Coroico and Ghuilsmnann Prov. Yun- gas, 1,600 metres alt.” [La Paz Depart- ment: Coroico, 1,725 m, 16°10’S, 67°44'W: Chulumani, 1,905 m, 16°24'S, 67°31'W]. (2) “Reyes, right bank of Rio Beni” [El Beni Department, 232 m; 14°19'S, 67°23'W]. (3) “Missiones [sic] Mosetenes” [approximately 15°31’'S, 67°25’W].° These ° The Moseten Indians inhabited upper reaches of the Rio Beni and its tributaries in the Andean foot- hills of the present department of La Paz (Métraux, 200 localities would not be unusual for S. prionotus, although the first two localities are the highest elevations recorded for the species (the species occurs at 1500 m in nearby Puno Department, Peru). All are in the upper reaches of the Rio Beni, whereas the two confirmed Bolivian local- ities for S. prionotus are farther north in the same drainage. Burt and Burt (1931: 273) listed two specimens of “Leiocephalus |=Stenocer- cus| scapularis” (AMNH 22450, 22532) from Rurrenabaque, Bolivia [E] Beni De- partment; 14°28’S, 67°34'W]. These were reidentified in 1971 by Thomas H. Fritts as Stenocercus caducus but they are pres- ently missing from the AMNH collection (Linda Ford, in litt., February 1999). Rur- renabaque is very close to Balzan’s locality (2) above. Based solely on presumed hab- itats and presently known distributions of S. prionotus and S. caducus in eastern Bo- livia (see text; Fig. 1), I suspect that AMNH 22450 and 22532 are most likely S. prionotus. They should be reexamined if they are ever located. As an outside pos- sibility, any of the unverified specimens from these Bolivian localities could rep- resent S. fimbriatus, which is now known from southern Peru (Appendix). PATTERNS OF SYMPATRY AND GEOGRAPHIC VARIATION IN STENOCERCUS PRIONOTUS, AND THE NEED FOR ADDITIONAL FIELDWORK Stenocercus prionotus varies geographi- (S OO cally in several characters, most notably in the height of the vertebral crest. Because 1942). According to Métraux (1942: 15), by the end of the 19th century when Balzan collected his spec- imens the decimated population of the Mosetenes was concentrated in the three “Misiones Mosetenes” of San Miguel de Muchanes (15°14’S, 67°39'W), San- ta Ana (15°31'S, 67°30’W), and Covendo (15°49’S, 67°06'W). The approximate coordinates given in the text are the average for these three sites. According to several maps with elevational contours indicated, Muchanes and Covendo are between 500 and 1,000 m and Santa Ana is less than 500 m in elevation. Mathews (1879) also discusses these missions. Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 of the potential role of the crest in either intra- or interspecific communication it seems appropriate to discuss patterns of geographic variation in S. prionotus in the context of the distribution of other species of the “Ophryoessoides group.” In this re- gard, patterns of sympatry among species of this group in Peru and Bolivia are es- pecially relevant. Patterns of Sympatry of Species in the “Ophryoessoides Group.” The most com- plex distributional patterns for the “Ophryoessoides group” are in northern and central Peru, where six species occur (Stenocercus aculeatus, S. fimbriatus, S. prionotus, S. huancabambae, S. scapularis, and an undescribed species; Fig. 2). In contrast to the other species, S. huanca- bambae and the undescribed species occur in comparatively dry deciduous forests west of the known distributions for the other species. Their distributions probably do not overlap the other species and they will not be considered further here. Based on known latitudinal and elevational dis- tributions, three species (S. aculeatus, S. fimbriatus, and S. prionotus) are probably broadly sympatric in northern Peru, al- though sympatry is documented only for pairs of these at three localities (Fig. 2; Table 2). The habitat preferences of the species in the region of sympatry are un- known. Whether Stenocercus scapularis is sym- patric with S. prionotus is less clear. Sten- ocercus scapularis is known from two widely separated areas in Hudnuco and Puno departments (Peru) (Fig. 1; Appen- dix), and it seems to be elevationally par- apatric to S. prionotus in northern Peru (Hudnuco Department). In this area, S. scapularis occurs above 1,000 m and S. prionotus is not known above 700 m. In Puno Department, S. prionotus is known from a series obtained by Hilda Heller at San Juan del Oro (1,520 m elevation), which elevationally overlaps the distribu- tion of S. scapularis in the same region (1,000-1,830 m). However, these two spe- cies have not been taken at the same lo- NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ¢ Cadle cality in southern Peru. Interestingly, al- though the largest series of S. prionotus available from a single locality (San Juan del Oro) is in the elevational range of S. scapularis, no specimens of that species were obtained by Heller during a month- long stay at the site. In southern Peru and Bolivia, Stenocer- cus prionotus is not known to be sympatric with other members of the “Ophryoesso- ides group.” However, based on the oc- currence of S. prionotus and S. fumbriatus at two nearby localities within Manu Na- tional Park (Cocha Cashu and Pakitza,° re- spectively), sympatry for these two species is expected along the Andean front in this region. In southern Peru and Bolivia, S. prionotus is also broadly sympatric with S. roseiventris, a large terrestrial species that is superficially similar to species of the “Ophryoessoides group.” However, these species have been taken together at only one locality, Explorer’s Inn. To summarize these patterns of sympat- ry, the distributions of three species of the “Ophryoessoides group” overlap broadly in northern Peru. Stenocercus prionotus is known to be sympatric with at least one other species of the group in Hudnuco and Loreto departments. No species of the “Ophryoessoides group” are known to be sympatric in southern Peru and Bolivia. Geographic Variation. Geographic vari- ation pertains to both quantitative and qualitative characters in Stenocercus prionotus. Specimens from the northern part of the range have higher average counts of midbody scales than southern specimens, which is shown graphically in Figure 8. However, there is broad overlap in the counts from opposite ends of the range. A reverse trend (lower counts in the north), occurs in the number of vertebral (dorsal crest) scales (Fig. 9); this reflects the reduced prominence, and hence small- ° The record of Stenocercus fimbriatus from Pak- itza (Appendix) extends the known range for this spe- cies south by approximately 200 km ‘from Balta in Ucayali Department, Peru (Avila-Pires, 1995). 201 er scales, of the crest in southern popula- tions. None of these differences in scale counts is statistically significant. The height of the vetebrl crest shows strong Glinal variation in Stenocercus prionotus. Populations from northern and central Peru have higher crests than those from southern Peru Sand Bolivia (Fig. 10). However, the absence of specimens be- tween Hudnuco Department in central Peru and Manu National Park (Madre de Dios Department) in southern Peru (Fig. 1) makes it impossible to analyze this trend in detail. Specimens from the former re- gion have a high crest typical of all speci- mens from that area and farther north, whereas specimens from Madre de Dios Department and farther south have dis- tinctly lower crests. The crest is not strongly sexually dimorphic in either re- gion. Whether the transition in crest height is abrupt or gradual between central and southern Peru is unclear. Indeed, it is un- clear whether intervening populations of Stenocercus prionotus exist. Both patterns of clinal variation have been extensively documented empirically and theoretically for many organisms (Endler, 1977). The pattern of geographic variation in S. prion- otus could conform to any of several in Endlers (1977: 4) classification scheme. However, present knowledge of distribu- tions and character variation is most simi- lar to Endler’s “differentiated disjunction” pattern, wherein disjunct populations of a species diverge in one or more characters. Further speculation is not fruitful in the absence of more extensive collections from the region of disjunction in eastern Peru. A Hypothesis of Causation for Geo- graphic Differentiation in the Vertebral Crest of Stenocercus prionotus. Variation in the height of the vertebral crest is a prominent distinction between northern and southern populations of Stenocercus prionotus. This variation calls for an expla- nation while at the same time recognizing that, in the absence of experimental or field studies, ascribing causation to pat- 48 e@ @ 47 ) 46 @ 45 @ @ 44 ee 43 ee ee 42 eee ee im) 2 gS 41 e 7p) > 3 40 @ @®e © ie} 3 S 39 ) ) @ 38 ee 37 ® @ 36 ee 35 2 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 6.5- 9.3- 7.2 9.6 11.1- 11.8 12.6- 12.9 25.4- 25.6 20.8- 21.3 17.5- 18.3 14.2 Latitude Figure 8. Geographic variation in the number of midbody scales in Stenocercus prionotus (@) and S. caducus (Ml). Each symbol represents one specimen; counts for all specimens examined are plotted. The horizontal axis represents degrees south latitude for each locality converted to a decimal value (e.g., 17°30’ converted to 17.5); geographically contiguous localities were arbitrarily combined to yield the latitudinal ranges given. The S. caducus specimens examined were biased toward Bolivian specimens. terns of geographic variation is difficult. Vertebral crests in iguanids are visual sig- nals. The crests in sexually dimorphic spe- cies are involved in intr aspecific behavioral encounters, especially intrasexual aggres- sion (e.g., Watkins, 1998). Nonetheless, despite strong geographic differentiation, the vertebral crest in Stenocercus priono- tus is not strongly sexually dimorphic in any part of its range. Thus, a role in in- trasexual behavioral signaling seems un- likely to be the primary function of the crest in this species. Intersexual signalling is a possibility, but this would not explain why northern and southern populations differ so strongly in this character. Populations of Stenocercus prionotus with the highest vertebral crests occur in the area of highest species density and documented Supe for the “Ophryoes- soides group” (Table 2 and above discus- sion). Not only do northern populations of S. prionotus have the highest vertebral crests within this species, but in northern and central Peru S. prionotus has a much more prominent vertebral crest than other species of the “Ophryoessoides group” (S. aculeatus approaches it most closely). In NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle 203 43 42 A 40 39 e e 38 e e e 37 ee 36 ee e ee 35 e e 34 e e Vertebral Crest Scales 6.5- 9.3- 11.1- 12.6- 7.2 9.6 11.8 12.9 17.5- 15.3 18.3 21.3 25.6 14.2 14.6- 20.8- 25.4- Latitude Figure 9. Geographic variation in the number of vertebral crest scales in Stenocercus prionotus and S. caducus. Symbols and axes are the same as Figure 8. contrast, in southern Peru and northern Bolivia the distribution of S. prionotus overlaps only S. scapularis and (perhaps) S. fimbriatus of the “Ophryoessoides group, but S. prionotus is not known to be sympatric with either of these species.‘ 7 The only case of sympatry of any of these with other species of Stenocercus of which I am aware involves S. prionotus and S. roseiventris at Explorer's Inn (cf. Table 2), but S. roseiventris is phenotypically dissimilar to species of the “Ophryoessoides group” in having shorter limbs, different color pattern, spiny tail, and different body form. The vertebral crest of S. prionotus in southern Peru and northern Bolivia is sim- ilar in development to that in allopatric populations of S. aculeatus. I conjecture that the strongly developed vertebral crests of Stenocercus prionotus in northern Peru may be related to the presence of multiple sympatric congeners in that part of the range, and that the crest functions as a species recognition charac- ter. If true, this explanation for variation in crest height would have no precedent among lizards. The only analogous situa- 204 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 (a) prionotus AS Huanuco, Peru 67 mm (es) (b) prionotus FE Puno, Peru 73 mm @ (Cc) prionotus Beni, Bolivia 81 mm ©) ae or (d) caducus Sta. Cruz, Bolivia 72 mm NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle tion seems to be dewlaps in Anolis, in which species recognition has been pos- tulated as a biological role for dewlap color and size in complex assemblages of these lizards (Williams and Rand, 1977; Losos and Chu, 1998). The Need for Additional Fieldwork. Only field observations and experimental studies of the function of the vertebral crest in Stenocercus prionotus can deter- mine whether the functional hypothesis advanced here is correct. Ideally, parallel studies should be conducted in the area of sympatry with other species of the “Ophryoessoides group,” as well as in the area with no known sympatric species of that group. That design would permit an evaluation of any context-dependent use of the crest in these populations. Two sym- patric species are known from the vicinity of Tingo Maria (Hudnuco Department; Table 2), a very accessible area for study. Two other reasons for additional com- prehensive fieldwork on these species are indicated. First, we need documentation of the extent to which species of the “Ophryoessoides group” are sympatric or syntopic. Particular targets should be the northern part of the range of Stenocercus prionotus, where S. aculeatus and S. fim- briatus are known to have overlapping ranges; and Manu National Park in south- ern Peru, a reasonably accessible area where S. fumbriatus and S. prionotus are known to be closely allopatric (but not sympatric) on a microgeographic scale (Ta- ble 2). Second, the character differences between the northern populations of S. prionotus and those of southern Peru and Bolivia could indicate that two or more species reside within my concept of this species. Fieldwork concentrated in the e= 205 geographic gap between Tingo Maria and vicinity (Hudnuco Department) and Cocha Cashu (Madre de Dios department), the two most proximate localities of the north- ern and southern population groups, is needed to verify whether intermediate populations occur. The possibility that these might be two allopatric species sep- arated by a broad geographic gap should not be dismissed wathout further evalua- tion. NATURAL HISTORY OF STENOCERCUS PRIONOTUS Few natural history observations exist for Stenocercus prionotus. Wade C. Sher- brooke (field notes) found the holotype running in the relatively open floor of a bamboo garden at about 1200 h. USNM 193685 was taken along a trail within rain- forest. Brief notes ascociated with the se- ries from Explorer’s Inn (Madre de Dios Department, Peru; USNM 247468-69, 247680, 269022) indicate that specimens were obtained during the day on the ground from clearings around the lodge, although USNM 269022, an adult female (91 mm SVL), was on a leaf 40 cm above the ground. MCZ 150243 was retrieved from a mist net near the edge of tall flood- plain forest at Cocha Gaskin: In the dry season (September—December) of a low- land rainforest in the vicinity of Tumi Chu- cua (Beni Department, Bolivia), Fugler (1986: table 5) found Stenocercus priono- tus (reported as Ophryoessoides aculeatus ) in varzea (seasonally inundated rainforest) but not in terra firme rainforests of the area; females with enlarged eggs were found in early November (Fugler, 1986: table 4). Hilda H. Heller provided the following Figure 10. Diagrammatic representation of geographic and size-related variation in the height of the vertebral crest in Steno- cercus prionotus and S. caducus. Drawings were made with a camera lucida to emphasize the form and height of the vertebral crest. Sketches are drawn to an approximately uniform interval between the external ear opening (EE) and the white antehumeral stripe (AS). For each specimen the geographic location and the SVL are given (all specimens are adult males): (a) MCZ 43759, (b) FMNH 64799, (c) USNM 280250, (d) CM 970. Note especially the differences between the northern specimen of S. prionotus (a) compared to southern ones (b and c), especially given the size differences among these; and the differences between size- matched specimens of S. prionotus and S. caducus (b and d). 206 notes on the San Juan [del Oro] locality from which she obtained a series of Sten- ocercus prionotus in the early 1950s (K. P. Schmidt archives, FMNH): Steep forest with deep undergrowth. Steep fields. Rainfall probably somewhat greater than at Pampa Grande,® due to its colder climate and steep ex- posure; I have no figures. Brushy second growth may be burned in fairly wide patches in August, and [the] resort is frequently made to burning, in- dicating a moderately dry winter period. As of the mid-1980s very little undisturbed forest was left in the vicinity of San Juan del Oro (personal observations). Although Heller provided detailed notes on some of the snakes and frogs from her collection, she makes no specific comments about the lizards. Most observations suggest that Steno- cercus prionotus prefers open habitats, such as areas of human disturbance and light gaps within forests (e.g., created by trails), rather than deep rainforests. Alter- natively, the observations may simply in- dicate the ease of observation and capture in more open habitats. A combination of tic coloration and escape behavior (rapid flight followed by immobility) pos- sibly makes S. prionotus very difficult to observe in closed-canopy rainforest, as re- ported for the similar species, S. fumbria- tus (Dixon and Soini, 1986; Avila-Pires, 1995) and S. caducus (Scrocchi et al., 1985).° However, we currently lack obser- vations to support these statements for S. prionotus. COMPARISON OF STENOCERCUS PRIONOTUS WITH S. CADUCUS A scaly flap associated with the posthu- meral pockets is a unique and unquestion- ably derived character shared by Stenocer- cus prionotus and S. caducus, which sug- gests that these are sister species (Figs. 5, *T have been unable to localize Pampa Grande. ° Cei (1993) claimed that Stenocercus caducus was arboreal, but Scrocchi et al. (1985) reported the be- havior of this species in more detail and stated that it was terrestrial. Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 11).° I am unaware of a similar structure in any other lizards. Some individual vari- ation in the precise form and size of the flap occurs in both species but it seems extremely improbable that these structures are not homologous in the two species. Moreoever, Stenocercus prionotus and S. caducus are similar in standard meristic characters (Table 1; Figs. 8, 9), and the similarity among Bolivian populations of both species has caused confusion about the identity of particular populations (see citations in the synonymy of S. prionotus). Geographic, ontogenetic, or individual var- iation of some characters within both spe- cies, especially the height of the vertebral crest and the number of midbody scale rows (Fig. 8), further clouds the distinc- tions between them. Differentiating the northern populations of S. prionotus from S. caducus is unequivocal and facile based solely on the size of the vertebral crest and on associated meristic counts. However, specimens of S. prionotus from southern Peru and northern Bolivia are more diffi- cult to distinguish from S. caducus. For example, animals from populations of S. prionotus in southern Peru and_ Bolivia have less prominent vertebral crests (hence, higher vertebral scale counts), and generally lower numbers of midbody scale rows, than do specimens from northern populations (Figs. 8, 9); in these respects they are more similar to S. caducus. Nev- ertheless, even accounting for these diffi- culties, a combination of three qualitative characters is sufficient to distinguish prionotus from S. caducus, and the species '° Im my comparisons I have emphasized Bolivian specimens referred to Stenocercus caducus, whereas the type locality is “Paraguay.” I have not fully con- vinced myself that specimens referred to this species from Bolivia, Par aguay, and Argentina are, in fact, all the same taxon. Considerable variation exists in some aspects of coloration and scale characters in these specimens. However, my concept of S. caducus cor- responds to that used in current literature (e.g., Gal- lardo, 1959; Scrocchi et al., 1985; Cei, 1993). Only a thorough study of S. caducus across its range will re- solve this issue. NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA * Cadle may differ in patterns of sexual size di- morphism."! The Form of the Vertebral Crest. De- spite geographic variation in the promi- nence of the vertebral crest, males and fe- males of Stenocercus prionotus have a dis- tinctly projecting serrate vertebral crest ex- tending from the nuchal region to the anterior portion of the tail (Figs. 6, 10, 12). The scales of the crest are strongly trian- gular in lateral view, are flaplike (i.e., they bend easily), project vertically from the dorsum, and are strongly differentiated from the adjacent dorsal scales. Although the crest is somewhat less developed in fe- males, it is prominent in both sexes. Spec- imens from northern Bolivia and southern Peru have a substantially lower crest than specimens from central and northern Peru (Fig. 10). Nonetheless, the form and pro- jection angle of the crest scales is the same as in the northern populations. In contrast, the scales of the vertebral crest in Stenocercus caducus are only mod- erately differentiated from adjacent dorsal scales in being more strongly keeled and mucronate. The crest in S. caducus is only slightly projecting in males (Fig. 10) and even less so in females (Fig. 12); the crest is mainly apparent on the neck and ante- rior body. In S. caducus, the scales of the crest are stiff and prismatic, and the main axis of projection is posterior rather than vertical, as in S. prionotus. Crest height in Stenocercus varies posi- tively with size and thus it is critical to compare similar-sized specimens when documenting differences among popula- tion samples or species. This realization has been critical to differentiating Steno- cercus prionotus from S. caducus in south- "Tam uncertain how Fugler (1983, 1986, 1989) distinguished Bolivian specimens he referred to Sten- ocercus aculeatus and S. caducus. In 1983 and 1986 he referred specimens from Tumi Chucua (Beni, Bo- livia) to S. aculeatus. In 1989 he listed these again, along with ROM specimens from San Marcos Ranch (Beni, Bolivia) identified as S. caducus. Fugler spec- imens from these localities that I have examined are all S. prionotus (see list of paratypes). bo 0 qq Figure 11. Axillary region of Stenocercus caducus showing the posthumeral flap (MCZ 34215). Anterior to the left. The posthumeral flap comprises the heavily outlined scales pos- teroventral to the forelimb. The opening of the posthumeral pocket is the heavily stippled area deep to the flap. Top, Pos- thumeral flap in its normal orientation covering the anterov- entral portion of the pocket. Bottom, the flap deflected ven- trally, with its posterior scales viewed from their tips. Approx- imately <8.5. ern Peru and Bolivia. Large adult males of S. prionotus from southern populations are scarce in collections. For example, al- though only 18 specimens of S. prionotus are available from northern Peru, one half of these are males with SVL =60 mm. In 208 contrast, twice as many specimens each of S. prionotus and S. caducus are available from southern Peru and Bolivia. Yet, only one third of the available specimens of ei- ther species from these areas are males =60 mm SVL, and no males of S. caducus were >72 mm SVL. Figure 10 shows differences in the height of the vertebral crest in a series of males of Stenocercus prionotus from southern Peru and Bolivia compared with similar-sized specimens of S. prionotus from northern Peru and with S. caducus (the largest males of S. caducus studied were 72 mm SVL; see also Fig. 6). The trend toward lower crests in S. prionotus from the southern part of its range is evi- dent, as is the difference between S. cad- ucus and S. prionotus. A comparison of all specimens suggests that the difference in the height of the vertebral crest between S. prionotus and S. caducus males begins to be apparent by approximately 65 mm SVL and becomes pronounced at around 70-75 mm SVL. No males of S. caducus >72 mm SVL were among the specimens examined, despite the availability of a large number of specimens from southern Bo- livia, including a series of 31 specimens (10 males =60 mm SVL) from the vicinity of Santa Cruz. A similar contrast in crest height appears in females of the two spe- cies (Fig. 12). Angulate Temporal Scales. Stenocercus prionotus has two (occasionally three) very strongly keeled, projecting angulate tem- poral scales on each side (Fig. 4). These are much larger than adjacent scales on the head and they are partially or com- pletely separated from the large posterior head scales (parietals, postparietals, and occipital) by one row of small keeled scales (occasionally partially doubled). Stenocer- cus caducus usually has two (occasionally three) angulate temporals that are smaller and less projecting than those in S. prion- otus. In S. caducus the angulate temporals may or may not be larger than adjacent posterior head scales and they are not Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 Figure 12. Size-matched females of Stenocercus prionotus and S. caducus from Bolivia. Top, S. prionotus (USNM 269022, snout-vent length [SVL] 91 mm). Bottom, S. caducus (UTA 38046, SVL 93 mm). Note the subtle difference in crest height between the two specimens and their otherwise similar patterns. strongly differentiated from other posteri- or head scales. Color Pattern of the Gular Region. Many specimens of Stenocercus prionotus have a regular pattern of alternating diag- onal light and dark stripes on the throat. These usually converge closely toward the midline (Fig. 7) and are most easily visu- alized in preserved specimens submerged in alcohol. This pattern consists of a dark stripe beginning at a point on the lower labials in line with, but broader than, the subocular dark bar. The stripe projects posteromedially, gradually fading and blending with the ventral ground color on the neck anterior to the pectoral region. The dark stripe is bordered on either side by a distinct pale stripe. Anteriorly, this se- ries is preceded by another dark and an- other pale stripe. The dark stripes are usu- ally approximately twice as wide as the pale ones, although not always (e.g., the dark stripes are only slightly wider than the pale ones in MCZ 150243). In life the pattern may manifest itself as a series of pale stripes on a darker background (e.g., the “gular area streaked by several light NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ¢ Cadle 209 cream colored lines” in the life colors of the holotype). The gular region appears uniform in many preserved specimens of Stenocercus prionotus, but I suspect this is a preser- vation artifact. Occasional specimens have pale spots in the pectoral region, and oth- ers are essentially unicolor and without ap- parent pattern (again, probably a preser- vation artifact). On the other hand, the throat pattern of Stenocercus caducus is highly variable and irregular. When a distinctive pattern is ,present, it most often consists of light ‘spots rather than alternating stripes (Fig. 13). Cope (1862) described the holotype of S. caducus from Paraguay as having a dark throat that was “light varied” (i.e., variegated, or spotted), and some speci- mens I examined have this pattern (Fig. 13). None of several color descriptions for Argentinian specimens of S. caducus men- tion stripes or spots on the throat. Scrocchi et al. (1985) described living examples as having pale spots in parallel transverse rows in the pectoral region or with pale spots on the abdomen, but did not com- ment on the throat pattern; Gallardo (1959) described the ventral coloration as “pale olive with some scattered pale spots; throat darker”; and Cei (1993) described the venter as “dark brownish with series of rounded pale spots, sometimes anastomos- ing along the length of a median line.” Al- though no authors mention alternating light and dark stripes on the throat in S. caducus, UTA 38046 does have this pat- tern (Fig. 13). But in this specimen the stripes are confined to the lateral edges of the throat (i.e., do not closely approach the midline as in S. prionotus). Apart from the throat pattern, the coloration of S. prion- otus and S. caducus seems to be very sim- ilar judging from descriptions of S. cadu- cus in the literature (Gallardo, 1959: Scrocchi et al., 1985; Cei, 1993). A Possible Difference in Sexual Size Di- morphism. Data presented in Table 1 sug- gests another contrast between Stenocer- Cus prionotus and S. caducus: S. prionotus Figure 13. Gular patterns in Stenocercus caducus. Top, typ- ical throat pattern consisting of light spots on a dark back- ground (BMNH 1927.8.1.163). Bottom, variant pattern consist- ing of stripes confined to the lateral portion of the throat (UTA 38046). Compare to Figure 7. 210 is not dimorphic in the maximum sizes at- tained by males and females, whereas fe- males of S. caducus apparently attain about 20 mm greater SVL than males. However, estimating maximum size is Subb= ject to considerable sampling error so this distinction should be considered to be only provisional. Nonetheless, males of S. prionotus with SVL >80 mm are known from the northern (USNM 193685) and southern (USNM 280250, BMNH 98.6.9.4) portions of its range, even though the three largest males from the largest population sample 3 specimens in FMNH from Puno Department, Peru) had an SVL of 73 mm (this sample includ- ed six adult females with an SVL of 78—89 mm). In contrast, a sample of 39 Stenocercus caducus from Bolivia included 12 adult males, none of which had an SVL >72 mm; in the total sample of S. caducus (N 43) 17 females had an SVL =80 mm (range 80-93 mm). Thus, unless a sys- tematic collecting bias against males exists, the different pattern of ewaal size dimor- phism provides another character distin- guishing S. prionotus and S. caducus. Data presented in Table 1 suggests that other species of the “Ophryoessoides group” may be size dimorphic (S. fimbriatus, S. huancabambae, and ?S. scapularis) or not (S. aculeatus), and either males (S. huan- cabambae) or females (S. caducus and S. fimbriatus) may attain a larger body size. DISTRIBUTIONS OF STENOCERCUS PRIONOTUS AND S. CADUCUS IN EASTERN BOLIVIA The ranges of Stenocercus prionotus and S. caducus approach one another in These sizes are somewhat larger than those pre- viously reported (81 mm; Gallardo, 1959; Scrocchi et al., 1985: Marcus, 1986). Cei (1993) stated that S. caducus reaches only 75 mm SVL in Argentina. Sexes were not given for any individual or aoa specimens in these reports. Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 eastern Bolivia, but do not overlap.!® Cur- rently, the two closest documented locali- ties are, for S. prionotus, near the junction of the Rio Madidi and the Rio Beni (BMNH 98.6.9.4), and for S. caducus, many specimens from the vicinity of Santa Cruz de la Sierra (see above discussion for S. prionotus and Appendix; Fig. 1). Sten- ocercus caducus is also known from the Bolivia—Brazil border in the region of the Serrania de Huanchaca in northern Santa Cruz Department, Bolivia." I am aware of no specimens of Steno- cercus referable to either S. caducus or S. prionotus between the Rio Beni valley and roughly a line connecting Santa Cruz and the vicinity of Serrania de Huanchaca. Southeast of the Rio Beni, the central part of the Beni basin (the Llanos de Mojos) is characterized by flooded savanna grass- lands, palm savannas, swamps, and other habitats that are inundated for significant portions of the year; terra firme forests are restricted to somewhat elevated levees along rivers (Clapperton, 1993: 196; Han- agarth, 1993). Stenocercus prionotus or S. Gadus seem unlikely to occur in this area except possibly in these gallery forests, al- though Fugler (1986) reported S. priono- tus (Be Ophryoessoides aculeatus) in sea- sonally flooded forest during the dry sea- son. Stenocercus prionotus and S. caducus probably are segregated by habitat in Bo- livia and their distabutone may not over- lap. Stenocercus prionotus is associated 'S All references to “Ophryoessoides aculeatus” in Bolivia (e.g., Fugler, 1983, 1986) that I verified have referred to Stenocercus prionotus. However, given the general confusion of species in this complex, some records not traced will have to be checked to rule out the possibility that they do not refer to S. caducus or perhaps some other species of the “Ophryoessoides group,” such as S. fimbriatus or S. scapularis (see Distribution Patterns in Stenocercus prionotus). '4 See the Appendix, UTA 38048. Michael Harvey (personal communication) recently obtained speci- mens of Stenocercus caducus at El] Refugio, a lowland locality at the southern end of the Serranfa de Huan- chaca (14°44'S, 61°O1’W). NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle with upper Amazonian and lower montane rainforests with annual rainfall greater than 2,000 mm in both Peru and Bolivia. On the other hand, confirmed localities of S. caducus are within the physiographic domain broadly referred to as chaco, in- cluding a mixture of dry forests, palm sa- vannas, gallery forests, deciduous forests, and ecotonal areas (Scrocchi et al., 1985; Marcus, 1986; Cei, 1993). Short (1975) and Parker et al. (1993) described the di- versity of chaco habitats. Gallardo (1979: table 12.1) listed S. caducus as a species “basically restricted to the chaco.” Average annual rainfall in this area is less than 1,000 mm. Stenocercus caducus is known from Parque Nacional Noel Kempff Mer- cado and vicinity in Bolivia (see footnote 14; Harvey, 1998). This area is character- ized by a complex mixture of habitat types, including deciduous forests and cerrado enclaves, and with an annual rainfall of 1,400-1,500 mm (Killeen, 1998). Harvey (1998) encountered S. caducus at granitic outcrops covered by semideciduous forests and more open habitats. The herpetofauna of this site is a mixture of species that are typical of Amazonian and of chaco envi- ronments (Harvey, 1998; personal obser- vations). The range of Stenocercus caducus ex- tends outside the strictly defined chaco re- gion (see Short, 1975, and Clapperton 1993, for discussion) on the southeastern edge of its range east of the Rio Paraguay and in the Andean foothills of southern Bolivia and northern Argentina (Fig. 1). Harvey (1997) reported S. caducus from “subtropical wet forests” (1,150—2,050 m elevation) in southern Bolivia. He charac- terized S. caducus as a “Chacoan species that invade|s|] the Andean foothills . . . [in- cluding] those distributed within the Gran Chaco or that occur in dry forests sur- rounding the Gran Chaco” (Harvey, 1997: 35). The montane wet forests (yungas) of this area are restricted to ridges high enough for cloud formation during much of ie year (generally >1,500 m elevation), and they are surrounded by deciduous dry 211 forest (Schulenberg et al., 1997). The cli- mate of this area is generally dry and it receives only about 1,200 mm of rainfall per year (Holst, 1997). The transition between the wet rainfo- rests of Peru and northern Bolivia (range of Stenocercus prionotus) and the chaco habitats (range of S. caducus) occurs in a very broad ecotone consisting of savannas, evergreen shrublands, and gallery forests of the Beni basin and Rio Mamore drain- age, from which no specimens of either S. prionotus or S. caducus have been report- ed. The piedmont forests of the Andes be- tween the known ranges of S. prionotus and S. caducus, which are wetter than ad- jacent lowland forests because of the mod- erating effect of the Andes, provide one potential route for contact or overlap of their ranges. The eastern distributional limits of Sten- ocercus caducus along the Bolivia—Brazil frontier are not well ‘aaclen stood. I am un- aware of verified records from Brazil, al- though the species does occur close to the Brazilian border in the vicinity of the Ser- rania de Huanchaca in Parque Nacional Noel Kempft Mercado. Some references to “Stenocercus caducus” from western Brazil (e.g., Mato Grosso State; Cope, 1887; Boulenger, 1903) likely refer instead to an undescribed species very similar to S. caducus (P. FE. Vanzolini and E. E. Wil- liams, personal communication; personal observations). However, the ranges of S. caducus and the undescribed species in eastern Bolivia—Paraguay and western Bra- zil are not well defined; the two species may be separated by the seasonally inun- dated savannas of the pantanal. Addition- ally, few specimens of S. caducus appar- ently exist from the chaco of northwestern Paraguay, although Aquino et al. (1996) re- ported specimens from Parque Nacional Defensores del Chaco (approximately 20°30'S, 60°20'W), as well as other Para- guayan localities in more mesic regions east of the Rio Paraguay. bo bo 1 IS THE DISTRIBUTION OF STENOCERCUS ACULEATUS DISJUNCT? In the process of diagnosing Stenocercus prionotus I reviewed the characters and distribution of S. aculeatus. In addition to variation in some characters of uncertain significance, some aspects of the distribu- tion of S. aculeatus seem peculiar (Fig. 2). First, the distribution of S. aculeatus ap- pears to be disjunct between northern Peru and eastern Ecuador. Although the type locality is in northern Peru (Moya- bamba, San Martin Department), most specimens are from eastern Ecuador (Fig. 2). The two areas from which specimens are known (Fig. 2) are separated by a broad geographic gap through which courses the main tributary of he upper Amazon, the Rio Marafion. Neither S. acu- leatus nor any other species of Stenocercus was obtained during herpetofaunal surveys in northern Loreto Department, Peru (Duellman and Mendelson, 1995), north- ern Amazonas Department, Peru (Rio Ce- nepa and Rio Santiago; J. E. Cadle and R. W. McDiarmid, unpublished data), or dur- ing a rapid biological assessment of the Cordillera del Condor region of southeast- ern Ecuador and northern Peru (Schulen- berg and Awbrey, 1997a). Stenocercus acu- leatus is known from many localities in ad- jacent regions of Ecuador. Second, all Ecuadorian localities for Stenocercus aculeatus are in the drainages of the Rio Pastaza and the Rio Curaray. No specimens are known from the Rio Napo drainage just to the north, even though no recognized physiographic or faunal break seems to separate the Rio Napo drainage from the Rio Curaray drainage. However, all localities that have been sampled comprehensively from the Rio Napo are on the left (northern) bank of the river (e.g., Duellman, 1978; Vitt and De la Torre, 1996; unpublished list from a large collection from the Jatun Sacha Bi- ological Station assembled and under study by Gregory Vigle). The absence of Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 fimbriatus and S. prionotus has concomi- S. aculeatus from Santa Cecilia (Duell-_ man, 1978) is probably real rather than sampling error, given the intensity of col- lecting over several years at the site. Thus, S. aculeatus possibly does occur on the right (south) bank of the Rio Napo and will be recorded once large collections are made there. The apparent geographic disjunction of | Stenocercus aculeatus between northern | Peru and eastern Ecuador may correspond | to some character differences among sam- | ples that should be studied more thor- | oughly (Cadle, unpublished data). For ex-_ ample, Peruvian specimens of S. aculeatus have very deep postfemoral pockets (Type 5) in both sexes, whereas the postfemoral pockets are more weakly developed in specimens from Ecuador (Type 2) Oo) lM both sexes). Ecuadorian specimens also appear to have more scales in the vertebral row and fewer subdigital scales on the fourth toe than do Peruvian specimens. All of these impressions are based on small sample sizes (Appendix). The significance of these differences is unclear without a more detailed study of variation among populations of Stenocer- cus aculeatus. However, one possibility is that two or more species are represented in specimens currently referred to S. acu- leatus, in which case the distributions of individual taxa may be not be contiguous. This is analogous to the previous confusion of S. fimbriatus and S. prionotus with S. aculeatus. Taxonomic recognition of S. tantly reduced the geographic distribution understood for S. aculeatus. Consequently, a more comprehensive systematic analysis of S. aculeatus with special reference to a comparison of Ecuadorian and Peruvian populations is warranted. If two species are recognized, the name Liocephalus an- gulifer Werner (1901) is available for the’ Ecuadorian populations. KEY TO SPECIES OF THE “OPHRYOESSOIDES GROUP” OF STENOCERCUS Because of the general confusion about the species Coneideres! herein (e.g., see the: NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA * Cadle synonymy of Stenocercus prionotus ), | pro- vide the following key as a guide for iden- tifications. The key will work for those spe- cies of Stenocercus in Peru or Bolivia with keeled ventral scales, enlarged posterior head scales, and one row of moderately to greatly enlarged supraoculars (“Ophryoes- soides group” as used herein). I have also included the three other currently recog- nized species having these characteristics, S. erythrogaster (Hallowell), S. dumerilii (Steindachner), and S. tricristatus (Du- méril), although these are not known from Peru or Bolivia and are unlikely to occur there. Character and distributional data in the key for S. dumerilii and S. tricristatus follow Avila-Pires (1995). I also include in the key an undescribed species with keeled ventrals and enlarged head plates and supraoculars from Ama- zonas Department, Peru, butel am un= aware of other undescribed species of the “Ophryoessoides group” from Peru or Bo- livia. However, an undescribed species similar to Stenocercus caducus (but lacking a posthumeral flap) is known from western Brazil (Mato Grosso) and is not included in the key. Additional study of S. iridescens from the Pacfic lowlands of Peru and Ec- uador is needed (Cadle, 1998: footnote 4) and, as indicated above, a thorough mod- ern study of variation in S. aculeatus (Am- azonian Ecuador and Peru) is also war- ranted. Other undescribed species may re- side within either of these named taxa. The key will permit identification of all Pe- ruvian and Bolivian taxa previously con- fused with S. aculeatus (e.g., Dixon and Soini, 1975, 1986 [S. fimbriatus]; Fugler, 1983, 1986, 1989 [S. prionotus]) and S. ir- idescens (e.g., S. huancabambae and S. limitaris; see Cadle, 1991, 1998). The key also should work for Ecuadorian species, with the caveat that I have paid less atten- tion to Ecuadorian Stenocercus except as necessary in conjunction with work on Pe- ruvian species. Of the species covered, only S. aculeatus, S. iridescens, and S. lim- itaris are definitely known from Ecuador. 213 The key should be viewed as a means of identifying a set of phenotypically similar, but not necessarily closely related, species within Stenocercus in the broad sense. All other species of Stenocercus in Peru and Bolivia have smooth (or at most only very weakly keeled) ventrals and more frag- mented supraoculars and head plates; see Fritts (1974), Frost (1992), and Cadle (1991, 1998) for discussion and _illustra- tions. Many of these species also have granular scales on the body or posterior surface of the thigh, neither of which is present in species covered by the key. The keys and discussions in Fritts (1974) and Cadle (1991, 1998) are useful for identi- fying these other species. The key assumes familiarity with char- acters of the mite pockets, head scales, and neck folds and crests outlined in Cadle (1991) (see also Materials and Methods). In most cases I have used characters that show minimal sexual dimorphism so that specimens of either sex can be identified; exceptions are noted. It is useful to keep in mind that, in most species of Stenocer- cus, scales of juveniles are more promi- nently keeled than in adults, even when the corresponding scales of adults, such as head scales and dorsal body scales, are smooth. Instances of possible confusion in the key are indicated. The extent of de- velopment of posthumeral and postfemor- al mite pockets varies according to sex and size in many species of Stenocercus, al- though such variation seems less extensive in this set of species than in many others; I have indicated the range of variation in- cluding juveniles and adults of both sexes in the key. Summary geographic distribu- tions are given for each species as a rough guide to known occurrences. However, these should be used cautiously as ancil- lary information in identifying specimens because distributions of species are some- times poorly circumscribed. For greatest utility the key should be used in conjunc- tion with illustrations herein and in Cadle (1991, 1998) and Avila-Pires (1995). 2 a to icy) 14 Canthal and supraciliaries forming a pro- nounced crest that ends in an enlarged, erect, postsupraciliary that may be dis- tinctly pointed or blunt. Posthumeral and postfemoral mite pockets absent (Type 1 rial loxovclay Tua uANNKOS)) 2 Canthals and supraciliaries not forming a pronounced crest; no enlarged, erect post- supraciliary. Posthumeral pocket absent (Type 1) to deep (Type 4). Postfemoral pocket absent (Type 1) to deep (Type 5) Enlarged postsupraciliary distinctly pointed. Two enlarged scales above ear opening. Tibia approximately equal to thigh length pan Stee oes Ne Stenocercus dumerilii (Steindachner) (northeastern Para, Brazil) Enlarged postsupraciliary blunt. No enlarged scales above ear opening; tibia distinctly SIorausre tdaenel (Hove oe EA ENS | Stenocercus tricristatus (Duméril) (known only from the holotype, probably from the state of Minas Gerais, Brazil) Superciliary scales projecting laterally shelf- like above the orbit in adults, rectangular in dorsal view.'!° Midbody dorsal scale rows more than 55 (59-70). Postfemoral pocket well developed (Type 3 or 5) __...--.-------- Bet ee RLS tel s Stenocercus scapularis (Boulenger) (intermediate elevations on the Andean slopes of central and southern Peru; known elevations greater than 1,000 m) Superciliary scales not projecting laterally, the anterior ones elongate, strongly over- lapping. Midbody dorsal scale rows fewer than 55 (30-53). Postfemoral pocket vari- ANS (kyae Il; 2, O; Cir &) 2 4 Posthumeral mite pocket deep (Type 4) and with an associated scaly flap extending from its anteroventral border and partally COMCE AMIN GPity sete ate see ee 5 Posthumeral mite pocket absent to deep (Type 1, 2, 3, or 4) but without an asso- Clatealet apy piesa lean eien mas MET eee A hol 5S 6 Vertebral crest strongly projecting in both sexes, serrate, extending from the nuchal region to the proximal portion of the tail; its individual scales triangular, flaplike. Throat often with alternating oblique dark and light stripes. Two enlarged, strongly keeled and projecting angulate temporal 'S The superciliaries in juveniles of Stenocercus sca- pularis have a more typical shape. The extent of shelf-like projection and change to a more rectan- gular shape seem positively cor ated with body size Aaa thus develop with age. The number of dorsal scale rows and the development of the postfemoral pocket are useful clues for subadults. Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 scales om each side ee ee Stenocercus prionotus Cadle (rainforested lowlands and Andean foothills of eastern Peru and northern Bolivia) Vertebral crest low, nonprojecting in both sexes (slightly higher in males), evident mainly on neck and anterior body; its in- dividual scales prismatic and lying more or less flat. Throat pattern variable, but usu- ally consisting of light spots on a darker background when evident. Angulate tem- poral scales keeled, but not greatly en- largedsand monprojecting seen myftiea See FS Ree ON Stenocercus caducus (Cope) (deciduous woodlands and ecotonal areas of southern Bolivia, northern Argentina, and the chaco of Paraguay) 6. A fringe of enlarged fimbriate scales on the distal posterodorsal surface of thigh. Sev- eral longitudinally oblique rows of large, strongly keeled scales on shank.'® Seales between vertebral and dorsolateral crests smooth or occasionally very weakly keeled. Let Dee Dds BS Stenocercus fimbriatus Avila-Pires (lowlands of eastern Peru and central western Amazonian Brazil) No fringe of fimbriate scales on thigh or strongly keeled oblique scales on shank. Scales between vertebral and dorsolateral crests moderately to strongly keeled at least posteriorly; dorsolateral crest may be weakly developed, but dorsal scales. still strongly keeled. 420. 1.0 0 Mie seit et tad vere i 7. Posthumeral mite pocket variable (Type 1, 2 3, or 4). Postfemoral mite pocket variable (Type 1, 2, 3, or 5). Head scales smooth or keeled. Angulate temporal scales keeled, may be projecting and bladelike. Internasals usually 4 or more (occasionally 3, never 2), often irregular in pattern and Shape wit! ano As, Wee ehtded deel ereecermet 8 Posthumeral mite pocket absent or weakly developed (Type 1 or 2). Postfemoral mite pocket absent (Type 1). Head _ scales smooth. Angulate temporal scales smooth; none bladelike and projecting. Two polyg- onal internasals in contact on the midline, each broader laterally than medially. Bb neta eatin. ee Stenocercus iridescens (Giinther) (Pacific lowlands and intermediate elevations of western Ecuador and northwestern Peru) 8. One to 3 strongly keeled, but nonprojecting, angulate temporal scales in line with the superciliary row between the lateral tem- ‘© The fimbriate scales form a projecting fringe on the distal portion of the thigh. Both the fimbriate scales and the oblique Realeer on the shank are rela- tively more prominent in juveniles than adults. NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA * Cadle porals and the posterior dorsal head scales. Two subequal canthal scales on each side. Head scales keeled, at least posteriorly 9 Two projecting bladelike angulate temporals in line with superciliary row. A single can- thal on each side (rarely, 2 are present but in that case 1 is much larger than the oth- er). Head scales smooth or keeled. _..... 11 9. Posthumeral pocket moderately developed in males (Type 2 or 3), absent in females (Type 1). Postfemoral pocket absent in fe- males (Type 1), moderate to deep in males (Type 3 or 5). Anterior gular scales weakly (co) Gaveredhy Ievslkevl ee ee peste) conte Stenocercus erythrogaster (Hallowell) (northern Colombia) Posthumeral and postfemoral pockets deep in both sexes (Types 4 and 5, respective- ly).!7 Anterior gular scales smooth to weak- IWakeele diye iw te wees Sol eh Pat ee ee 10 10. Interparietal indistinct, parietal eye not visi- ble. Three occipitals. Dark subocular bar absent. Three angulate temporals separat- ed from large posterior head scales by a row of tiny scales __. Stenocercus new species (known from a single specimen [Appendix] from the inter-Andean valley of the Rio Maranon near Balsas, Amazonas Depart- ment, Peru) Interparietal distinct, parietal eye visible. Two occipitals. Dark subocular bar pre- sent. One angulate temporal much larger than others and in contact with at least 1 other enlarged posterior head scale. ___. PEER athe Stenocercus limitaris Cadle (intermediate elevations [600—2,200 m] of the Andes on the Pacific versant of southwest- ern Ecuador and northwestern Peru) 11. Head scales smooth to slightly wrinkled in adults; weakly keeled, wrinkled, or rugose in juveniles. Prominent dorsolateral crest on body from neck to base of tail and con- tinuous with both supra-auricular crest and antehumeral crest. Postfemoral pock- et moderate to deep (Type 2, 3, or 5). ae Tu Stenocercus aculeatus (O’Shaughnessy) (rainforested lowlands and intermediate ele- vations of northern Peru adjacent to the Andes and in eastern Ecuador) Head scales strongly keeled or multicarinate in juveniles and adults. Dorsolateral crest, when present, weak and restricted to neck and anterior body. Postfemoral pocket GRES|O) (CLIN GONE Bo cee densa NRTA Stenocercus huancabambae Cadle '7 An undescribed species in the next couplet of the key is known only from a single adult male. The dis- tributions of species in couplets 9 and 10 should be used as ancillary data for identification. 215 (dry inter-Andean valleys of the upper Rio Marafion in Cajamarca and west central Amazonas departments, northern Peru) ACKNOWLEDGMENTS Loans and other assistance were facili- tated by Linda Ford, Darrel Frost, and Charles W. Myers (AMNH); E. Nicholas Arnold and Colin J. McCarthy (BMNH); John Wiens (CM); Cassy Redhed, Alan Resetar, and Harold Voris (FMNH); Wil- liam E. Duellman, Christopher J. Raxwor- thy, and John E. Simmons (KU); Frank Burbrink and Douglas Rossman (LSUMNS): Ross MacCulloch and Robert W. Murphy (ROM); Roy W. McDiarmid, Steven W. Gotte, W. Ronald Heyer, and Robert P. Reynolds (USNM); and Jona- than Campbell and Michael B. Harvey (UTA). Victor Morales permitted me to ex- amine a specimen of Stenocercus fimbria- tus in his care. I am grateful to Wade C. Sherbrooke for providing copies of his field notes and other information on spec- imens he collected. I owe a great debt to the late Ernest E. Williams, who was ex- tremely generous with discussion, notes, il- lustrations, and encouragement. Williams, Paulo E. Vanzolini, and Richard Etheridge long ago distinguished Stenocercus prion- otus and two other species I described (S. huancabambae and S. limitaris) but kindly let my work on the group unfold with their gracious consent. Vanzolini supplied a copy of a portion of Balzan (1931), helped interpret Balzan’s localities, and pointed out Métraux’s work to me. I am indebted to several people for the special efforts they made in tracking down information about particular collections: Bruce Patter- son (FMNH) supplied information on the collections of Colin Sanborn and Hilda Heller from Puno Department, Peru; Alan Resetar and Cassy Redhed (FMNH) dug into the Schmidt archives and found ad- ditional information about Heller’s collec- tion; Charles W. Myers (AMNH) did the same for Harvey Bassler’s journeys in northern Peru and provided the base map used to prepare Figure 1; Robert S. Voss 216 checked AMNH sources for information on Keays’s Peruvian localities; and Colin J. McCarthy (BMNH) clarified the confusion about P. O. Simons’s “Palca” locality, if not the locality itself. Robin Andrews, Tom Jenssen, and A. Stanley Rand discussed as- pects of geographic variation and the uses of vertebral crests in S. prionotus with me. Laszlo Meszoly drew Figures 4, 5, and 11. For comments on the manuscript I thank Richard Etheridge and Charles W. Myers. The research was supported in part by a faculty grant from the School of Arts and Sciences of Harvard University; publica- tion costs were supported by the Colles Fund of the MCZ. APPENDIX: SPECIMENS EXAMINED Institutional abbreviations are as fol- lows: AMNH American Museum of Natural History, New York ANSP Academy of Natural Sciences of Philadelphia BMNH The Natural History Muse- um, London CM Carnegie Museum of Natural History, Pittsburgh, Pennsyl- vania FMNH The Field Museum, Chicago KU Natural History Museum, University of Kansas, Law- rence LSUMNS Louisiana State University Museum of Natural Science, Baton Rouge MCZ Museum of Comparative Zo- ology, Harvard University, Cambridge ROM Royal Ontario Museum, To- ronto USNM National Museum of Natural History, Washington, D.C. UTA University of Texas at Arling- ton Bracketed information was inferred from sources listed in the Materials and Methods. For specimens of other species of the “Ophryoessoides group” examined Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 (S. huancabambae, S. iridescens, and S. limitaris), see Cadle (1991, 1998). Bold- face numbered localities 1—4 are known or suspected areas of sympatry between the species listed and Stenocercus prionotus. They correspond to numbered localities in Figures 1 and 2 and in the text discussion. Stenocercus aculeatus ECUADOR: Prov. MORONA-SANTIAGO: Chiguaza [ea. 1,000 m; 01°59’S, 77°58’W] (USNM 200882-84). [PProv. Napo/TuncurAnuA]: Llanganates area!’ (FMNH 23527). Prov. Pastaza: Rio Pastaza, Abita- gua [01°23'S, 78°05'W] (FMNH 25803-05, 26892, 28011, 28057 [=17 specimens]). Rio Pastaza, Alpay- acu [01°28'S, 78°07'W] (FMNH 3926-27: MCZ 8081). Canelos [01°35'S, 77°45’W; 530 m] (MCZ 38530). Montalvo, Rio Bobonaza [314 m; 02°04’S, 76°58'W] (USNM 200892). [PPRov. PASTAZA]: Banos, Mera Trail [? = between Banos and Mera"; approx- imately 01°30'S, 78°10'W] (FMNH 28012). PERU: [Depto. LA LipertaD]: E Peru, Pampa Seca, Rio Mixiolla [=Rio Mishollo] Valley, Upper Huallaga region, 4300 ft [2, 1,311 m; approximately 08°16'S, 76°58'W]?) (AMNH 57085). [DEPpro. Lor- 18 T have been unable to localize this. The Cordil- lera de los Llanganates is a high range (to >4,500 m) in the Cordillera Oriental north of the Rio Pastaza (Paynter, 1993). The locality may refer to lower ele- vations in this range. '° Bafios is a famous collecting locality at the foot of Volcan Tungurahua at 1,820 m. That would be an altitudinal record for Stenocercus aculeatus. I inter- pret the locality as stated in the FMNH catalogues — as being on the trail between Bajos and Mera, which is at 1,160 m. See Brown (1941) and Chapman (1926) for discussion. °° Harvey Bassler collected Stenocercus for the AMNH at two localities on the Rio Mixiolla (=Rio Mishollo): Pampa Seca and La Pinita (see Stenocer- cus fumbriatus), as listed in AMNH catalogues. The Rio Mishollo originates in southeastern La Libertad Department, flows eastward, and joins the Rio Hual- laga in southwestern San Martin Department. The elevations given for these localities, 1,067 m and 1,311 m, indicate that they lie in the narrow stretch of the valley that straddles the boundary between La Libertad and San Martin departments (departmental | maps produced by the Instituto Geografico Nacional, _ Lima). I identify these localities as Pampaseca and | Piflita, respectively, in extreme southeastern La Lib- ertad Department, as indexed by Stiglich (1922). — Both localities are in Ongén District and the coor- dinates given are those for the town of Ongon. Stig- lich (1922) states that Pifiita is a small village on the © Quebrada Pedernal, a left tributary of the Rio Mish- ollo. Apart from indicating that Pampaseca is a farm, | NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle ETO]: NE Peru, Front Range between Moyabamba and Cahuapanas, 3000 ft [915 m; approximately 05°37'S, 77°00'W] (AMNH_ 57083). Northeastern Peru: Icuta on Balsapuerto-Moyabamba trail, 3500 ft [1,067 m; 05°58’S, 76°40'W; given as “Icuto” or “Icu- to Cuesta” by Lamas, 1976] (AMNH 56413). Stenocercus caducus BOLIVIA: No SPECIFIC LOCALITY: _BMNH 1946.8.29.76 (holotype of Leiocephalus bolivianus Boulenger); CM 4583-84. DEpTo. CHUQUISACA: Sud Cinti, trail from Rinconada Bufete to El] Palmar [1,170-2,000 m; approximately 20°50'S, 64°21'W] (UTA 39102). DErro. SANTA Cruz: Buena Vista, ca. 500 m [17°27'S, 63°40’W] (MCZ 20625-26, 29023; FMNH 16165, 21486, 21511; BMNH 1927.8.1.163— 164: CM 4527, 4550-51, 4558, 4587-88, 4605, 4607, 4616, 4626, 4634-36, 4641). Las Yuntas [=Las Jun- tas:2! 18°38'S, 63°08’W] (CM 970). Provincia Chiqui- tos, Santiago (Serrania and nearby), 700-750 m [18°19’S, 59°34"'W] (FMNH_ 195983). [| Provincia | Chiquitos, Canton El] Cerro, Finca Dos Milanos, 17°27'30"S, 62°20'00"W (UTA 38046). Provincia Sara, eastern Bolivia, 600 m [17°27'S, 63°40'W] (BMNH 1907.10.31.7-8). Provincia Sara, Santa Cruz de la Si- erra [17°48'S, 63°10’W] (CM 966, 969, 13018). Prov- incia Sara, Rfo Surutu W of Buena Vista [17°24’S, 63°51'W] (CM 4590). Provincia Sara, Rio Colorado [17°38'S, 63°54’W] (CM 4598). [Provincia] Velasco, Inselbergs near Florida [14°38’S, 61°15’W] (UTA 38048). [DEepro. Tara]: Villa Montes [21°15'S, 63°30'W] (MCZ 28634). Misi6n San Francisco [21°15’S, 63°30'W] (BMNH 98.7.7.5; specimen col- lected by Alfredo Borelli, whose San Francisco = Vil- la Montes fide Paynter, 1992). PARAGUAY: [DEpTO. CAAaGuAzv]: Pastoreo [ap- proximately 25°23'S, 55°52’W] (MCZ 34214—-15). [DEPro. CENTRAL]: Asuncién [25°16’S, 57°40'W] (FMNH 9496). Colonia Nueva Italia [25°37’S, 57°30’W] (FMNH 42281). Stenocercus erythrogaster COLOMBIA: [DEptTo. MAGDALENA]: Rio Frio [30-450 m; 10°55'N, 74°10’W] (MCZ 29707). Santa Marta Mountains [approximately 10°50'N, 73°40'W] (MCZ 11303). Rio Toribio, Hacienda “Papare,” sec- ond river on road from Cienaga to Santa Marta [11°03’N, 74°14’W] (FMNH 165153). [DEPTo. SAN- TANDER]: San Gil [1,095 m; 06°33’N, 73°08’W] (ANSP 24136, MCZ 36877). Stiglich (1922) gives no further information about its location. *1 The specimen was collected in November or De- cember 1913 by José Steinbach, who collected at a locality known as “Las Juntas” during that same pe- riod (Paynter, 1992). The two localities are assumed to be the same. The variant spelling “Yuntas” does not appear in any sources consulted. bo 7 Stenocercus fimbriatus PERU: No SPECIFIC LOCALITY: (FMNH 56070). DeEpro. HUANUCO: ca. 35 km NE Tingo Maria, Heda. Santa Elena, ca. 1000 m [approximately 08°57’S, 76°02'W] (LSUMNS 26966-67). Approximately % mile E Universidad Agraria de La Selva, Tingo Maria, vicinity of Rio Huallaga [3, 09°18’S, 75°59'W], USNM 193684. [DEpro. LA LIBERTAD]: E Peru, La Pinita, Rio Mixiolla, tributary of upper [Rio] Hualla- ga, 3500 ft [1,067 m; approximately 08°16'S, 76°58’W; see footnote 20] (AMNH 56797-98). [DEPTo. Lor- ETO]: E. Peru, Contamana, Ucayali River valley [134 m: 07°15’S, 74°54'W] (AMNH 56803). E Peru, E of Contamana on trail to Contaya, 700 ft [213 m; ap- proximately 07°15’S, 74°54’W]| (AMNH 56781-82). E Peru, Pampa Hermosa, mouth of Rio Cushabatay, 500 ft [152 m] [1, 07°12’S, 75°17'W] (AMNH 56788, 56790-92, 56794—96, 56801—02). Mishana, Rio Na- nay, Estacion Biolégica Cauicebus, 150 m [03°53’S, 73°27'W] (USNM 222377). Mishuana [=Mishana; 150 m, 03°53’S, 73°27'W]| (KU 212628). DEPTO. Ma- DRE DE Dios: Pakitza Station [Rio Manu], Manu Na- tional Park [4, 11°56’S, 71°17’W] (Victor R. Morales 18235). Drpro. UcayYALi: Rio Curanja, Balta, ap- proximately 300 m [approximately 10°08'S, 71°13'W] (LSUMNS 17519, 25402-04, 26720-23). Alto [Rio] Purtis, Alto [Rfo] Curanja, Igarape Champuiaco [9°34'S, 70°36’W] (MCZ 61226). Peru/Brazil frontier, Utoquinia Region, 1000 ft. [305 m; approximately 08°00'S, 74°00’W]|22 (AMNH 56789, 56799-800). Stenocercus scapularis** PERU: No SPECIFIC LOCALITY: (FMNH 56444). [DEpPTo. JUNiN]: Chanchamayo, 1200 m_ [approxi- mately 11°03'S, 75°47’'W] (FMNH 40608—11). Pere- ne, 1200 m [10°58’S, 75°13’W] (MCZ 49580-81). Tarma, Chanchamayo, 1300 m [11°25'S, 75°42’W] (FMNH 45522). [DEpTo. PUNO]: Sagrario, Rio Qui- tin [approximately 1,020 m; 13°55’S, 69°41'W] ~ The region referred to is north to northeast of Pucallpa. The variant spellings Utoquinia, Utoquinea, and Uroquinea are in the literature and are applied to a right-bank tributary of the Rio Ucayali, a village on the Rio Ucayali, and an airstrip on the Rio Uto- quinia near the Brazilian border. The entire region is less than 500 m in elevation except for a small raised area near the Brazilian border that attains nearly 800 m and that is apparently the source of the Rio Uto- quinia. 23 The occurrence of Stenocercus scapula ris at Rur- renabaque, El Beni Department, Bolivia, as reported for two specimens in the AMNH (Burt and Burt, 1931: 273) is apparently based on a misidentification. These specimens are probably either S. prionotus (most likely) or S. caducus (see Distribution Patterns in Stenocercus prionotus). 218 (FMNH 40408). “Camp 4” [between Santo Domingo and La Pampa; approximately 13°44’S, 69°37'W]*! (FMNH 40409). Juliaca, Lake Aracona, 16,600 ft. [shipping point only; correct locality is on the right bank of the Rio Inambari, 1,830 m, 13°30'S. 70°00'W |? (AMNH 1701). 4 According to the field catalogue in the FMNH Mammal Division the collector, Colin Sanborn, was in Santo Domingo on 20 October 1941 and in La Pampa on 23 October (see also notes in Stephens and Traylor, 1983). The specimen FMNH 40409 was col- lected 21 October, and thus “Camp 4” is assumed to be between these points. > The specimen was collected in 1900 by H. H. Keays, who collected many mammals and other ver- tebrates in southern Peru, primarily for the American Museum of Natural History. It is clear that most of the specimens labeled with the locality “Juliaca” (a town on the Peruvian altiplano near Lake Titicaca) actually came from farther north in the Rio Inambari valley. Allen (1900: 219; 1901: 41) provides the fol- lowing information: The Museum has recently received two small col- lections of mammals made by Mr. H. H. Keays, at Juliaca, in southeatern Peru, a little to the west- ward of Lake Titicaca. Mr. Keays writes: “Our camp is situated in the loop of the ‘Inambary River. The country is very broken, with deep narrow can- ons, and is cover ed with a dense undergrowth of shrubs and vines, with here or there a palmetto or a cedar rising above the surrounding vegetation.” He gives the altitude as 6000 feet [1,830 m], and the position as latitude 13°30’ S., longitude 70° W. . it is necessary to correct a misleading statement in my former paper in respect to the locality where the ... collections were made. Mr. Keays’s post- office address was Juliaca, and through lack of ex- plicit information, it was inferred that the Inca Mines, where he collected, were in the immediate vicinity of Juliaca ... the Inca Mines are situated about 200 miles northeast of Juliaca, on the east side of the Andes, on the Inambary River, a trib- utary of the Amazon, and at a much lower altitude than Juliaca. The altitude and geographical position were correctly given in the fannie paper, but in place of Juliaca,, ... read Inca Mines. Keays’s information quoted by Allen places the local- ity on the right bank of the Rio Inambari in the foot- hills of an outlying Andean spur separating the Rio Inambari from upper tributaries of the Rio Tambo- pata. I have not located a Lake Aracona and suspect that this is an error for Lake Aricoma, a high Andean lake on the route between Juliaca and the location of Keays’s camp. However, it is not at all clear why this name is associated with the locality. No notes or cor- respondence of Keays are in the AMNH mammal department archives for further clarification (R. S. Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 Stenocercus sp. PERU: Deprro. AMAZONAS: 17 km ENE Balsas [06°49'S, 78°00'W] (ROM 16458). LITERATURE CITED ALLEN, J. A. 1900. On mammals collected in south- eastern Peru, by Mr. H. H. Keays, with descrip- tions of new species. Bulletin of the American Museum of Natural History, 13: 219-227. . 1901. On a further collection of mammals from southeastern Peru, collected by Mr. H. H. Keays, with descriptions of new species. Bulletin of the American Museum of Natural History, 14: 41-46. AQUINO, A. L., N. J. SCOTT, AND M. MOTTE. 1996. Lista de anfibios y reptiles del Museo Nacional de Historia Natural del Paraguay (Marzo, 1980— Setiembre, 1995), pp. 331-400. In O. R. Marti- nez (ed.), Colecciones de Flora y Fauna del Mu- seo Nacional de Historia Natural del Paraguay. San Lorenzo, Paraguay: Museo Nacional de His- toria Natural del Paraguay. AVILA-PIRES, T. C. S. 1995. Lizards of Brazilian Ama- zonia (Reptilia: Squamata). Zoologische Verhan- delingen, Leiden, 299: 1—706. BALZAN, L. 1931. Viaggio di Esplorazione nelle Re- gioni Centrali del Sud America. Milan, Italy: Fra- telli Treves. xix + 368 pp. BOULENGER, G. A. 1898. A list of the reptiles and batrachians collected by the late Prof. L. Balzan in Bolivia. Annali Museo Civil Storia Naturali di Genova, series 2, 19: 128-133. . 1901. Further descriptions of new reptiles collected by Mr. P. O. Simons in Peru and Bo- livia. Annals and Magazine of Natural History, series 7, 7(42): 546— 549. . 1903. List of the batrachians and reptiles col- lected by M. A. Robert at Chapada, Matto Gros- so, and presented by Mrs. Percy Sladen to the British Museum (Percy Sladen Expedition to Central Brazil). Proceedings of the Zoological Society of London, 1903, II(1): 69-70. BROWN, F. M. 1941. A gazetteer of entomological sta- tions in Ecuador. Annals of the Entomological Society of America, 34(4): 809-851. Burt, GC. E., AND M. D. BuRT. 1931. South American lizards in the collection of the American Muse- um of Natural History. Bulletin of the American Museum of Natural History, 61: 227—395. CADLE, J. E. 1991. Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from orth ern Peru: new species and comments on rela- tionships and distribution patterns. Proceedings Voss, personal communication). Stephens and Traylor (1983) discuss “Inca Mines” under their entry for Santo Domingo, a settlement on the Limbani—Aster- illo road, which was apparently close to the Inca Mines. See also Vaurie (1972). NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle 219 of the Academy of Natural Sciences of Philadel- ae 143: 1-96. 1998. New species of lizards, genus Steno- cercus (Iguania: Tropiduridae), fron western Ec- uador and Peru. Bulletin of the Museum of ea Zoology, 155(6): 257-297. ee Meelo93: Reptiles del Noroeste, Nordeste y Taste de la Argentina. Herpetofauna de las Selvas Subtropicales, Puna y Pampas. Torino, Italy: Mu- seo Regional di Scienze Naturali. 949 pp. CHAPMAN, F. M. 1926. The distribution of bird-life in Ecuador. 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WILLIAMS, E. E., AND A. S. RAND. 1977. Species rec- ognition, dewlap function, and faunal size. Amer- ican Zoologist, 17: 261-270. ON THE SUBFAMILY XYLOPHAGAINAE FAMILY PHOLADIDAE, BivatviA, MoLtusca) RUTH D. TURNER HARVARD UNIVERSITY mote ~ VOLUME 157, NUMBER 4 CAMBRIDGE, MASSACHUSETTS, U.S.A. 31 October 2002 (US ISSN 0027-4100) . | OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY PUBLICATIONS ISSUED | BreEvVIORA 1952— BULLETIN 1863— : Memoirs 1865-1938 ' Jounsonia, Department of Mollusks, 1941-1974 ) OCCASIONAL PAPERS ON MOLLusKs, 1945— SPECIAL 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, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4. Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Emest 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 of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the 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 of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. es of the New England Zoological Club 1899-1947. (Complete sets only.) Proceedings of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2002. ON THE SUBFAMILY XYLOPHAGAINAE (FAMILY PHOLADIDAE, BIVALVIA, MOLLUSCA) RUTH D. TURNER’ CONTENTS BicitonallNotes int © ean eee 223 NUNS trac ty. ea RAAT Ra Sea ars Betleel pee Rh er 223 TO LUG LOI ie een eaves Weare ee ae ete 224 Species Groups in the Genus Xylophaga ___. 225 \V GREET OS ypc is ei, a a ae 227 Preliminary Report on the Results of Experiments on the Ecology of Deep- Sea Wood Borers and the Role of Wood Hhay ove} IDYStS50) (SSE Lene eee ee 234 SOUT ey, ecb ee ee 236 Genus Xylophaga Turton 1822 _. 236 Xylophaga concava Knudsen _... 236 Xylophaga gerda Turner new species ____ 237 Xylophaga grevei Knudsen .......---- 238 Xylophaga clenchi Turner and Culliney 239 Xylophaga supplicata Taki and Habe _. 241 Xylophaga whoi Turner new species 242 Xylophaga profunda Turner new species — 243 Xylophaga abyssorum Dall 245 Xylophaga duplicata Knudsen __- 247 Xylophaga muraokai Turner new species — 247 Xylophaga atlantica Richards 249 Xylophaga washingtona Bartsch 250 Xylophaga rikuzenica Taki and Habe _... 252 Xylophaga depalmai Turner new species 253 Xylophaga guineensis Knudsen 256 Xylophaga mexicana Dall __..........----.---------- 257 Xylophaga tipperi Turner new species __. 259 Xylophaga bayeri Turner new species __.. 260 Xylophaga japonica Taki and Habe 261 Genus Xylopholas Turner 1972 _............-...-.- 262 Xylopholas altenai Turner 263 Genus Xyloredo Turner 1972 264 MPIDRBOID) TOGA VOTRE 265 Xyloredo ingolfia Turmer —_.....------------------ 266 Kyloredownaceli Tamer 2-22) 267 Achenexalercles tea ee ee ee 268 Witeratuner© ite cern een res arene ner mee 268 ‘Department of Mollusks, Museum of Compara- tive Zoology, Harvard University, Cambridge, Mas- sachusetts 02138. EDITORIAL NOTE. Professor Emerita Ruth Dixon Turner died on 30 April 2000 and was for the last several months of her life severely disabled; in fact, her active work as a researcher was considerably fore- shortened by medical problems beginning in about 1995. Among her Nachgelassene Werke was an im- portant manuscript on the systematics of the deep- sea pholadid bivalve genera Xylophaga, Xyloredo, and Xylopholas, a manuscript that she had been preparing for a number of years and one that had the active support of the U.S. Department of Defense’, then, Office of Naval Research (ONR). Professor Turner was unquestionably a leading world authority on these taxa and had posted this document, in its pre- liminary draft form, on a Web site; after her retire- ment and the beginning of the illnesses that plagued her, the manuscript was removed from the Web site with the intent of readying it for formal publication. Two outside authorities, Dr. Jorgen Knudsen of the Zool- ogisk Museum, Kgbenhavns Universitet, Kobenhavn, Denmark, and Dr. K. Elaine Hoagland, then at the Association for Systematic Collections, Washington, D.C., were solicited to make criticisms, and these, along with my own, were incorporated into a more advanced revision of the text prepared by Ms. Helene Ferranti, a long time coworker and associate of Pro- fessor Turner. Ms. Ferranti agreed to revise this Nachlass in accordance with the comments of the re- viewers and to update its content and organization. Having collaborated with Professor Turner on the subject of deep-sea bivalves, Ms. Ferranti is credited herein as the person responsible for the final com- pletion and revision of this valuable text. The new species described, for which specimens are available for study in the Museum of Comparative Zoology (MCZ), and the taxonomic suggestions incorporated into the text are to be credited to Professor Turner. Kenneth J. Boss Editor ABSTRACT. The provisional grouping of the species of the bivalve genus Xylophaga suggested by Turner and Culliney is further elaborated, with 37 species assigned to six groups depending on characteristics of Bull. Mus. Comp. Zool., 157(4): 223-307, October, 2002 Dore) bo bo the mesoplax, siphons, muscle scars, and method of reproduction. Three cases of variation in species of Xylophaga are discussed: variation in response to dif- ferent substrates, variation that possibly is genetic, and variation in a normal growth series. Some obser- vations are made regarding the ecology of deep-sea wood borers based on experiments carried out with wood panels; these support the hypothesis that wood contributes to the growth and diversity of deep-sea organisms and that the Xylophagainae contribute sig- nificantly to the food chain by converting wood to a usable form. A section on systematics considers 19 species of Xylophaga, of which 7 are new, as well as the monospecific genus Xylopholas and three species of Xyloredo. Detailed descriptions are given of new species. INTRODUCTION The Xylophagainae, a subfamily of the Pholadidae, is composed of the genera Xy- lophaga Turton, 1822, Xyloredo Turner, 1972, and Xylopholas Turner, 1972. Species in the genus Xylophaga range in depth from just below low tide (X. dorsalis Tur- ton) to depths of 7,000 m (X. grevei Knud- son), whereas species in Xyloredo and Xy- lopholas are restricted to the deep sea (depths of 239 to more than 2,000 m). Species in Xyloredo range in depth from 1,737 to more than 2,000 m (X. nooi Turn- er, 1972) and species in Xylopholas range from 239 to 366 m (X. altenai Turner, 1972), with one lot dredged in 2,550 m off Port Victoria, S40 Tomé, Gulf of Guinea. However, the Sio Tomé specimens were boring in coconut shells and may not have been living at that depth. The Xylophagainae are marine, cosmo- politan, and range from moderate to abys- sal depths. All of the Xylophagainae, so far as known, are wood borers, and all are sublittoral. Only in high latitudes do they compete with shipworms (Teredinidae) in cold boreal waters. So far as known, the Xylophagainae do not occur intertidally, or in floating wood. Wood containing speci- mens of the Xylophagainae usually has been obtained by dredging. Occasionally, specimens may be obtained from water- logged wood that has been brought up and thrown ashore during a storm after being on the bottom for some time. 4 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Until recently, the Xylophagainae were considered to be deep-sea organisms of lit- tle or no economic importance. They were rare curiosities, of interest mostly in their role of recycling wood on the continental shelf and slope and in the abyss, largely beyond the depth range of the teredinids. The Xylophagainae often were referred to as shipworms and because of the ephem- eral, patchy distribution of wood in the deep sea were thought to have little im- pact on ecological processes. With the ex- tension of human activities into the deep sea for fishing (especially trap fishing for lobsters and crabs), as well as for archae- ology, mining, monitoring currents, and other activities, these borers are now gen- erally considered to be pests. Species in the genus Xylophaga (Xylo = wood, phaga = eating) are restricted to wood, woody plants, and structures made of wood found in the deep sea. In com- mon with the teredinids (shipworms), they have symbiotic bacteria in their gills (Wa- terbury et al., 1983). These bacteria are be- lieved to have the ability to digest cellulose and probably to fix nitrogen. Collaborative work with Dr. Waterbury, microbiologist at Woods Hole Oceanographic Institute, was unable to culture cellulose-digesting bac-— teria from the gills of X. atlantica, but ev- idence was found of cellulose enzymes in the gill tissue. : The three genera of Xylophagainae may be briefly characterized as follows: Genus Xylophaga Turton. Siphons rela- | tively short, of equal length or with the excurrent siphon truncated, and often | capable of retraction between the valves. Burrow seldom more than five times the length of the valves and often with a chimney of fecal pellets lining the posterior end of the burrow. Genus Xylopholas Turner. Shell typical but with the animal extended and with lateral plates on the siphons. Genus Xyloredo Turner. Shell typical but animal elongated and producing a tere- | dinidlike burrow that is lined with a cal-_ careous tube marked with distinct growth rings and margined anteriorly with a periostracal band. The Xylophagainae are often confused with teredinids, but the gills and digestive and reproductive organs in the Xylopha- gainae do not extend posteriorly beyond the valves. In addition, the Xylophagainae do not have pallets to close the entrance to their burrows or apophyses for the at- tachment of the foot muscles. In common with the teredinids, but unlike species in the pholadid genera Martesia and Ligno- pholas, the only other genera of wood-bor- ing pholadids, the Xylophagainae have a large wood-storing cecum and probably utilize the wood in which they bore as food. For details of anatomy, see Purchon (1941) for Xylophaga dorsalis Turton, and Turner (1955) for X. atlantica. SPECIES GROUPS IN THE GENUS XYLOPHAGA Genus Xylophaga Turton. Xylophaga Turton, 1822, Conchylia Insularum Britanicarum, p. 253 (type species, Teredo dorsalis Turton, 1819). Species in this genus are characterized by teredolike shells that lack apophyses and have a divided mesoplax that is vari- able in shape and size. A chrondrophore and internal ligament are present. The si- phons are variable, united for part or all of their length, with the excurrent siphon of- ten truncated. The visceral mass and gills do not extend beyond the valves posteri- orly. The wood-storing cecum is large. In his discussion of the taxonomy of Xy- lophaga, Knudsen (1961) believed that the use of subgenera was not feasible and would only lead to the creation of a large number of monotypic subgenera that would be of limited value. This is still par- tially true but new species described in this report and the additional material now available concerning other species have made possible a provisional grouping of the species, as suggested by Turner and Culliney (1971). See also Hoagland and XYLOPHAGAINAE ° Turner 925 Turner (1981) and Hoagland (1983). This grouping is helpful when discussing rela- tionships and geographic distribution. The characters used for grouping the species are those mainly of the mesoplax and si- phons in conjunction with the muscle scars and methods of reproduction (see Text- Fig. 1). The mesoplax is a transverse plate, usually wider than long, that straddles the valves at the umbos and partially or com- pletely covers the posterior end of the an- terior adductor muscle. The mesoplax may be composed of one or two parts. The im- portant character of the mesoplax is the presence or absence of a ventral portion and tubes: the more detailed characters, such as the presence of lobes, seem to be of specific value only. Siphonal characters include the relative length of the two si- phons, the presence or absence of cirri at the apertures, and the type of siphonal folds, which may or may not have lappets or fringes. Not all characters are known for all spe- cies and a few species seem to be transi- tional between groups. There is no ques- tion that more material is needed before definite statements can be made concern- ing the formal use of subgenera. However, the grouping of species as presented here does offer an opportunity to speculate on the possible origin and evolution of the ge- nus and to focus attention on the types of information that should be considered in future studies. Comparative anatomical and molecular studies are greatly needed but it probably will be some time before these can be completed because well-pre- served specimens of deep-sea Xylophaga are rare and difficult to obtain. If we consider species with simple si- phons of equal length and a mesoplax of two simple flat to slightly curved plates to be the basic type, it is possible to group the species in what appears to be a devel- opmental series of six groups. This list does not include all nominal Xylophaga. The groups may be characterized as fol- lows: bo Text-Figure 1. siphonal retractor pedal retractor posterior adductor ventral adductor umbonal-ventral ridge Bulletin Museum of Comparative Zoology, Vol. 157, umbonal reflection extended. Group 1. Group 2. Mesoplax composed of two simple flat or slightly curved plates lo- cated posterior to the anterior adductor muscle and standing erect. Siphons of equal length or with the excurrent si- phon slightly shorter, cirri on one or both siphons present or absent. Group 1 includes X. erecta Knudsen, X. con- cava Knudsen, and X. gerda Turner n. sp. Mesoplax composed of two plates that may be curved, flat, sculp- tured, or smooth, set at an acute angle to each other, lack dorsal tubes and a ventral portion, but cover the anterior adductor muscle dorsally. Siphons of the same length or with the excurrent si- phon slightly shorter and with large cirri on the sides of the excurrent siphon and small cirri at the incurrent siphon. Group 2 includes X. grevei Knudsen, X. wolfft Knudsen, X. hadalis Knudsen, X. galatheae Knudsen, X. murrayi Knud- sen, X. africana Knudsen, X. panamen- sis Knudsen, and X. clenchi Turner and Culliney. Group 3. Group 4. No. 4 anterior adductor incurrent siphon umbonal- ventral sulcus mantle Nomenclature of parts of Xylophaga. (1) Internal view of left valve showing relative position of muscle scars. (2) External view of left valve. (3) Dorsal view of animal with siphons retracted. (4) Lateral view of entire animal with siphons nearly flat plates set at an acute angle to each other forming an inverted V, with tubes extending from the posterior dor- sal surface or longitudinally folded with anterior lobes or pores. Mesoplax lack- ing a ventral portion and set in a tentlike | fashion over the anterior adductor mus-_ cle. Siphons nearly the same length and usually with small cirri on both open-— ings. Group 3 includes X. supplicata | Taki and Habe, X. lobata Knudsen, X. tubulata Knudsen, X. bruuni Knudsen, | X. obtusata Knudsen, X. whoi Turner n. | sp., and X. profunda Turner n. sp. | Mesoplax composed of two plates that have a small to large ventral | portion, the dorsal portion being smooth, folded, or lobed. Siphons of the - same length or with the excurrent si- phon slightly shorter and with cirri or papillae at one or both openings. Group_ 4 includes X. abyssorum Dall, X. dupli- cata Knudsen, X. muraokai Turner n. sp., X. foliata Knudsen, and X. atlantica’ Richards. Mesoplax composed of two. } Group 5. Mesoplax composed of two plates that are more or less triangular in outline, with a ventral portion ranging from very narrow to more than one half the width of the dorsal portion. The ex- current siphon may vary in length from one half to three quarters that of the incurrent siphon and have cirri, or it may be truncated just posterior to the valves and have dorsal lobes or folds ex- tending from the truncation along the dorsal surface of the incurrent siphon for part or all of its length. Group 5 in- cludes X. washingtona Bartsch, X. ri- kuzenica Taki and Habe, X. aurita Knudsen, X. turnerae Knudsen, and X. praestans E. A. Smith. Group 6. Mesoplax composed of two more or less ear-shaped plates some- what coiled posteriorly. Excurrent si- phon truncated near the posterior end of the valves, continuing as lateral lobes extending from the truncation along the dorsal surface of the incurrent siphon. These lobes may vary in width but are always fringed. Group 6 includes X. dor- salis Turton, X. depalmai Turner n. sp., X. guineensis Knudsen, X. mexicana Dall, X. tipperi Turner n. sp., X. bayeri Turner n. sp., X. globosa Sowerby, X. ja- ponica Taki and Habe, and X. indica Smith. Most species are known only from the type series and these have all been studied by the author except X. indica, and the spe- cies described by Taki and Habe. Howev- er, paratype specimens received through the kindness of Dr. Habe are in the col- lection of the MCZ. They include X. ja- ponica, X. rikuzenica, X. teramachi, and X. supplicata, although unfortunately all lack the mesoplax except the last. Two species, X. teramachi Taki and Habe (Taki and Habe, 1950) and X. tomlini Prashad (Pras- had, 1932); are known only from the valves and remain unassigned. A map showing the distribution of species of Xylophaga is provided in Text-Figure 2. Nineteen of the 37 species of Xylophaga XYLOPHAGAINAE * Turner ORT listed in the groups above as well as the monospecific genus Xylopholas and the three species of Xyloredo are considered in the section on systematics. Some spe- cies are discussed more fully than others but the distinctive characters have been given for all. For example, Xylophaga mex- icana Dall and Xylophaga abyssorum Dall are fully described because these names were based on valves only and were vir- tually nomina dubia. By matching the valves of the holotypes with complete specimens, it has been possible to fix the names of these species. If additional char- acters or records are given for a well-de- scribed species, a reference is made to the original description. Detailed descriptions are given for new species. VARIATION Knudsen (1961) aptly stated that very little was known about variation in species of Xylophaga. Unfortunately, large series of any one species seldom have been avail- able for study because material usually is obtained from small pieces of wood or oth- er plant material that has been dredged or occasionally thrown ashore as driftwood. Of the 30 species listed by Knudsen and the 7 species described as new in this pa- per, 24 are known from fewer than 10 specimens; only 7 species are known from series of more than 100 specimens. Most are known from only one or two localities and often all specimens are from a single piece of wood, and may all be of the same set, that is, have settled at the same time. Consequently, it is not surprising that all specimens in any one lot are quite similar. Only since the beginning of deep-sea test- ing and the use of the submersible DSV Alvin to place experimental wood islands at great depths has it been possible to ob- tain sufficient material to study intraspe- cific variation in this subfamily. The earliest work of this sort was done by the U.S. Naval Civil Engineering Lab- oratory (USNCEL) and the Navy Ocean- ographic Office (NOO). Three cases of variation based on this material are re- iN) bo 8 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 dorsalis a praestans 0 A 0 grevei bruuni indica Q tomlini or o erecta Rm: rikuzenica supplicata japonica tubulata clenchi atlantica ° abyssorum depalmai washingtona S pir aoka proto . Sue obtusata duplicata . guineensis africana Oe wolffi ct ~obata —___ murrayi ae o' wliata concava aurita turnerae AS galathea uy 27 hadalis globosa Text-Figure 2. Distribution map of species of Xylophaga. ported here. The first case involves varia- tion in response to different substrata, the second case exhibits variation that possibly is genetic because the substrate and all other parameters were as nearly uniform as possible, and the third case illustrates variation in a normal growth series. Variation Due to Different Substrates. Variation in Xylophaga washingtona in re- sponse to the substrate can be denounce at= ed with material from the USNCEL tests. A series of 10 panels of different types of wood were attached to a submersible test unit (STU) that was submerged from April 1965 to May 1966 off San Miguel Island, California (34°06'N, 120°42’W) at 2,370 ft (730 m) (see Table 1 and Text-Figs. 3-9). mhes2)xe6r< 0) d-inl (0S P52 alo a7— mm) wood panels were all attached to the same rack on the STU so that they would be resting just above the mudline. Con- sequently, all factors affecting the borers were as nearly identical as possible except’ the substrate (i.e., the species of wood) on which the borer larvae settled and into” which they would bore. Text-Figures 3-8 | illustrate typical specimens from each of. the wood panels; Text-Figure 9 shows specimens from a phenolic laminated rod. Ihe 1g interesting to note that the dorsal plates in all specimens are remarkably uni- | form, varying only slightly in length/width | proportions. Even the specimens taken from the phenolic laminated rod could be | identified by the dorsal plates. The general shape of the valves with the high posterior slope also remained rather constant except in the extremely steno- | morphic (stunted) specimens from Afam- beau and the phenolic laminated rod. It is difficult to explain the proportionate size of the larval valves on specimens boring into harder materials except that these specimens had not greatly increased in di-_ XYLOPHAGAINAE * Turner 229 TABLE 1. VARIATION OF XYLOPHAGA WASHINGTONA BURROWS IN DIFFERENT TYPES OF WOOD. No. Burrow Burrow Specimens Length Diameter Wood eanined (m m) (mm) Remarks Cedar 50 25.0 4.5 many dead, often three or four specimens in one enlarged cavity where burrows ran together. Heavily attacked, particularly at one end. Ash 125 21.0 5.0 many dead, burrows running together, panel heavily attacked, particularly at one end. Maple 75 11.0 5.5 well distributed, with a little more concentration around the hole at one end. Pine 35 15.0 5.0 clustered at one end, newly settled to adult. Oak +50 15.0 6.0 evenly distributed, shells yellowish-green from wood. Fir 46 12.0 4.5 many newly settled, often cut into burrows of oth- er specimens. Redwood 46 14.0 4.5 specimens stained dark red brown by wood. Greenheart 75 2.75 1.75 concentrated around edges. Afambeau 19 0.75 0.05 concentrated at one end, all very small, many in umbo stage veliger, 15 small depressions with- out animals. Antidesma pulvinatum 150 0.10 0.05 many newly settled, just beginning metamorpho- sis. = [+__—____| 1mm 1mm Text-Figure 3. Typical specimens of Xylophaga washingtona collected from submerged cedar (top) and pine (bottom). 230 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Ash Maple Oak 5mm Fir Text-Figure 5. Typical specimens of Xylophaga washingtona collected from submerged oak (top) and fir (bottom). XYLOPHAGAINAE *° Turner 231 Redwood Greenheart Afambeau ba Saar 0.1 mm Text-Figure 7. Typical specimens of Xylophaga washingtona collected from a submerged panel of Afambeau. 232 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 eae Sa Sy 0.1mm Text-Figure 8. Typical specimens of Xylophaga washingtona collected from a submerged panel of Antidesma pulvinatum. ameter as they bored so that the larval valves were not inturned with the growth of the umbos. Evidently little or no attri- tion had occurred at the umbonal area or the embryonic valves would have been worn away. The most variable characters in this se- ries are the size of the valves and the num- ber of denticulated ridges on the anterior slope in relation to the length of the valves. Specimens from cedar, pine, ash, maple, oak, and fir (Text-Figs. 3-5) all were drawn to the same scale, as shown by the 5-mm scale bar, and are arranged in order of de- creasing size. The number of denticulated ridges on the anterior slope of these spec- imens varies from 12 to 20. Specimens from redwood, greenheart, Afambeau, and Antidesma pulvinatum (Text-Figs. 6-8) were drawn to their own scales and, al- though these specimens are much smaller, they have as many or more ridges on the anterior slope. The specimen removed from the phenolic laminated rod (Text-Fig. 9) has 44 denticulated ridges. Correlated with the increased hardness of the sub- strate and the additional denticulated ridg- es is a proportional increase in the size of the posterior adductor muscle and its scar. This suggests the enlargement of the pos- terior adductor muscle in response to in- creased activity of boring. The general shape and sculpturing of the muscle scar was similar in all specimens. No noticeable variation was found in the siphons, except size, regardless of the substrate. However, specimens that were able to bore deeply into the wood usually formed a chimney composed of compacted fecal material lin- ing the posterior end of the burrow. Two periods of settlement apparently occurred on some of the panels, because specimens of two age groups could be found. However, it is impossible to say whether the specimens removed from the Antidesma were of the second set or if NL Boe Phenolic Laminated Rod XYLOPHAGAINAE ° Turner 233 Text-Figure 9. Typical specimens of Xylophaga washingtona collected from a submerged phenolic rod. they had simply been unable to increase in size because of the hardness, chemical composition, or both of the wood. Because the specimens apparently were alive at the time the wood was removed from the wa- ter and because only a few rows of dentic- ulated ridges were present, the inference was made that these specimens probably belonged to a second set. Certainly the lar- val shells shown in Text-Figure 8 must be from a second set. The number of speci- mens examined and the maximum length and diameter of the burrows for each type of wood are given in Table 1. Variation in the Mesoplax of Xylophaga depalmai Turner n. sp. Approximately 300 specimens of X. depalmai n. sp. were ob- tained from tests conducted by the NOO about 2—3 miles east of Fort Lauderdale, Florida (26°04'N, 80°04'W) in depths from 100 to 500 ft (30.5 to 152.5 m) (see Table 14 under the description of X. de- palmai n. sp. for information giving panel numbers, depth, and dates of exposure). In this species, the general shape of the valves, the siphons, and the muscle scars show little variation but the mesoplax is extremely variable. The mesoplax is typi- cally bilaterally symmetrical, ear-shaped, longer than wide, with the two halves coiled inward at the posterior end, and with a long medial line where the two halves meet (Plate 24, Figs. 16, 17). In nu- merous specimens, the mesoplax was not bilaterally symmetrical, but one half was considerably shorter than the other and of- ten appeared malformed (Plate 24, Figs. 11-15). In several specimens, the ventral surface of the two halves of the mesoplax was fused by the periostracal covering, al- though the dorsal surface still appeared di- vided. In other specimens, the mesoplax was elongated, the coiled posterior ends, instead of curling inward toward each oth- er, remained nearly straight or curled slightly outward, with the ventral surface being completely fused (Plate 24, Figs. 1— 5). In two specimens, the two halves of the mesoplax had completely fused dorsally and ventrally, although the lines of fusion remained clearly visible. The tapered pos- terior end of this cornucopialike mesoplax coiled slightly ventral and to the left. Such variation is in marked contrast to the uniformity seen in the mesoplax of X. washingtona Bartsch. It is impossible to say whether this variation is genetic or eco- logic but we are able to say that all types of the mesoplax of X. depalmai were found in a single panel retrieved from a depth of approximately 300 ft (91.44 m). Variation in the mesoplax is a factor that must be 234 taken into consideration when evaluating species in this genus. Variation Exhibited in Growth Series. In most species of Xylophaga, the dorsal plate is quite simple and the mesoplax of the young is similar to that of the adult, as seen for X. washingtona. Dons (1929a,b) described briefly and illustrated a similar situation in X. praestans Smith and X. dor- salis Turton. Some species have more elaborate dor- sal plates and for some of these (i.e., X. muraokai Turner n. sp., X. bayeri Turner n. sp., and X. abyssorum Dall), it is pos- sible to build up what appears to be growth series. All begin with a simple pos- terior covering to the anterior adductor muscle that is difficult to differentiate among the species in the young stage. As calcification of the periostrascal covering progresses, the adult form gradually ap- pears. In the three species mentioned, the characters of the valves and the siphons of the young agree with those of the adult specimens. Therefore, although living ma- terial has not been available and develop- mental studies have not been possible, it seems reasonable to consider these as growth series. Xylophaga muraokai (Plate 19, Fig. 3) is relatively simple, with the dorsal “portion becoming thickened and joining laterally with the basal portion while the ventral flanges enlarge. In X. bayeri (Plate 31, Fig. 3), the broad trans- verse dorsal portion becomes very con- spicuous and in X. abyssorum (Plate 15, Figs. 4, 5), the dorsal incurving of the ee eral arms produce an elaborately lobed mesoplax. Nothing is known of the allo- metric changes ihe take place during growth of ine peculiar dorsal plates of xX tubulata Knudsen, X. bruuni Knudsen, X. obtusata Knudsen, and X. whoi Turner n. sp. Detailed studies of these growth series will have to await improved techniques for handling living material under deep-sea conditions, and the ability to rear deep-sea species in the laboratory. In the meantime, it seems best to consider these forms as Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 members of growth series rather than dif-_ ferent species, particularly because the se-_ ries in each case was built up from mate- i rial taken from a single piece of wood. | PRELIMINARY REPORT ON THE RESULTS OF EXPERIMENTS ON THE ECOLOGY OF DEEP-SEA WOOD BORERS AND THE ROLE OF WOOD IN | THE DEEP SEA | The results of the first exposures of wood panels at the Woods Hole Oceano- graphic Institution (WHOI)—Alvin per- | manent bottom station south of Woods | Hole (39°46'N, 70°41'W, in 1,830 m) were | reported in 1973 (Turner, 1973). At that time, the Xylophagainae were postulated — to be the most important deep-sea organ-_ isms involved in converting woody plant material to an available food source for | other organisms. Pursuing this theory and to continue the studies of the ecology and life history of the Xylophagainae, wood panels were ex- posed at the Alvin permanent station in the Tongue of the Ocean, Bahama Islands, on 19-22 January 1974 (Alvin dives 492, 493, 494, and 495) at a depth of 2,032 m. The first of these panels was picked up on 7 March 1974, frozen immediately, and re- turned to Woods Hole where it was ex- amined. Newly settled larvae and meta- morphosing X ylophaga with one to two rows of denticulated ridges were removed from the panel. The specimens were ap- proximately 300 ym in length and the greatest penetration was about twice the depth of the shell. The debris rings sur- rounding the burrows were much coarser than tose made by teredinids. The distri- bution of the entrance holes was some- what patchy and varied from 5 to 20 cm’. The specimens were too young to identify because none of the dorsal plates had been formed but examination with scanning electron microscopy showed a well-devel- oped distinctive sculpture on the larval shell. This first panel from the Tongue of the Ocean established that Xylophaga were t just beginning to settle on the wood a max- imum of 48 days after it was implanted in ‘the bottom and that settlement of larvae could occur in early March, at least at this site. Three more panels were removed from the Tongue of the Ocean station (Tower 1—west arm) on 19 April 1975 during Al- vin dive 552. I was an observer on this dive and as we approached the panels I noticed an increase in the number of shrimp and galatheid crabs. The panels had numerous crabs crawling all over them. Some of the crabs had crawled under the plastic mesh bags covering the panels and had grown so lar: ge they could not escape. (Note: After the. near loss of the panels at the northern station because of the heavy attack of bor- ers, the decision was made to put the pan- els in plastic mesh bags so that the pieces could be retrieved if they began to disin- tegrate.) The specimens inside the bag were carried to the surface with the pan- els. When the panel was disturbed, the specimens on the outside of the mesh fell off. The largest of the 12 crabs was 43 mm in length. The diamond-shaped opening of the mesh was 5 X 10 mm. The smallest crab measured § ae in length; others measured 40, 33, 32, 30, and 24 mm. It is obvious that the se were finding suffi- cient food either in or on the wood B grow at a fairly rapid rate. The first young crabs to find the wood may have fed on the newly settled Xylo- phaga larvae and this might explain the patchy distribution of tne borers in the panels. However, the larger crabs would not have stayed on the wood unless there was something for them to eat. The crabs had to be under 10 mm in length to get under the mesh and if the larvae were not settling until early March it would be at least early May before the borers had grown sufficiently to be a good food source for the crabs. Therefore, I think we can postulate that the largest crab measured grew at least 33 mm in a period of 10 months. Examination of the panels showed XYLOPHAGAINAE ° Turner 935 rather heavy attack of three species of Xy- lophagainae. These included Xyloredo nooi Turner and two Xylophaga species, Xylophaga clenchi Turner and Culliney and X. profunda Turner n. sp. The X. nooi were typical with valves that reached 5 mm in length and burrows that were 18— 22 mm in length. The calcareous lining of the burrow of the largest specimen was 13 mm long and 2.5 mm in diameter at its anterior end. The smaller species of Xy- lophaga, X. clenchi Turner and Culliney, also had been obtained previously from wood exposed in the Tongue of the Ocean by John DePalma of the: NOO. This is a fairly small species. The valves were 8—10 mm in length and several of the specimens were carrying larvae on the umbonal area of the valves. The larvae measured 0.2 mm in length. The large species of Xylophaga, X. profunda Turner n. sp., had not been seen before. The valves were 14 mm in length, and one specimen measured 40 mim to the tip of the siphons. The burrows were 45-50 mm in length. Both X. clenchi and X. profunda lined the posterior end of their burrows with consolidated fecal pel- lets and the burrows of all dead specimens contained one or more specimens of cap- itellid worms that were feeding on the pel- lets as well as the remains of the Xylopha- ga. Often the spaces between the valves of che borers were filled with the smaller fe- cal pellets of the worms. Breaking these balls of pellets apart, I always found one or two capitellid worms. In the X ylophaga burrows and on the surface of the wood, I also found two other polychaete worms. One belonged to the family Chrysopetali- dae and the other to a family of polynoid worms. A preliminary examination of the stom- ach contents of a broken specimen of a galatheid showed that the crab had ingest- ed some fine chips of wood because iden- tifiable cells remained in the material. Consequently, we can postulate that the crabs were feeding on the Xylophaga, probably dead ones. “The tissues of the Xy- lophaga are so soft that they are unrec- 236 ognizable in such a preliminary examina- tion. The crab’s stomach also contained se- tae of the chrysopetalid worms, a small nematode, and some sponge spicules. The chrysopetalids probably were feeding on the capitellid worms. I think five points are worthy of notice: 1) Xylophaga in the Tongue of the Ocean as well as at the northern Alvin station are growing much faster than one would ex- pect. 2) The larvae of X. profunda n. sp. were settling in early March. 3) Xylophaga clenchi broods its young and was carrying young in mid-April. 4) Probably the crabs and worms were also growing faster than normal for the deep sea. 5) A food chain based on wood and Xylophaga was being developed. This lends support to my hypothesis that wood is an important source of enrich- ment in the deep sea, that it contributes to both the diversity of organisms in a giv- en area and to their rate of growth, and that the Xylophagainae are the most im- portant organisms involved in converting the wood to a usable form. To my lenovell edge, this isthe first documented food chain for invertebrates in the deep sea. On the basis of these simple experiments, it now seems conceivable that the slow growth rates usually attributed to deep-sea animals may be due to lack of food, at least for epifaunal forms, rather than being an inherent characteristic of the species in- volved. SYSTEMATIC ACCOUNT GENUS XYLOPHAGA TURTON 1822 Xylophaga concava Knudsen Plate 1 Xylophaga concava Knudsen, 1961, Galathea Report, 5: 167-169, figs. 4, 5 (Galathea, station 726, Gulf of Panama [5°49'N, 78°52’W] in 3,270-3,672 m). Holotype, Zoological Museum, University of Co- penhagen; paratype, MCZ 235796. Distinctive Characters. Posterior slope of valves concave when viewed dorsally. Mesoplax composed of two rather wide, erect, curved plates that extend above the Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 umbos. Siphons nearly the same length, excurrent slightly shorter with a few large | cirri, incurrent siphon with many small cir ri (Plate 1, Figs. 2, 3). Chimney apparently | lacking, not mentioned by Knudsen and. not found with the single specimen re- ported here. | Remarks. At the time Knudsen de-- scribed this species he had 4 Specimen: from Galathea station 726 and 25 speci-_ mens from Galathea station 739. Both of | | these stations were given as in Gulf of Pan- ama. However, station 726 is about 95. miles west of the Gulf of Tibuga, Colom- bia, whereas station 739 is about 90 miles. west of Ensenada Guayabo, Panama. A single specimen of X. concava was taken’ by the R/V Pillsbury at station 526. This locality is about midway between the two Galathea stations. | The Pillsbury specimen agrees closely with the description and figures given by Knudsen (1961) except that the incurrent. siphon has a double row of about 25 small cirri around the aperture and the excur- rent siphon has 6 large cirri (Plate 1, figs. 2, 3). Concerning the siphons, Knudsen’ stated that “both openings are at the distal end, close together, and around them 15— 16 small cirri are present.” In Knudsen’s- illustration, the distal ends of the siphons _ appear to be contracted; neither the two openings nor the cirri are apparent. The contracted condition of Knudsen’s speci-| mens probably accounts for the differenc-| es noted here. Xylophaga concava is closely related to| Xylophaga g gerda Turner n. sp. but differs in the size Ok the mesoplax, the type of | siphonal openings, and the chimney. (See Remarks under x. gerda.) Xylophaga con-. vava also is related to X. erecta Knudsen: (1961) from the Sulu Sea. Knudsen (1961) | reported that no cirri were visible on the. siphons of X. erecta, that the posterior ad- ductor scar was much broader in X. con-_ cava, and that the posterior slope of the valve viewed dorsally was convex in X. er ecta rather than concave. Knudsen’s de-’ scription of X. erecta was based on 20) specimens and unfortunately no further records have been obtained. Range. From off Ensenada Guayabo, Pannen south to off the Gulf of Tibuga, Colombia, in depths from about 915 “he 3,670 m. Specimens Examined. COLOMBIA: Galathea, sta- tion 739, Gulf of Panama, about 90 miles W of En- senada Guayabo (7°22'N, 79°32’W) in 915-975 m (dried specimens); Pillsbury, station 526, about 110 miles W of Cabo Marzo (6°53’'N, 79°27'W) in 3,193— 3,211 m: Galathea, station 726, Gulf of ee (5°49'N, 78°52’W) in 3,270-3,670 m. Xylophaga gerda? Turner new species Plates 2, 3 Holotype. MCZ 328378. Paratypes, MCZ 316741, 316742. Type Locality. Gerda, station 499, about 3 miles off Southwest Point, Great Baha- ma Island, Bahama Islands (26°37'N, 78°56’W) in 155 fathoms (283.96 m). Distinctive Characters. Posterior slope of valves concave when viewed dorsally. Mesoplax composed of two narrow, erect curved plates at the posterior end of the anterior adductor muscle. Siphons of equal length, with a periostracal sheath and four or five cirri surrounding the ap- ertures. Chimney composed of fecal ma- terial agglutinized to a periostracal base Welate 2, Fig. 7). Description. Shell globose, fragile, reaching 3.0 mm in length and 2.8 mm in height, alae inflated. Pedal angle 110— 115°. Anterior slope with up to 45 rather evenly spaced denticulated ridges. Umbo- nal_ventral sulcus narrow and only slightly depressed. Disc and posterior slope sculp- tured with fine, incised growth lines. Pos- terior slope high, flaring, and somewhat ear-shaped. Inner surface of valves smooth and glis- tening. Umbonal—ventral ridge low and in- distinct except near the wide, low ventral condyle. Chondrophore and internal liga- 2 Named for R/V Gerda, Rosenstiel School of Ma- rine and Atmospheric Sciences, University of Miami, Miami, Florida, whose station 499 is the type locality. XYLOPHAGAINAE * Turner 237 TABLE 2. MEASUREMENTS OF XYLOPHAGA GERDA. Length Height (mm) (mm) Location Tail 1.0 Gerda, station 266 15 1.4 Gerda, station 266 1.8 2.0 Gerda, station 266 2.5 2.3 paratype 3.0 2.5 holotype 3.0 2.8 Pillsbury, station 328 3.3 3.0 Pillsbury, station 944 3.8 3 Pillsbury, station 944 ment small. Posterior adductor muscle scar large with faint transverse impres- sions, which are best seen externally on an entire specimen. Disc separated from pos- terior slope by a shelflike ridge (Plate 3, figs. 2-5). Pedal and siphonal retractor scars not visible. Mesoplax composed of two erect, nar- row, curved, slightly calcified plates, locat- ed just posterior to the anterior adductor muscle and not extending above the um- bos. Siphons long, probably not capable of retraction between the valves, of equal length, united nearly to the tip, with a thin periostracal sheath. Siphonal apertures of about equal size, each with four or five comparatively large cirri, which appear as a common ring of cirri when the siphons are retracted. A brown periostracal cylin- der containing fecal material may extend nearly one hale the length of the excurrent siphon (Plate 2, Figs. 124) Chimney built in sections, composed of fine fecal material agelutinized on a periostracal lining with “leaves” of periostracum extending to the outer surface (Plate 2, Figs. 6, 7). Arrange- ment of the gills and labial palps typical for the genus, foot large but not muscular, cecum very large and showing through the foot (Plate 2, Fig. 5). Pedal and siphonal retractor muscles weak, their arrangement typical for the genus. Measurements. See Table 2. Remarks. On the basis of the shell and the mesoplax, this species is closely related to X. concava Knudsen from the Gulf of Panama. It differs in being much smaller 238 (none of the 19 specimens reached 4 mm in length, whereas Knudsen gives 8.6 mm for X. concava) and in having a narrow me- soplax that does not extend above the um- bos. In addition, the excurrent and incur- rent siphons of X. gerda are the same length, are separate at the tip, and the ap- ertures of both siphons have four or five relatively large cirri. In X. concava, the si- phons are joined for their entire length and the excurrent siphon is slightly shorter. The type of chimney produced by X. gerda is unlike any other known to date in this genus. Unfortunately, all the specimens of X. gerda are small, extremely fragile, and rather poorly preserved. Consequently, it has been impossible to do anatomical work beyond that mentioned in the description. One specimen from which the valves were removed appeared to have an accessory genital organ similar to that described by Purchon (1941). Range. Probably throughout the Carib- bean in depths from about 283 to 2,072 m. Specimens Examined. BAHAMA ISLANDS: Ger- da, station 499, about 3 miles off Southwest Point, Great Bahama Island (26°37'N, 78°56’W) in 155 fath- oms (283 m). UNITED STATES, FLORIDA: Gerda, station 266, off Fowey Rocks, Florida Keys (25°39'N, 79°58’W) in 185-187 fathoms (338-342 m). LESSER ANTILLES: Pillsbury, station 944, 45 miles N of Port Louis, Guadeloupe Island (16°32.2'N, 61°36.8'W) in 364-421 m. PANAMA: Pillsbury, sta- tion 328, about 25 miles N of Punta San Blas, Gulf of San Blas (9°55.8’N, 78°59.8’W) in 2,069-—2,072 m. Xylophaga grevei Knudsen Plate 4 Xylophaga grevei Knudsen, 1961, Galathea Report, 5: 176, figs. 16-18 (Galathea, station 495, Banda Trench, south of Ceram [5°26'S, 130°58’E] in 7,250-7,290 m). Holotype, Zoological Museum, University of Copenhagen. Distinctive Characters. Mesoplax com- posed of two triangular plates that are flat dorsally, in contact the length of their me- dian edge, bent downward on their outer edge to meet the umbonal reflection, and lack a ventral portion. Posterior adductor muscle scar with oblique radiating impres- Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 TABLE 3. MEASUREMENTS OF XYLOPHAGA GREVEI. | Length Height (mm) (mm) Location 1.9 1.6 Galathea, station 444 12.5 11.0 holotype sions. Siphons nearly the same length, the | aperture of the excurrent siphon much_— smaller in diameter than the incurrent si- phon and with about 6 cirri; incurrent si- | phon with about 35 small cirri. Young car- | ried on the umbonal area of the adult. | Measurements. See Table 3. | Remarks. Through the kindness of Jor-_ gen Knudsen, it was possible to borrow the preserved dredged wood from the Zoolog- | ical Museum, University of Copenhagen. In a small piece taken by the Galathea at station 444, I found several additional specimens of X. grevei. They are much) smaller than that figured by Knudsen, the anterior slope is much narrower, and the > denticulated ridges are more widely spaced. However, they appear to be young but sexually mature specimens of that spe-- cies. From one to five rather large young: were attached posterior to the umbos on | the dorsal surface of the parent shells. Xylophaga wolffi Knudsen, based on) only two specimens, also was from Gala- thea, station 444. It has valves, muscle | sears, and siphons similar to those of X. | grevei. The outstanding difference be-_ tween these species is the flat plates of the mesoplax of X. wolffi, which are set in| tentlike fashion at an acute angle to each_ other. It has not been possible to compare the type of X. wolffi with this new material | | | from Galathea, station 444, but it appears _ that these two forms (i.e., X. grevei and X._ wolffi) may be equivalent to the condition found in X. clenchi Turner and Culliney, where occasional specimens have a bent, flat-topped mesoplax (Plate 4, Figs. 5, 6). Further collecting may show X. grevei and _ X. wolffi to be forms of a single species. The denticles on the ventral edge of the | mesoplax of X. wolffi described by Knud- sen may be an age factor but more mate- rial is needed to prove this. Range. Mindanao Sea south to the Ban- da Trench, Banda Sea in depths from about 545 to 7,290 m. Specimens Examined. PHILIPPINE ISLANDS: Galathea, station 444, Sulu Sea, W of Basilan Island (7°54'N, 121°30’E) in 5,050 m. Xylophaga clenchi? Turner and Culliney Plates 5-8 Xylophaga clenchi Turner and Culliney, 1971, Amer- ican Malacological Union Annual Report for 1970, p. 66 (U.S. NOO test site, Tongue of the Ocean, about 4 miles off northeastern tip of Andros Island, Bahama Islands [24°54’N, 77°49'W] in 1,737 m). Holotype, MCZ 316743; paratypes, MCZ 316744, 316745. Distinctive Characters. Mesoplax small, composed of triangular, nearly flat plates lacking a basal portion; the two plates usu- ally meeting at an acute angle in frontal view. Burrow lined with a chimney of coarse, loosely consolidated fecal pellets. Young held on posterior dorsal surface of the adult. Excurrent siphon shorter than incurrent siphon and with two large papil- lae on either side. Description. Shell globose, reaching 14 mm in length and 13.5 mm in height, thin, fragile, with a thin, light brown periostrac- um that is thickened along the dorsal and posterior margin of the valves. Umbos prominent and strongly incurved. Pedal gape angle about 108°. Beaked portion of the anterior slope sculptured with numer- ous denticulated ridges that are widely spaced in the young, becoming increasing- ly compacted toward the ventral margin in older specimens. Specimens 5 mm _ in length have up to 25 ridges. Posterior por- tion of the anterior slope rather narrow. Umbonal-ventral sulcus moderately to deeply impressed becoming shallower with age and bounded posteriorly by a broad, low, rounded ridge. Disc and pos- *>Named for Dr. William J. Clench, Curator of Mollusks, 1926-1966, Museum of Comparative Zo- ology, Harvard University. XYLOPHAGAINAE ° Turner 239 terior slope sculptured with uniform, rath- er pronounced growth ridges that are par- ticularly prominent in young specimens. Inner surface of valves smooth and glis- tening. Umbonal-ventral ridge prominent, distinctly segmented, and with a large ven- tral condyle. Chondrophore and internal ligament well developed. Posterior adduc- tor muscle scar kidney-shaped in outline and with irregular transverse, often anas- tomosing impressions that become more numerous with age. Pedal retractor scar more or less oval in outline and located just anterior to the embayment in the pos- terior adductor scar. Siphonal retractor scars small, not impressed, located just posterior to the umbonal—ventral ridge about midway between the umbo and the ventral condyle. Mesoplax small, composed of two flat to slightly arched triangular plates lacking a ventral portion, set in a tentlike fashion and held in place by the periostracum and the anterior adductor muscle. Plates usu- ally meet dorsally at 90° (Plate 5, Fig. 3), occasionally at an obtuse angle. They are rarely bent longitudinally (Plate 8, Fig. 3). Outer surface of plates sculptured with ridges paralleling the anterior margin and lining up with the denticulated ridges of the valves where they come in contact. Siphons short, extending only about one third the length of the valves. Excurrent siphon about one half the diameter of and slightly shorter than the incurrent siphon, with a well-developed sphincter muscle surrounding the aperture and two large papillae on each side (Plate 7, Fig. 3). In- current siphon has minute cirri surround- ing the aperture and a second set of larger cirri anterior to them within the siphon (Plate 6, Fig. 4). Posterior end of burrow lined with coarse, loosely compacted fecal pellets. Young held by byssus threads on the pos- terior dorsal surface of the adult shell (Plate 7, Figs. 1, 2). Measurements. See Table 4. Remarks. This species is most closely re- lated to X. africana Knudsen from the 240 TABLE 4. MEASUREMENTS OF XYLOPHAGA CLENCHI. Length Height (mm) (mm) Location Des 2a Ingolf, station 67 B.D 2.0 Atlantis II, station 124 3 3.5 holotype 4.0 3.5 Tongue of the Ocean 5.0 4.8 Tongue of the Ocean 10.0 9.5 Tongue of the Ocean 10.0 10.0 Pillsbury, station 104 14.0 13.5 Pillsbury, station 394 Gulf of Guinea, West Africa (1°42'N, 7°51'E in 2,550 m), but the mesoplax dif- fers in having a longer median line where the two plates meet, a weaker sculpture, and in lacking the rounded nodule on the ventral surface. Unfortunately, the valves of the two known specimens of X. africana were too fragmentary to allow comparisons on this basis. The siphons of these two species are similar but X. clenchi differs in having only two rather than three large cir- ri on either side of the excurrent siphon and in having two rings of small cirri bor- dering the aperture of the incurrent si- phon. The incurrent siphon of X. africana lacks cirri, according to Knudsen (1961). Two other species, X. wolfft Knudsen from the Sulu Sea and X. murrayi Knudsen from off Zanzibar, also have similar dorsal plates. The former differs from X. clenchi in having denticles on the basal margin of the mesoplax with corresponding denticles on the umbonal reflection. In addition, on each side of the excurrent siphon of X. wolffi there are “7 finger-like tentacles on a common base and somewhat larger ten- tacle dorsally.” The siphons of X. murrayi differ in having a circle of about 35 “short tentacles” surrounding both openings. Knudsen (1961) compared X. wolffi with X. supplicata Taki and Habe but exami- nation of paratypes of X. supplicata re- ceived from Habe showed that species to have tubules on the mesoplax. (See also under X. swpplicata Taki and Habe.) Xylophaga clenchi Turner and Culliney also appears to be closely related to X. panamensis Knudsen but the latter differs Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 | in having the plates of the mesoplax | smooth. The posterior adductor muscle | scar of X. panamensis is oval and irregu- | larly lobed anteriorly, the weak umbonal— ventral ridge is bounded posteriorly by a | groove, and the condyles are lacking. The | siphons of X. panamensis are unknown. | The variation in the mesoplax of X. clen- | chi is interesting and presents some prob- | lems. In most specimens, the plates are flat | or nearly so and none has a ventral portion | or cavity of any kind. The angle at which | the plates meet dorsally is typically acute; however, specimens from shallower waters | (i.e., less than 900 m) occasionally have plates that meet at an obtuse angle. Two | of the five specimens from Atlantis II, sta- tion 124, have the dorsal plate bent lon-— gitudinally at a right angle (Plate 8, Fig. 3), whereas others are typical. Because the shell characteristics of these specimens fit within the range of variation of the typical | forms, they are considered to be ecologic: variants. Unfortunately, none of the spec-— imens from shallower water has extended siphons, so comparisons cannot be made on that basis. | Although the holotype of X. clenchi is small, it was selected because it was the most nearly perfect and the only specimen’ for which dorsal plates, young, siphons, | and chimney all were present. The large’ specimen from R/V Pillsbury, station 394, | and the two from Pillsbury, station 104, | were dead when collected. At least one | specimen from all other localities was alive | at the time of collection. The specimens | from the Tongue of the Ocean were from test panels submerged by the NOO from 4 April 1962 to 17 February 1965. For a description of the test site see DePalma (1969). One specimen taken by the R/V | Atlantis IT at station 119 and three speci- mens taken at station 131 were from epi- benthic sled hauls in which no wood or plant material was found. The specimens | were alive and apparently normal although they were badly crushed, for they were mi- | nute and very delicate. This raises the_ question as to whether or not at least some species of Xylophaga can survive in a firm muddy bottom if wood is not available. So far as known, these are the only living specimens of Xylophaga not taken from wood or other plant material, except those that were boring in plastic at the navy test site off California and possibly those spec- imens of Xylophaga foliata Knudsen that came from a station in Macassar Strait from which it was reported that no wood was taken. The specimens from off Iceland were removed from wood dredged by the Ingolf Expedition in 1896. All specimens were small, the largest being 2.5 mm in length. Many had from 1 to 10 large young attached to the valves in the umbonal area (Plate 7, Fig. 2). This record extends the range of the species far to the north but it has been impossible to find any characters to distinguish these specimens from those occurring to the south. The difference in the size of the young attached to the dorsal surface of the spec- imens figured in Plate 5, Figures 1 and 2, and Plate 7, Figures 1 and 2, reflects the age of the larvae, with those in the latter plate having well-developed umbos and appearing fully mature. It is interesting to note the position of the larvae on the two adults and to speculate that perhaps the larvae move gradually toward the umbos as they mature. Range. From off Iceland south to Ve- nezuela in depths ranging from 35 to 4,862 m. Specimens Examined. ICELAND: Ingolf, station 67, S of Eyrabakki (61°30'N, 22°30’W) in 1,836 m. UNITED STATES, NEW JERSEY: Albatross, station 2550, about 160 miles E of Barnegat Bay (39°44'N, 70°30’W) in 1,977 m. VIRGINIA: Albatross, station 2731, off Cape Henry (36°45’N, 74°28'W) in 1,428 m. NORTH CAROLINA: Atlantis II, station 131, about 420 miles E of Currituck Sound (36°28.9’N, 67°58.2’W) in 2,178 m; off Cape Hatteras (35°44'N, 75°15'W) in 35 m. GEORGIA: Pillsbury, station 104, about 80 miles SE of Brunswick (31°00'N, 79°50’W) in 247 m. BAHAMA ISLANDS: Tongue of the Ocean, about 4 miles off NE tip of Andros Island (24°54'N, 77°49’W) in 1,737 m; Tongue of the Ocean (24°53.2'N, 77°40.2’W) in 2,066 m. BERMUDA: At- lantis II, station 124, about 750 miles E of Cape Charles (due N of Bermuda) (37°26’N, 63°59'W) in XYLOPHAGAINAE ° Turner 241 TABLE 5. MEASUREMENTS OF XYLOPHAGA SUPPLICATA. Height (mm) 8.5 8.8 Length (mm) Location paratype MCZ 194820 4,862 m:; Atlantis II, station 119, just S of Bermuda Pillsbury, station 328, about 25 miles off Punta San Blas, Gulfo San Blas (9°55.8'N, 78°59.8’W) in 420— 640 m. VENEZUELA: Pillsbury, station 719, about 100 miles N of Pertigalete Bay (11°35'N, 64°35.4’W) in 770-890 m. Xylophaga supplicata Taki and Habe Plate 9 Metaxylophaga supplicata Taki and Habe, 1950, II- lustrated Catalogue of Japanese Shells No. 7, p. 47, text-figs. 1, 2 (Tosa Bay, Shikoku, Japan, in 100 fathoms). Holotype, T. Habe Collection; paratype, MCZ 194820; Knudsen 1961, Galathea Report, 5: 188. Distinctive Characters. Mesoplax com- posed of two flat triangular plates set at a sharp angle to each other, dorsal margin more than one half the total length, and with minute tubules at the posterior end. The tubules sit deep within the cavity formed by the umbos and cannot be seen when the shell is viewed anteriorly. Chon- drophore of left valve with a large tooth (Plate 9, Figs. 5, 6). Measurements. See Table 5. Remarks. Taki and Habe (1950) de- scribed the dorsal plate of X. supplicata simply as a “small triangular protoplax [=mesoplax]” that is attached vertically. A paratype received from Dr. Habe has small tubes on the dorsal posterior end of the mesoplax. Xylophaga bruuni Knudsen, 1961, from the Mindanao Sea is very close to if not synonymous with X. supplicata. Unfortu- nately, Taki and Habe did not mention the tubes in their original description and Knudsen did not see the types. Also, un- fortunately, Knudsen had only a single specimen; hence, it will be necessary to obtain more material before a definite statement can be made concerning the sta- 242 tus of X. bruuni. (See also Remarks under Xylophaga whoi Turner n. sp.) Specimens Examined. JAPAN: Tosa Bay, Shikoku, in 100 fathoms (183 m). Xylophaga who/* Turner new species Plates 10, 11 Holotype. MCZ 275015. Type Locality. R/V Atlantis, station 3471, off Cardenas, Matanzas Province, Cuba (23°21'N, 80°56’W) in 500 fathoms (914 m). Distinctive Characters. Mesoplax com- posed of two flat triangular plates with a short median line, set at a sharp angle to each other and with a large hollow tube extending outward from the posterior dor- sal surface of each plate. Description. Shell globose, reaching 7 mm in length and 6.2 mm in height, Shan fragile, and with a very thin, light tan per- iostracum. Pedal gape angle walvoue 95°. Beaked portion of the anterior slope sculp- tured with numerous low denticulated ridges; the posterior portion with fine in- distinct ridges. Umbonal-ventral sulcus only slightly impressed and irregularly sculptured. Sculpture on the disc and pos- terior slope consisting of pronounced growth ridges. Umbonal reflection narrow, closely impressed near the umbos, free and vertical anteriorly. Inner surface of valves smooth and shin- ing. Umbonal-ventral ridge becoming ev- ident only near the ventral margin but rap- idly increasing in size to a large ventral condyle. Chondrophore and internal liga- ment well developed; chondrophore of fhe left valve with a small tooth on the poste- rior upper margin. Posterior adductor muscle scar large, elongate, oval in outline, irregularly sculptured, set high on the pos- terior slope and well in from the posterior margin. Pedal retractor scar rather large, elongate, somewhat irregular, and located ‘An acronym for Woods Hole Oceanographic In- stitution, whose research vessel Atlantis collected the holotype. Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 TABLE 6. MEASUREMENTS OF XYLOPHAGA WHOI. Length Height (mm) (mm) Location 2.5 ORS) Gerda, station 266 3.1 3.0 Gerda, station 266 6.9 6.2 holotype 7.0 6.8 Pillsbury, station 394 Me 7.0 Pillsbury, station 944 9.5 9.0 Pillsbury, station 944 12.5 11.5 Pillsbury, station 944 13.8 13.1 Pillsbury, station 944 14.9 14.0 Pillsbury, station 944 just anterior to the posterior adductor scar. Siphonal retractor scar not impressed. Mesoplax composed of two triangular plates that are set at an acute angle to each | other dorsally and have two large, hollow, tubular projections extending ‘from the posterior dorsal margin (Plate 10, Figs. 3— 10). Length of dorsal margin less heen one | half the length of the plate, outer surface — sculptured with distinct ridges paralleling — the anterior margin; inner surface smooth, with a small internal opening into the tube (Plate 10, Figs. 11, 12). Siphons short, of about equal length, and probably not capable of extending more than one half the length of the shell Aperture of incurrent siphon large and ap- parently lacking cirri, that of thes excurrent siphon small ana with a few fine cirri on each side ventrally. Measurements. See Table 6. Remarks. Knudsen (1961) described three species of Xylophaga with tubulate mesoplaxes, and studies for this report have shown that X. supplicata Taki and Habe also is tubulate. Three of these spe- cies are from the western Pacific (X. tub- ulata Knudsen from Macassar Strait, X. bruuni Knudsen from the Mindanao Sea, and X. supplicata Taki and Habe from Ja- pan), and one from the eastern Pacific (X. obtusata Knudsen from the Gulf of Pana- ma). Xylophaga whoi Turner n. sp. is the first tubulate species reported from the At- lantic. It is most closely related to X. bru- uni Knudsen from the Mindanao Sea but differs in that the mesoplax has a propor- tionately shorter dorsal margin and larger tubes. In addition, the posterior adductor muscle scar is set well in from the poste- rior margin and the umbonal-ventral ridge is evident only near the ventral margin of the shell but increases rapidly to a large condyle. Xylophaga whoi Turner n. sp. dif- fers from X. obtusata Knudsen in the po- sition of the posterior adductor scar, the shape of the mesoplax, the larger tubes on the mesoplax, the narrower anterior slope, and smaller umbonal reflection. The third tubulate species described by Knudsen, X. -tubulata from Macassar Strait, differs from X. whoi in having very large tubules that extend to the anterior margin of the me- soplax and in having the plates bent lon- gitudinally at a right angle so they are flat dorsally rather than meeting at an acute angle. Examination of a paratype specimen of X. supplicata Taki and Habe, 1950, from Japan shows that this species has minute tubes on the mesoplax similiar to those in X. bruuni and it may be synonymous with that species. (See also under X. supplicata Taki and Habe.) The specimen of X. whoi Turner n. sp. taken by the R/V Pillsbury from off Punta Piedras, Colombia, had 13 young attached to the umbonal area (Plate 11, Fig. 2). Among the other species in this group (Group 3), Knudsen (1961) reported that the type of X. obtusata had two young and the type of X. bruuni had four young on the dorsal surface of the adult shell, whereas X. tubulata had five young at the ventral base of the siphons in a depression of the mantle tissue. The young at the base of the siphons of X. tubulata were very small, about one half the size of those on X. bruuni. They possibly had only recently been extruded from the excurrent siphon of the adult and had not yet crawled to the dorsal surface of the adult. It has not been proven that the juveniles clinging to the mantle and shells of adult Xylophaga are definitely the young of the specimen to which they are attached, and this has been questioned by some workers. However, until the species concerned are cultured in XYLOPHAGAINAE ° Turner 943 the laboratory, it is the only assumption that can be made safely. The only way the young of another specimen or species could get into the burrow would be via the incurrent siphon. For the veliger larvae of another specimen to get to the umbonal area, they either would have to pass through the gills into the epibranchial chamber and then out the excurrent si- phon, or go through the digestive tract to the excurrent siphon. Neither of these al- ternatives seems likely. It is, of course, pos- sible that the young of X. twbulata com- plete their development at the base of the siphons of the parent. Range. From off southern Florida to Colombia in depths from about 336 to 910 m. Specimens Examined. UNITED STATES, FLOR- IDA: Gerda, station 266, off Fowey Rocks (25°39'N, 79°58'W) in 185-187 fathoms (338-345 m). CUBA: Atlantis, station 3471, off Cardenas, Matanzas Prov- ince (23°21'N, 80°56’W) in 500 fathoms (914 m). LESSER ANTILLES: Pillsbury, station 944, 45 miles N of Port Louis, Guadeloupe Island (16°32.2'N, 61°36.8S'W) in 364-421 m. COLOMBIA: Pillsbury, station 394, off Punta Piedras (9°28’N, 76°26’W) in 230-350 fathoms (419-640 m). Xylophaga profunda Turner new species Plates 12, 13 Holotype. MCZ 316751. Type Locality. Tongue of the Ocean, off NE tip of Andros Island, Bahama Islands (25°54'N, 77°49’W) in 1,722 m. From test panel submerged from 26 July 1962 to 17 February 1965. Distinctive Characters. Valves with a well-impressed umbonal—ventral sulcus, bounded posteriorly by a low broadly rounded ridge. Posterior slope sculptured with narrow, concentric grooves. Mesoplax of two triangular plates, lacking a basal portion, set at an acute angle to each other, folded longitudinally with the anterior margin bent inward and with a small pore in each anteriorly. Umbonal—-ventral ridge on inner surface of valves narrow, high, and strongly segmented. Description. Shell globose, reaching about 11 mm in length, thin, fragile, and QA4 TABLE 7. MEASUREMENTS OF XYLOPHAGA PROFUNDA. Length Height (mm) (mm) Location 5.9 5.6 holotype Tongue of the Ocean, off NE tip of 8.0 7.9 Andros Island, Bahama Islands Tongue of the Ocean, off NE tip of 95 88 Andros Island, Bahama Islands Tongue of the Ocean, off NE tip of 10.0 9.8 Andros Island, Bahama Islands Tongue of the Ocean, off NE tip of 10.9 10.8 Andros Island, Bahama Islands with a relatively heavy, light brown perios- tracum. Pedal gape angle about 95°. Beaked portion of anterior slope sculp- tured with numerous strong, denticulated ridges that are more closely spaced toward the ventral margin in adults. Umbonal— ventral sulcus narrow, deep, bounded pos- teriorly by a low, broad, rounded ridge. Posterior slope sculptured with widely spaced, incised grooves. Umbonal reflec- tion narrow, high, and strongly segmented. Incised grooves of posterior slope ex- pressed internally as faint ridges just pos- terior to the umbonal-ventral ridge giving a “back bone and ribs effect.” Posterior ad- ductor muscle scar broadly oval, set high on the posterior slope, and lightly marked with irregularly anastomosing depressions. Pedal retractor scar lightly impressed, dumbell-shaped, and located just anterior to the midportion of the posterior adduc- tor. Siphonal retractor scar large, elongate, lightly impressed, and located about mid- way between the posterior adductor and the umbonal-ventral ridge. Chondrophore large, that of the left valve with two small teeth on the posterior edge. Mesoplax composed of two triangular plates lacking a basal portion, set at an acute angle to each other, folded longitu- dinally with the anterior margin bent in- ward and with a small pore in each ante- riorly at the outer edge of the fold. Measurements. See Table 7 Remarks. This species is superficially similar to X. abyssorum Dall but differs in Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 having a low, broad, rounded ridge poste- _ rior to the umbonal—ventral sulcus, in lack- | ing the deep groove posterior to the um- bonal—-ventral ridge on the inner surface, in having a pronounced sculpture on the posterior slope, and in being larger. In ad- dition, the mesoplax of X. profunda lacks a ventral portion, has only a single pore in each plate, does not produce tubes, and appears narrow when viewed anteriorly. Xylophaga profunda Turner n. sp. is probably most closely related to X. lobata Knudsen from the Sulu Sea. Xylophaga profunda differs in having a mesoplax that is much longer than wide, whereas in X. lobata the structure is wider than long. Both species have a strongly segmented umbonal-ventral ridge and a similarly © marked posterior adductor scar, although in X. profunda the markings are more ex- tensive and elaborate. Knudsen (1961) compared his X. galatheae from the Tas- man Sea with X. lobata, noting differences in the mesoplax and muscle scars. Unfor- tunately, he had only a single complete specimen of X. galatheae, so he could not determine range of variation. Xylophaga galatheae possibly is a young X. lobata but more material from intervening areas is needed to ascertain this. On the basis of the general shape and attachment of the dorsal plates, X. profun- da, along with X. lobata and X. galatheae, appear re belong in species Group 3, al- though X. profunda is not closely related to any species in the group and is placed here tentatively. The major differences in- clude the sculpturing of the posterior ad- ductor muscle scar, the presence of ante- rior pores, and the lack of posterior tubes on the mesoplax. The chondrophore of the left valve of X. profunda has two small teeth, a character that relates it to X. whoi and X. supplicata, both of which have a single large tooth. Unfortunately, Knudsen digh not mention the presence of teeth in the three tubulate species he described. Consequently, it cannot be stated definite- ly that this is characteristic of the group. The siphons of X. profunda are short, probably of the same length and capable of complete retraction ation the valves. Unfortunately, none of the specimens was sufficiently well preserved for anatomical work. Four specimens carried young on the umbonal area, the smallest had 5 and the largest had 75. The young have pro- nounced umbos, distinct concentric sculp- ture, and measure about 0.30 mm in length. Xylophaga profunda Turner n. sp. is known from five specimens taken from one panel suum cee in the Tongue of the Ocean at 25°54'N, 77°49’'W, Aa from 14 specimens taken from an asbestos-backed panel submerged at 24°53.2'N, 77°40.2'W. Specimens Examined. BAHAMA ISLANDS: Tongue of the Ocean, off NE tip of Andros Island (25°54 N, 77°49'W) in 1,722 m; Tongue of the Ocean, Tower 3 (24°53. 2'N, 77°40. 2'W) in 2 5066 m. Xylophaga abyssorum Dall Plates 14-16 Xylophaga abyssorum Dall, 1886, Bulletin Museum of Comparative Zoology, 12: 317, pl. 9, fig. 7, 7a (Blake, station 215, off St. Lucia, Lesser Antilles [13°51'N, 61°03'W] in 226 fathoms). Holotype, MCZ 8135; Turner, 1955, Johnsonia, 3(34): 156, pl. 93. Distinctive Characters. Valves with a prominent ridge just posterior to the um- bonal-ventral sulcus and with a slightly to strongly concave profile posterior to the ridge when viewed dorsally. Umbonal— ventral ridge on the inner surface devel- oped only near the ventral condyle and bounded posteriorly by a deep groove. Mesoplax composed of two more or less triangular plates having a ventral portion and a variously lobed dorsal portion; lobes varying with age and coalescing to form pores or tubes (Plate 15, Figs. 4, 5). Description. Shell globose but appear- ing constricted posteriorly when viewed dorsally, reaching 5.5 mm in length, thin, fragile, and with a thin, light brown per- iostracum. Pedal gape angle about 95°. Beaked portion of the anterior slope sculp- tured with fine, denticulated ridges, widely spaced in young specimens but compacted XYLOPHAGAINAE ° Turner IA5 toward the ventral margin in older speci- mens. Umbonalventral sulcus narrow, rather shallow, and with a narrow promi- nent ridge just posterior to it. Profile of valves concave posterior to the ridge when viewed dorsally. Posterior slope sculptured with faint concentric growth lines, umbo- nal reflection erect, the ventral margin of the mesoplax meeting it and ateached by a periostracal fold. lWnibes inflated, particu- larly in young specimens. Inner surface of valves white and glazed. Muscle scars well marked. Posterion ad- ductor scars somewhat pear-shaped, set well in from the posterior margin, and marked with irregular, elongate depres- sions extending inward from the posterior margin. Anterior adductor scar covering most of the umbonal reflection. Pedal re- tractor scar nearly circular and located just anterior to the widest position of the pos- terior adductor scar. Siphonal retractor scar small, elongate, lightly impressed, and located anterior and ventral to the pedal retractor. Umbonal—ventral ridge not well developed except near the ventral condyle but bounded posteriorly by a deep groove that is a reflection of the external ridge. Chondrophore well developed, internal ligament strong. Mesoplax variables with a well-devel- oped, more or less triangular basal portion, occasionally with lateral notches in young specimens. Dorsal portion developing and varying with age; in young specimens, con- sisting of lobes extending anteriorly from the posterior ends of they plates (Plate 14, Fig. 2) followed by lobed lateral folds (Plate 14, Figs. 3-5, 7), which grow and eventually eodlesce to form pores or short tubes (Plate 15, Figs. 4, 5, and Plate 16, Figs. 4, 5). Aperture of the tubes in living specimens covered by a periostracal mem- brane. Siphons short, about equal in length, and capable of complete retraction within the valves. Diameter of the incurrent si- phonal aperture about twice that of the ex- current siphon. The margins of both ap- pear to have cirri. 246 TABLE 8. MEASUREMENTS OF XYLOPHAGA ABYSSORUM. Length Height (mm) (mm) Location 3.0 2.6 Pillsbury, station 944 3.0 2.9 Tongue of the Ocean, off NE tip of Andros Island, Bahama Islands Pillsbury, station 944 Pillsbury, station 944 holotype Gerda, station 266 Pillsbury, station 944 3.3 3.0 Se) ei 4.0 3.5 4.0 3.6 5.4 4.5 Measurements. See Table 8. Remarks. Dall based his description of X. abyssorum on two isolated valves of a dead specimen, with the main distinguish- ing character being the pronounced ridge posterior to the umbonal—ventral sulcus. All specimens that have been assigned to this species since that time have been so named on the basis of this ridge. Among the specimens taken from boards sub- merged in the Tongue of the Ocean off Andros Island, Bahama Islands, by the U.S. NOO (DePalma, 1969), was a single specimen the valves of which coincided with those of the holotype. Two specimens were taken from wood dredged off Gua- deloupe Island, Lesser Antilles. All of these specimens have a pronounced ridge, concave posterior slope, and lobed meso- plax. With this new material, it has been possible to redescribe X. abyssorum giving the characters of the mesoplax and a range of variation. Plate 14, Figure 6, illustrates the dorsal view of the right valve of the holotype of X. abyssorum; Figure 3 illus- trates the dorsal view of the right valve of the specimen from the Tongue of the Ocean. These two valves so closely resem- ble each other that, except for a slight dif- ference in size, they could be from the same specimen. Variations in the valves and dorsal plates of specimens from Flor- ida and Guadeloupe Island are illustrated on Plates 14-16. Plate 15, Figure 5, illus- trates the position in which the young are carried on the adult. The largest number of young on any one specimen was five; Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 the average measurement of the young | was 0.3 mm in length. Xylophaga abyssorum has a small shell. but is a distinctive species, particularly in the adult stage. It is probably most closely | related to X. lobata Knudsen from the | Sulu Sea but differs in having a sharp ridge | posterior to the umbonal—-ventral sulcus | and a far more elaborately lobed mesoplax | in the adult. From X. profunda Turner n. | sp., the only other species with lobes on | the dorsal portion of the mesoplax, X._ abyssorum differs in having broad dorsal plates with two or more pores or tubes in- each. In addition, X. profunda is a much— larger species, the umbonal—ventral ridge on the outer surface of the valves is broad | and rounded, and the deep groove on the inner surface is lacking. The posterior ad- ductor muscle scars of X. profunda are lightly and irregularly marked, whereas the muscle scars of X. abyssorum are well marked. In the young stage, the mesoplax of X. abyssorum is similar to that of X. bay- eri Turner n. sp. and X. profunda, but the valves of these species do not have a pro- nounced ridge. The shape of the valves of X. abyssorum somewhat resembles those of X. japonica Taki and Habe but the me- soplax of that species is ear-shaped and not lobed. (See under X. japonica.) Range. Based on the valves of dead specimens lacking a mesoplax, the range of this species extends from off Atlantic City, New Jersey, south to St. Lucia, Lesser An- tilles (Turner, 1955: 157). Living speci- mens are known only from Florida, the Bahamas, and Guadeloupe Island, Lesser Antilles, in depths ranging from 342 to GO soi, } I) | | | f Specimens Examined. UNITED STATES, FLOR- IDA: Gerda, station 266, off Fowey Rocks (25°38'N, 79°58'W) in 185-187 fathoms (338.3-342 m). BA- HAMA ISLANDS: Tongue of the Ocean, about 4 miles off NE tip of Andros Island (24°54'N, 77°47'W) in 1,722 m. LESSER ANTILLES: Pillsbury, station 944, 4.5 miles N of Port Louis, Guadeloupe Island (16°32.2'N, 61°36.8'’W) in 360-420 m. TABLE 9. MEASUREMENTS OF XYLOPHAGA DUPLICATA. Length (mm) Height (mm) Location Bia 3.5 holotype 2) 5.0 Gulf of Tehuantepec, Mexico 5.5 5.0 Gulf of Tehuantepec, Mexico Xylophaga duplicata Knudsen Plate 17 Xylophaga duplicata Knudsen, 1961, Galathea Re- port, 5: 175, figs. 14, 15 (Galathea, station 745, Gulf of Panama [7°15'N, 79°25’W] in 915 m). Ho- lotype, Zoological Museum, University of Copen- hagen. Distinctive Characters. Plates of the mesoplax elongate oval, with a large ven- tral portion, somewhat inflated, diverging anteriorly, and standing off from the sur- face of the valves. Anterior adductor mus- cle extending into the cavity of the meso- plax. Anterior slope sculptured with nu- merous exceedingly fine, closely set den- ticulated ridges. Umbonal reflection narrow and erect. Umbonal—ventral sulcus narrow, slightly impressed, and bounded posteriorly by a faint, rounded ridge. Pos- terior adductor muscle scar oval and smooth. Umbonal-ventral ridge narrow, high, and segmented; ventral condyle small. Siphons of equal length, united ex- cept at the tip, the posterior three-fourths covered with a brown periostracal sheath; siphonal openings (apertures set on two short tubes) each with six to eight cirri. Measurements. See Table 9. Remarks. At the time Knudsen de- scribed this species he had only two spec- imens from the Gulf of Panama. Two ad- ditional specimens received from D. Shas- ky were taken by the San Juan Expedition at station N-12 from a sunken log dredged in 60 fathoms (109 m) in the Gulf of Te- huantepec, Mexico (15°08'N, 93°28’W). Although the specimens were in rather poor condition, the valves and siphons agree with those described by Knudsen. The mesoplax stands off from the surface of the shell and, as stated by Knudsen, is XYLOPHAGAINAE ° Turner IAT double, that is, has a basal portion, and the anterior muscle extends into the cavity of the mesoplax. The siphons are the same length, combined in a common sheath ex- cept at the tip, and are covered with a thin, brown periostracum. The white spots mentioned by Knudsen are not evident, but the few cirri surrounding the siphonal openings are similar. Range. This record extends the geo- graphic range of the species about 500 miles to the north. The depth range is from 1O9 to 915 m. Xylophaga muraokai> Turner new species Plates 18, 19 Holotype. MCZ 316746; paratypes, MCZ 316747, 316748, 316749, 316750. Type Locality. U.S. Naval Civil Engi- neering Laboratory Test Site I, about 81 miles SW of Port Hueneme, California, or about 25 miles S of San Miguel Is- land, Santa Barbara Islands (33°44'N, 120°45’W) in 5,640 ft (1,720 m). The ho- lotype was taken from a panel exposed on STU I-2 from October 1963 to October 1965. Distinctive Characters. Plates of the mesoplax wedge-shaped, the basal portion large and the small dorsal portion covering only the posterior part of the muscle. Si- phons smooth, of unequal length, the ex- current slightly shorter and with 8-10 prominent cirri. Description. Shell globose, valves reach- ing 14 mm in length and 13 mm in height; the width of apposed valves about 12 mm. Valves thin but strong, white, with a thin clear, transparent to bright yellow perios- tracum. Angle of the pedal gape about 115°. Anterior slope sculptured with nar- row, finely denticulated ridges that are widely spaced during early growth, becom- ing more closely spaced toward the ventral margin in older specimens. Holotype with 17 ridges on the anterior slope. Umbonal— ventral sulcus rather narrow, only slightly > Named for James Muraoka, Biologist, U.S. Naval Engineering Laboratory, Port Hueneme, California. 248 depressed and sculptured with fine growth lines. Disc and posterior slope sculptured with growth lines only. Inner surface of valves smooth and shin- ing. Umbonal-ventral ridge narrow and marked with distinct transverse ridges that appear knobby in some specimens. Chon- drophore and internal ligament well de- veloped. Posterior adductor muscle scar large, elliptical, covering most of the pos- terior slope, and marked with irregular, faint depressions that generally cross the short axis of the scar. Pedal retractor mus- cle scar broadly oval in outline and located just anterior to and about midway on the posterior adductor scar. In some speci- mens, the two scars appear to be adjacent. Siphonal retractor muscle scars not clearly defined but located just posterior to the umbonal-ventral ridge about midway dor- soventrally. Ventral ductor muscle scar long, narrow, and traversing the ventral endl of the bonalevenerall ridge. Ante- rior adductor muscle scar covering the umbonal reflection and the ventral flange of the mesoplax. Mesoplax small and divided. The ventral portion is a large, curved, wedge-shaped shield, fitting closely against the surface of the valves. Posteriorly, the mesoplax curves upward and forward covering the posterior portion of the anterior adductor muscle (Plate 18, Figs. 3-6). Siphons smooth, united, and capable of extending over three times the length of the valves. Excurrent siphon only slightly shorter than the incurrent siphon. Aper- ture of the incurrent siphon bordered by 18—22 long, slender cirri that, in preserved specimens, curl inward and are not easily seen. At the base of these are numerous small cirri and, anterior to them on the inner wall of the siphon, 8-10 broad, branched cirri. Distal margin of excurrent siphon with two broad, short cirri on each side adjacent to the incurrent siphon, and 8-10 long, slender cirri, the longest locat- ed middorsally, with those on either side becoming progressively shorter. The anterior adductor muscle inserts Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 TABLE 10. MEASUREMENTS OF XYLOPHAGA MURAOKAI. Length Height (mm) (mm) Location 6.5 6.2 paratype 8.5 8.0 paratype 11.8 12.0 paratype 12.0 11.5 paratype 12.0 11.9 paratype 12.8 12.8 holotype 14.0 13.0 paratype both on the anterior reflection and on the basal portion of the mesoplax. Siphonal re- tractor muscles straplike, extending ante-— riorly from the base of the siphons along the midportion and over the outer surface of the visceral mass to insert on the valves just posterior to the umbonal—ventral ridge. Ventral adductor muscle, formed as a thokened area of the fused mantle mar- gin, inserts over and to each side of the ventral condyles. Pedal retractor muscles pass through the visceral mass and insert just anterior to the posterior adductor muscle. Measurements. See Table 10. Remarks. The young of X. muraokai Turner n. sp. superficially resemble those of X. concava Knudsen, found in the Gulf of Panama, but differ in having a mesoplax with a large ventral portion, in having the excurrent siphon slightly shorter than the incurrent siphon, and in having a few large cirri on the excurrent siphon. From X. du- plicata Knudsen, also from the Gulf of Panama, X. muraokai differs in lacking the brown periostracal sheath covering the si- phons, in having an elongate rather than nearly circular posterior AR ennettor scar, and in having the dorsal portion of the meso- plax reduced, covering only the posterior portion of the anterior adductor muscle. As with all Xylophaga known to date, specimens of X. muraokai show consider- able variation in size and sculpture de- pending on the type of wood in which they are boring. A specimen boring in green- heart with valves only 6 mm in length had 45 closely spaced denticulated ridges on the anterior slope, whereas a specimen of the same size in white pine had only 7 ridges. Although it is impossible to tell the exact age of these specimens, we know that the greenheart panel was exposed for 35 months and the white pine panels were exposed for 24 months. Other specimens in a white pine panel (exposed on the same rack for the same length of time as the greenheart panel) reached 12 mm in length and had only 24 ridges. Large numbers of X. muraokai were taken from panels submerged for 2 years at 5,640 ft (1,720 m). The openings of the burrows averaged about 1 mm in diameter and increased rapidly. When specimens were uncrowded, the anterior end of bur- rows that had reached 15 mm in length averaged 12 mm in diameter; burrows 55 mm long averaged about 15 mm in diam- eter anteriorly. The tunnels often ran to- gether and frequently two to six specimens occupied a large irregular cavity. Appar- ently, several specimens occasionally used a single opening to the surface. Xylophaga muraokai Turner n. sp. is known only from the USNCEL Test Site I. It has never been taken from panels sub- merged at Test Site II, which is just north of San Miguel Island in about 2,370 ft (722 m). Differences in temperature, dissolved oxygen content of the water, and hydro- static pressure between the two sites are probably the responsible factors. It is im- possible to say whether or not they are all of equal importance. Xylophaga washingtona Bartsch, a spe- cies of wide geographic and depth range, has been taken from wood exposed at Test Site I. It is the only species found at Test Site II, where it is abundant. For a com- parison of the test sites, see Table 11. For a description of the test sites, see Muraoka (1964, 1965, 1966a, 1966b, 1966c, 1967). Range. Known only from USNCEL Test Site I. Specimens Examined. UNITED STATES, CALI- FORNIA: USNCEL Test Site I (33°44'N, 120°45’W) about 81 nautical miles SW of Port Hueneme (about 25 miles S of San Miguel Island, Santa Barbara Is- XYLOPHAGAINAE *° Turner YAY TABLE 11. COMPARISON OF THE U.S. NAVAL CIVIL ENGINEERING LABORATORY TEST SITES (AVERAGE VALUES TAKEN FROM MURAOKA, 1967). Tempera- Dissolved Hydrostatic Depth ture Oxygen Pressure Locality (ft) (EG) (ml/L) (psi) Test Site I 5,640 2.5 1.26 2,482 Test Site IT 2,370 5.0 30 1,043 lands); STU I-1 at 5,300 ft (1615.4 m) exposed March 1962—February 1965; STU I-2 at 5,640 ft (1,720 m) exposed October 1963-—October 1965; STU I-4 at 6,800 ft (2,072.6 m) exposed June 1964—7 July 1965; STU I-5 at 6,000 ft (1,828.8 m) exposed 25 August 1965-12 February 1966. Xylophaga atlantica Richards Plate 20 Xylophaga atlantica Richards, 1942, Nautilus, 56: 68, pl. 6, fig. 4 (east coast of the United States). Ho- lotype, Academy of Natural Sciences Philadelphia 178741; Turner 1955, Johnsonia, 3(34): 152-154, pl. 91, figs. 1-6 (type locality, Mount Desert Island, Maine). (See Turner, 1955: 153.) [Not Turner, 1954, Johnsonia, 3(33): 5-6, pl. 4 = X. clenchi Turner and Culliney; see below. | Distinctive Characters. Mesoplax small, anterior to and between the umbos, com- posed of two triangular plates that are in contact the length of the dorsal margin and meet at a broadly obtuse angle. Ven- tral portion of the mesoplax narrow, form- ing a small posterior cavity into which the posterior end of the anterior adductor muscle extends. Umbonal—ventral sulcus shallow with a median threadlike groove. Posterior adductor muscle scar elongate and irregularly marked. Excurrent siphon slightly shorter than the incurrent, the ex- current aperture with 15—20 large papillae, incurrent aperture with a double row of numerous minute papillae (Plate 20, Fig. 83), Remarks. Between 16 June 1964 and 16 July 16 1965, the U.S. NOO submerged test panels off Mark Island, Penobscot Bay, Maine, in water 200 ft in depth. The panels were set in a vertical array at depths of 50, 100, 150, and 195 ft. Dissection of these panels showed that the bottom one had 85 specimens of X. atlantica, the one 250 at 150 ft had six specimens, and the one at 100 ft had only one specimen. This sug- gests that, like X. washingtona Bartsch, X Feet Turner n. sp., x depalmai Tain: er n. sp., and probably all Xylophaga, the larvae of this species do not rise very high in the water column and that the greatest decrease in attack occurs within a few feet of the bottom. Probably a panel touching or partially submerged in the substrate at this locality would Thewe been heavily at- tacked. These tests also showed that the valves of X. atlantica may reach a length of at least 10.5 mm in a year. Several spec- imens removed from wood dredged off Ipswich, Massachusetts, in about 80 m av- eraged 14 mm in length. The maximum length for X. atlantica appears to be 15 mm. The Ipswich specimens were well pre- served and allow a more detailed descrip- tion of the siphons than that given in the paper by Turner (1955) cited above. The siphons can be extended 1.5—2 times the length of the valves. The excurrent siphon is slightly shorter than the incurrent si- phon and is combined with it for most of its length. The excurrent aperture is sur- rounded by 15—20 relatively large papillae that on the dorsal surface appear to be grouped in two lobes. The incurrent si- phon is surrounded by an outer rim of small papillae and an inner one of slightly spac stouter papillae (Plate 20, Figs. 1, Xylophaga atlantica is oviparous and does not brood its young. The reproduc- tion and larval development of X. atlantica are fully discussed by Culliney and Turner (1976); they detail methods of laboratory culture and illustrate the various larval and growth stages. A reexamination of the specimen reported by Turner (1954: 5-6, pl. 4) has shown it to be X. clenchi Turner and Culliney, a species that usually is found with young attached to the parent. The adult of X. clenchi differs from that of X. atlantica in having a mesoplax that lacks a ventral portion, in the shape and type of marking on the posterior adductor muscle Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 scar, in having a broad, rounded ridge pos- | terior to the umbonal—ventral sulcus, and | in the arrangement of the papillae on the siphons. On the basis of the material then avail- — able, I formerly considered this species | most closely related to X. washingtona | Bartsch (Turner, 1956). It is now evident | that these species belong to two different | species groups. In X. atlantica, the excur- | rent siphon is only slightly shorter than the | incurrent siphon and cirri surround the | aperture of both siphons (Plate 20, Figs. 1-3). In addition, the posterior ndameton | scar is irregularly marked, whereas that of | X. washingtona has regular herringbone markings. Range. From Newfoundland south to Cape Henry, Virginia, in depths ranging from about 15 to 1,242 m. Specimens Examined (New Records Since Turner, 1955). CANADA, NEWFOUNDLAND: % miles off Argentia in 18.3 m (test panel). NOVA SCOTIA: North Bay, Cape Breton Island (46°20'N, 61°50’W) in 128 m. UNITED STATES, MAINE: off Mark Is- land, Penobscot Bay, in 15.2 m, 30.5 m, 45.7 m, and 59.5 m (test pa anels). MASSACHUSETTS: 15 miles off Ipswich in 73 m; NE of Cape Cod Light off Truro in 40 m: E of Nantucket Island (41°23'N, 68°46’W). Xylophaga washingtona Bartsch Plate 21 Xylophaga washingtona Bartsch, 1921, Proceedings of the Biological Society of Washington, 34: 32 (San Juan Island, Washington). Holotype, U.S. National Museum (USNM) 334478; Turner, 1955, Johnson- ia, 3(34): 154, pl. 92; Turner, 1956, Nautilus, 70: 10; Tipper, 1968, Ecological Aspects of Two Wood- Boring Molluscs from the Continental Terrace Off Oregon, Thesis, Department of Oceanography, Oregon State University, pp. 8-13, 64-115. Distinctive Characters. Mesoplax small, anterior to and between the umbos, com- posed of two triangular plates that are in contact for the length of the dorsal margin and are set at a moderately obtuse angle to each other. Ventral portion of the me- soplax usually greater than one half the width of the dorsal portion and keeled. Umbonal-ventral sulcus broad and _shal- low. Posterior adductor muscle scar elon- gate oval in outline with regular herring- XYLOPHAGAINAE ° Turner 251 TABLE 12. PENETRATION INTO WOOD OF XYLOPHAGA WASHINGTONA AT VARIOUS DEPTHS AND EXPOSURE TIMES. Locality Depth (m) Exposure Reference 3 miles off Oceanside, California 109 4 months Turner, 1956 STU* II (1 and 2), California 722 STU* I (1-4), California 1,615—2,066 19 miles off Depoe Bay, Oregon 200 25 miles off Depoe Bay, Oregon 500 40 miles off Depoe Bay, Oregon 1,000 * STU, submersible test unit. bone markings. Proximal end of the com- bined siphons usually having a thin perios- tracal sheath. Excurrent siphon one third to one half the length of the incurrent si- phon, truncated, and having a narrow ridge extending from each side for a short distance along the dorsal surface of the in- current siphon. Excurrent siphonal open- ing small, located at the end of a short fibe extending between the lateral ridges, and apparently lacking papillae (Plate. OL. igs.) 125)'3); Ineur tent siphonal opening margined by 15 inwardly extending papil- lae (Plate 21, Fig. 4). Remarks. In recent years a great deal has been added to our knowledge of the distribution of X. washingtona, ane inter- esting observations have been made con- cerning its ecology and variation. The many specimens taken from the STU pan- els exposed by the USNCEL off San Mi- guel Island, California, have shown that a great deal of variation exists in the shape and sculpture of the valves as well as the size of the posterior adductor muscle in response to the hardness of the substrate in which they are boring. This is discussed in the section on Variation. From these tests we have also learned that larval and adult X. washingtona can tolerate a wide range in dissolved oxygen concentration. They were the only borers taken from the STU II test site where the concentration was 0.30 ml/L. The temperature at this site was 5.0° C (Muraoka, 1965). Based on adults dissected from new wood exposed at various sites with known dates and 6.5, 13.4 months 4, 13, 25, and 35 months 38 and 72 days 2 months 2 months Muraoka, 1965, 1967 Muraoka, 1964, 1966a, 1966b, 1966c Tipper, 1968 Tipper, 1968 Tipper, 1968 depth of submergence, it has been possi- ble to determine the depth at which the larvae can successfully penetrate. These data are summarized in Table 12. A single large specimen was taken in 18.3 m from ‘wood that had been removed from old street cars and used to make Hermosa Reef, % mile west of Hermosa Beach, Santa Monica Bay, California. This is the shallowest record known for the set- tlement of larvae. The apparent scarcity of specimens suggests that this is the upper limit of the depth range. Living specimens have been taken from a fir log dredged from 73 m off Vancouver Island: Breen Colombia. Although no testing has been done in this area, it is likely that, in these colder waters, X. washingtona could occur at shallower depths. Muraoka (1966c) found that “wood specimens which were exposed near the sediment were damaged considerably more severely than Moca specimens which were exposed about 3 feet above the sed- iment.” This, he said, “indicates that the deep sea borers are very active in large numbers immediately above the aethinrent layer and that their numbers tend to de- crease in seawater as the distance from the sediment layer increases.” Tipper (1968) showed that the settlement of larvae of X. washingtona off Depoe Bay, Oregon, was dence in proximity to the sea sediment interface and that the drop in borer pen- etration usually occurred within the first 6 cm upward from the interface. He also showed that, from the initial penetration bo bo te) very close to the sea—sediment interface, the attack progressed upward with increas- ing time of exposure. Analyses of these data suggest that 1) the free-swimming lar- vae of X. washingtona probably do not rise more than 1O—25 ft above the sea floor; 2) as the attack increases, competition occurs for space; and 3) the second crop of larvae, prevented from settling on the lower levels because of the activity of the siphons of specimens already in the wood, are kept swimming and eventually settle on surfac- es higher up in the water column. Tipper (1968) also pointed out that the depth of penetration decreased with in- creased density of the wood. Cedar, pine, fir, and oak panels exposed for 50 days at a depth of 200 m showed an average pen- etration of 3.8 mm in cedar, 3 mm in pine, IS mm) in fir, and 0:5 mim in oak. De- crease in depth of penetration also is re- lated to increase in depth of exposure. This, he postulated, may be correlated with lower temperatures resulting in slow- er growth rates or with the increased den- sity of the wood from compression at great depth. These observations agree with those ob- tained from the tests off southern Califor- nia (see section on Variation in the Intro- duction). From the STU samples we found that the softer the wood (except balsa), the deeper the Xylophaga penetrated and the greater was the development of the chim- ney of fecal pellets. In a cedar panel ex- posed on STU I-5 for 6 months, two sets of X. washingtona were found; the earlier set had tunnels averaging 10-12 mm in length, and the second set had tunnels av- eraging 5 mm in length. Another panel on STU ie 1 exposed fork 35 months had spec- imens with tunnels reaching 35 mm in length. These tunnels averaged 10 mm in ghoumeter at the anterior end; a chimney of rather coarse fecal pellets lined the pos- terior 15 mm of the burrow. Considering the greater depth of the STU panels and realizing that the rate of growth decreases with age, these growth rates agree with those reported by Tipper (1968). Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Based on data gathered to date from test panels, it would appear that X. wash- ingtona probably breeds throughout the year and that the entire larval life is spent in the sea. Young specimens have never been found attached to the shells of the hundreds of adults examined. Range. From Vancouver Island, British Columbia, south to the Santa Barbara Is- lands, California, in depths ranging from 20 to 2,000 m. Specimens Examined (New Records Since Turner, 1955). CANADA, BRITISH COLOMBIA: Satellite Channel, between Saltspring and Vancouver Islands in 93 m (C. Carl). UNITED STATES, OREGON: about 30 miles W of Seaside (46°00’N, 124°48’W) in 464.5 m; about 50 miles W of Tillamookhead (45°54'N, 125°09’W) in 1,554.4 m; about 40 miles W of Silver Point (45°52'N, 124°54'W) in 822.9 m (all Oregon State University); 19 miles W of Depoe Bay (44°59’ N, 124°54’W) in 200 m; 25 miles W of Depoe Bay (44°52'N, 124°36'W) in 500 m; 40 miles W of Depoe Bay (44°52/’N, 125°01'W) in 1,000 m (all R. | Tipper). CALIFORNIA: Hermosa Reef, % mile off Hermosa Beach in 18.3 m (J. Fitch); 3 miles off Camp Pendleton pier, Oceanside, in 100 m (F. Snod- | grass); USNCEL STU II (1 and 2), 75 miles W of} Port Hueneme or about 5 mi NW of San Miguel L., Santa Barbara Islands (34°06’N, 120°42’W) in 722 m;_ USNCEL STU I (1, 4, and 5), 81 miles SW of Port. Hueneme or 25 miles SW of San Miguel Island, San- ta Barbara Islands (33°44'N, 120°45’W) in 1,524— 2,066 m (both J. Muraoka); Allan Hancock station 1372-41, foe % mile E of Empire Landing, Santa Catalina Island, Santa Barbara Islands (33°25' 50"N, | 118°24'50"W) in 84 m (Allan Hancock Foundacon 2 miles off Eel Point, San Clemente Island, Santa Barbara Islands in 72.2 m (F. Snodgrass). Xylophaga rikuzenica Taki and Habe | Plate 22 | Xylophaga rikuzenica Taki and Habe, 1945, Japanese | Journal of Malacology (Venus), 14: 112 (off Riku- | zen, Honshu, Japan). Holotype, T. Habe Collec-_ tion; paratype, MCZ 194821. Neoxylophaga riku- zenica (Taki and Habe) 1950, Illustrated Catalogue | of Japanese Shells, no. 7, p. 46, text-figs. 4, 5. | Distinctive Characters. Mesoplax com-. posed of two inflated triangular plates with | a large basal portion and ventral keel; plates in contact both dorsally and ven-| trally for the length of the medial margin (Plate 22, Figs. 5-7). Posterior Adductel muscle scar elongate oval, tapering dorsal-| ly, and with regular herringbone markings. Valves and dorsal plates covered with a heavy, brown periostracum. Umbonal— ventral sulcus wide and deep (Plate 22, Figs. 2, 4). Remarks. Taki and Habe (1945) briefly described this species in Japanese but in- cluded no figures. In 1950, they created the genus Neoxylophaga with rikuzenica as the type species, mentioning the small, tri- angular mesoplax and thick, brown perios- tracum as generic characters. Turner (1956) Plated X. rikuzenica to X. wash- ingtona Bartsch on the basis of the dorsal plates. A paratype specimen, received through the kindness of Dr. Habe, shows that the posterior adductor muscle scar is similar to that of X. washingtona Bartsch, X. praestans Smith, and X. aurita Knud- sen, the other species in this group with simple, unfringed lappets extending from the truncation of the excurrent siphon. Xy- lophaga rikuzenica is closely related to but differs from X. washingtona mainly in hay- ing a heavy, brown periostracum covering the valves and mesoplax as well as in hav- ing a much wider, deeper umbonal—ventral sulcus. Unfortunately, the siphons of X. ri- kuzenica are unknown so comparisons cannot be made on that basis. However, all species with regular herringbone mark- ings on the posterior adductor scar known to date have similar siphons so X. rikuzen- ica can be assumed to belong to the same group. No Xylophaga have been taken north of Vancouver, British Columbia, but continuous collecting across the north Pa- cific may show that X. rikuzenica merges with X. washingtona. Until this is done, the two species are being maintained be- cause they are recognizable and may not merge. Range. Known only from off Rikuzen and Toyama Bay, Honshu, Japan, in depths ranging from 100 to 700 fathoms (183 to 1,270 m) (Taki and Habe, 1950). Specimens Examined. JAPAN: off Rikuzen, Hon- shu in 183 m. XYLOPHAGAINAE ° Turner 253 Xylophaga depalmai® Turner new species Plates 23, 24 Holotype. MCZ 316735. Type Locality. 3.2 miles off Fort Lau- derdale, Florida (26°04'N, 80°04'’W) in 152.4 m (500 ft) in a test panel. Distinctive Characters. Shell globose, anterior slope with fine, closely set, den- ticulated ridges; umbonal—ventral sulcus shallow and bounded posteriorly by an in- distinct ridge; posterior slope smooth. Umbonal-ventral ridge narrow and high. Muscle scars barely impressed and smooth. Mesoplax composed of two inflat- ed, elongate plates that are coiled poste- riorly, vary in shape, and are occasionally fused. Excurrent siphon truncate; lappets on dorsal surface of incurrent siphon with exceedingly fine serrations. Description. Shell globose, valves reach- ing 9.8 mm in length and 9 mm in height, thin, fragile, and with a thin transparent periostracum. Pedal gape angle about 95— 105°. Anterior slope sculptured with ex- ceedingly fine, closely set, denticulated ridges; specimens &§ mm long had 40-66 ridges. Posterior portion of the anterior slope rather wide and the ridges extending to the umbonal—ventral sulcus more coarsely denticulate. Umbonal—ventral sul- cus barely impressed, nearly smooth, and bounded posteriorly by a fine, indistinct ridge. Disc and posterior slope faintly marked with growth lines. Posterior slope with a low, smoothly curved dorsal margin that is not reflected. Inner surface of valves smooth and glis- tening. Umbonal—ventral ridge narrow, high, and not enlarging at the vente: al mar- gin to form a able condyle. Posterior adductor muscle scar barely impressed, rather small, oval, smooth, and set high on the posterior slope. Pedal retractor scar dumbbell-shaped or as two adjacent cir- ° Named for John DePalma, U.S. Naval Oceano- graphic Office, who was responsible for the tests off Fort Lauderdale, Florida; in the Tongue of the Ocean, Bahama Islands; and in Penobscot Bay, Maine. 254 TABLE 13. MEASUREMENTS OF XYLOPHAGA DEPALMAI. Length Height (Gaaaa (mm) Location 6.0 5.5 off Fort Lauderdale, Florida 7.6 7.0 off Fort Lauderdale, Florida 8.0 WED holotype 8.2 8.2 off Fort Lauderdale, Florida 8.5 7.0 off Fort Lauderdale, Florida 9.8 9.0 off Fort Lauderdale, Florida cles just anterior to the posterior adductor scar. Ventral adductor scar large, elongate, irregularly oval, located posterior to the umbonal—ventr: all ridge and paralleling the ventral margin of the valve. Siphonal re- tractor scar small, broadly oval, and locat- ed just posterior to the umbonal—ventral ridge at the level of the pedal retractor scar. Anterior adductor scar covering the umbonal reflection and the lower suintace of the mesoplax. Chondrophore and inter- nal ligament well developed. Mesoplax composed of two _ inflated, posteriorly coiled plates that may be asym- metrical, have a well- developed ventral portion, are longer than wide, with the medial margin Re the plates parallel and occasionally with the two parts fused (Plate 24, Figs. 3-17). Burrow length of specimens with valves 4—8 mm in length varying from 1.5 to 4 times the length of the valves. Posterior end of the burrows usually with a smooth, firmly packed chimney composed of fine wood fragments. Siphons capable of complete retraction within the valves. Excurrent siphon trun- cated just posterior to the valves and lack- ing cirri. Finely serrated lappets extend from the truncation along either side of the dorsal surface of the incurrent siphon. Incurrent siphon with fine cirri surround- ing the aperture. Base of the siphons cov- ered by a fine periostracal sheath that ex- tends along the sides of the siphons and contains Bre: irregular, glasslike granules (Plate 23, Figs. 1, Ay 5), IMeaerements See Table 13. Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 | Remarks. This species is most closely re- | lated to X. guineensis Knudsen from the Gulf of Guinea, West Africa. Xylophaga depalmai differs in being larger, having an | elongate mesoplax, the posterior end of which fits between the umbos, and in hay- ing more coarsely serrated lappets on the siphons. (See also Remarks under X. gui- neensis.) From X. tipperi Turner n. sp., to. which it is also related, X. depalmai differs | in having an elongate inflated mesoplax | that fits between the umbos rather than a) broad flattened one that covers them, in) having finely serrated lappets on the si- phons, in having the granules on the si-. phons only in the periostracum rather than. being embedded in the tissue, and in pro-. ducing a chimney. From X. mexicana Dall, ich it also somewhat resembles, X. dem palmai differs in the shape of the meso- plax, in having narrower, more finely fringed lappets on the siphons, an incon- spicuous ridge posterior to the umbonal— ventral sulcus, and an exceedingly thin: periostracum. In addition, the granular in- clusions on the siphons of X. mexicana are: coarse, chalky, white, and arranged in a single line extending nearly the length of ines siphons. | Variation exhibited in the mesoplax of X. depalmai is unusual and unexpected, for. in most species of Xylophaga the mesoplax, is remarkably uniform unless the specimen: is injured or stenomorphic. The extreme: forms (Plate 24, Figs. 11-15) are suffi-| ciently different to be considered of spe- cific value in this genus. However, in the. more than 300 specimens obtained from) the test site off Fort Lauderdale, connect-, ing forms exist between them. The char-, acters of the shell, the siphon, and soft: parts are similar in all specimens. Conse-| quently, because the large majority are’ typical (Plate 24, Figs. 16, 17), the ex| tremes are considered to be variants. Xylophaga depalmai Turner n. sp. was) the most common species occuring in the. Fort Lauderdale test. It ranged from 1 to} 3.2 miles off shore and occurred in boards placed near the bottom in depths from 100 XYLOPHAGAINAE ° Turner 255 TABLE 14. DATA FOR ALL OF THE SPECIMENS OF XYLOPHAGA DEPALMAI N. SP. FOUND IN THE COLLECTING PANELS OFF FORT LAUDERDALE, FLORIDA. Distance From Bottom Depth of Shore Depth Panel Panel no. (miles) in Feet (m) in Feet (m) 2 panels without 2.5 300 on bottom numbers (91.44) Ww44 1.8 100 on bottom (30.48) H50 2.3 200 100 (60.96) (30.48) H56 23, 200 on bottom (60.96) G62 2.6 300 2.00 (91.44) (60.96) G68 2.6 300 on bottom (91.44) 59 2.3 200 on bottom (60.96) 492, 3.2 500 2.00 (152.4) 60.96) 493 By 500 200 152.4) 60.96) 497 3.2 500 300 152.4) 91.44) 498 Bi) 500 300 152.4) 91.44) 500 3.2 500 400 152.4) (121.92) D301 3.2 500 400 152.4) (121.92) 502 BP 500 400 (152.4) (121.92) 504 3.2 500 on bottom (152.4) 505 BP) 500 495 (152.4) to 500 ft (30.5 to 152.4 m). In addition, a few specimens were found in boards 100, 200, and even 300 ft off the bottom, which means that, in this species at least, the lar- vae range well up in the water column. As can be seen from Table 14, panels on the bottom in 300 ft (91.4 m) were the most heavily attacked. This may reflect cyclic variation in population density because the panels were exposed at different times, or that, because of the configuration of the bottom and the currents, more wood was available on the bottom in this vicinity, supporting a native population and a ready source of larvae. However, this also may nae fa eee Sept. 1961 June 1962 +250 Jan. 1964 Oct. 1964 1 Jan. 1964 Oct. 1964 2 Jan. 1964 Sept. 1964 5 Jan. 1964 Sept. 1964 6 Jan. 1964 Sept. 1964 31 24 Jan. 1964 4 Jan. 1965 3 20 Oct. 1965 13 Oct. 1965 3 20 Oct. 1965 13 Oct. 1966 10 20 Oct. 1965 13 Oct. 1966 5 20 Oct. 1965 13 Oct. 1966 il 20 Oct. 1965 13 Oct. 1966 3 20 Oct. 1965 13 Oct. 1966 2 20 Oct. 1965 13 Oct. 1966 3 20 Oct. 1965 13 Oct. 1966 8 20 Oct. 1965 13 Oct. 1966 6 indicate that this is the optimum depth for this species. According to DePalma (1969), the salin- ity at the test site was uniformly high and the temperature of the water ranged from 24 to 30° C at the surface, from 20 to 25° Ghat 100i mand) frome(itos95~ Cratel ds m. The northward-flowing Florida Current averaged 1.5 knots. Range. From Florida north to Massa- chusetts in depths ranging from about 30 to 174 m. Specimens Examined. UNITED STATES, FLOR- IDA: off Fort Lauderdale (26°04'N, 80°04’W). See Table 14. Additional records include: FLORIDA: 160 256 miles W of Tampa, OTEC station 3089 in 126 m; and OTEC station 3086 in 96 m. RHODE ISLAND: From rosewood panels tied to lobster pot set at 39°57'N, 69°19'W in 45.7 m (Knutton, from Al Ea- gle’s ship Reliance). MASSACHUSETTS: Delaware II, station 369 (42°37'N, 66°27'W) from wood dredged from 174 m. Xylophaga guineensis Knudsen Plates 25, 26 Xylophaga guineensis Knudsen, 1961, Galathea Re- port, 5: 195— 196, fig. 38 (Galathea, Station 52, off West Africa, 01°42’N, 07°51’E in 2,550 m). Holo- type, Zoological Museum, University of Copenha- gen. Distinctive Characters. Valves globose, with exceedingly fine, closely spaced, den- ticulated ridges; umbonal—ventral sulcus barely impressed, bounded by a weak an- terior ridge and a well-marked, narrow posterior ridge. Mesoplax composed of two inflated cornucopialike plates. Description. Valves globose, reaching 2.2 mm in length, thin, and fragile. Pedal gape angle about 90°. Anterior slope sculp- tured with exceedingly fine, evenly spaced, denticulated ridges, there being about 45 in a specimen 2 2. mm long. Uinbonalvene tral sulcus only slightly impressed, bound- ed anteriorly by a low, inconspicuous, rounded ridge and posteriorly by a narrow, well- deaned ridge, occasionally almost bladelike. Disc and _ posterior slope smooth. Inner surface of valves smooth and glis- tening. Muscle scars barely visible. Poetes rior Ralductor scar smooth, lobed anteriorly with the small pedal retractor just anterior to it in the upper embayment. Umbonal— ventral ridge and condyle prominent, with a parallel groove posterior to the ridge. This groove is the internal expression of the ridge bounding the sulcus on the outer surface. Mesoplax composed of two smooth, in- flated, cornucopialike plates, the horns of which curl toward each other (Plate 25, Figs. 3, 6, 8, 9); the two halves usually equal and mirror images but occasionally unequal and varying in size and shape. Siphons about equal to the length of the Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 TABLE 15. MEASUREMENTS OF XYLOPHAGA GUINEENSIS. Length Height (mm) (mm) Location 0.9 Pillsbury, station 260 1.3 NED, Pillsbury, station 260 1.3 1.5 paratype (estimated from Knudsen, 1961, fig. 38b) 1.5 1.3 Atlantique Sud, station 33 | Q® G9) Atlantique Sud, station 33 | shell, capable of complete retraction with- | in the valves. Excurrent siphon truncated, about one third the length of the incurrent. siphon, with narrow and minutely serrated lappets extending from the truncation along the dorsal surface of the incurrent. siphon. Base of siphons covered by a filmy periostracal sheath with fine, irregular, whitish granules imbedded in it along the) sides of the siphons (Plate 25, Figs. 1, 2). Measurements. See Table 15. Remarks. When Knudsen described X.. guineensis he had only “very fragmented shells of 5-6 individuals removed! from a piece of wood; not a single complete shell present.” One specimen had a complete mesoplax although only the umbonal area of the valves was present. The species was diagnosed on the basis of the mesoplax. and the large sculptured, bright yellow prodissoconch. In the course of examining wood | dredged from off the western boast of Af) rica, T obtained four dead specimens, one with a complete mesoplax (Atlantique Sud, station 33), and seven small living speci-| mens, two complete (Pillsbury, station 260), all of which appear to be this species. The two plates of the mesoplax of these specimens are mirror images of each other (Plate 26, Figs. 2-4). As shown under X.| depalmai (Plate 24, Figs. 11, 12), the me-| soplax is not always bilaterally symmetrical. Therefore, these new specimens from off, the western coast of Africa are being con- sidered X. guineensis and a description of : the species based on entire specimens is’ given. Although stenomorphs are commonly found in Xylophaga (see section on Vari- ation), the specimens dissected from the wood dredged by the Atlantique Sud Ex- pedition and the Pillsbury did not appear to be stunted or malformed in any way. They were not crowded and the wood was not particularly hard. Consequently, on the basis of the large number of evenly spaced ridges on the anterior slope, the well-de- veloped mesoplax, the strength of the ridge bounding the ara bomal-rentiel sul- cus posteriorly, and the small size of the specimens, I consider this to be a small species. All the major characters of the siphons mentioned in the description could be seen with a dissection scope at 250%. However, the fringe on the lappets could only be detected under a compound mi- croscope at 430. It was impossible to de- termine whether or not the fringe extend- ed the entire length of the lappet on the two specimens that had extended siphons. The siphons appear similar to those of X. depalmai. Xylophaga guineensis is most closely re- lated to X. depalmai taken from waters off Florida and the Bahamas. Xylophaga gui- neensis differs in being smaller; in having a smooth mesoplax composed of inflated, tubular, cornucopialike plates; in having more numerous and closely spaced, den- ticulated ridges on the anterior slope when comparing specimens of equal size; and in having a larger, more prominent, strongly sculptured prodissoconch. (See also Re- marks under X. depalmai.) The specimens that Knudsen described were taken from wood dredged in 2,550 m but the three lots of new material were from 147, 145, and 46 m. Although X. gui- neensis may extra into deep water, it is probably a fairly shallow water species, and this would agree with the known depth range of X. depalmai, the species to which it is most closely related. The fact that Knudsen’s specimens were all dead and fragmented further substantiates the possibility that the wood was carried into XYLOPHAGAINAE ° Turner 257 deeper water after it was invaded by X guineensis. The continental shelf in this area is narrow and such a movement of wood on the bottom could easily take place. Range. Gulf of Guinea, West Africa, in depths ranging from 46 to 147 m (living material) and in 2,550 m (dead). Specimens Examined. CAMEROONS: Pillsbury, station 260, off Santa Isabel Island (3°45'N, 9°05’E) in 46 m. GABON: Atlantique Sud, station 146, about 46 miles NNE of Port Gentil (0°03'S, 9°07’E) in 147 m; Atlantique Sud, station 33, 35 miles W of Ambri- zette (7°16'S, 12°17'E) in 145 m. Xylophaga mexicana Dall Plates 27, 28 Xylophaga mexicana Dall, 1908, Bulletin Museum of Comparative Zoology, 43(6): 425 (Albatross, station 3422, off Acapulco, Mexico [16°47'N, 99°59'W] in 141 fathoms [257.9 m]). Holotype USNM 122947; Turner, 1955, Johnsonia, 3(34): 150, pl. 90. Distinctive Characters. Umbonal—ven- tral sulcus bounded posteriorly by a nar- row, sharp ridge. Mesoplax ear-shaped, in- flated, the two halves not meeting medially except at the posterior end. Excurrent si- phon truncated; lappets extending along the dorsal surface of the incurrent siphon finely fringed. Siphons with a single row of opaque white granules imbedded along the sides. Posterior quarter of the incur- rent siphon papillose. Description. Shell globose, valves reach- ing 10.5 mm in length and 9.5 mm in height, thin, fragile, ane with a thin, light straw-colored periostracum on the poste- rior slope. Pedal gape angle about 110°. Anterior slope with exceedingly fine, close- ly set, denticulated ridges. Umbonal—ven- tral sulcus rather narrow, moderately deep, and bounded posteriorly by a narrow, sharp ridge. Disc and posterior slope sculptured with fine growth lines. Poste- rior margin broadly rounded. Posterior slope becoming proportionately more elongate with increased size and age, as indicated in the measurements. Umbonal reflection free and recurved anteriorly, ap- 258 pressed posteriorly, and covering the an- terior portion of the umbonal area. Inner surface of the valves white, smooth, and glazed. Umbonal—ventral ridge narrow, slightly segmented, and broadening only slightly toward the ventral margin to form the condyle. Posterior ad- ductor muscle scar well impressed, smooth, broadly oval in outline, set high on the posterior slope and at the posterior margin of the valve. Pedal retractor scars near ly circular, double, and adjacent to the embayment on the anterior margin of the posterior adductor scar. Ventral adductor scar large, irregularly oval in outline, and well posterior to the umbonal—ventral ridge. Siphonal retractor scar small, bi- lobed and located posterior to the um- bonal-ventral ridge at a level correspond- ing to the pedal retractor scars. Anterior adductor scar flection. Chondrophore and internal liga- ment well developed. Mesoplax somewhat variable but usually slightly longer than wide, with the medial margin of its two halves diverging. Plates ear-shaped, inflated, coiled posteriorly, and covering the umbos. Siphons capable of complete retraction within the valves and of extension to at least 1.5 times the length of the valves. Ex- current siphon truncated, about one third the length of the incurrent siphon, and lacking cirri. Lappets extending from the truncation of the excurrent siphon finely fringed. Posterior portion of the incurrent siphon papillose; inner edge of siphonal opening margined with numerous minute cirri. Siphons. with thin periostracal sheath posteriorly and with a single row of white, opaque, irregular granules embedded along the sides. Measurements. See Table 16. Remarks. Xylophaga mexicana Dall was described on the basis of two right valves that were in rather poor eomeition and lacked a mesoplax. Consequently, it has been impossible up to now to relate it to other species. Valves of living specimens taken off California agree with those of the covering the umbonal re- - Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 TABLE 16. MEASUREMENTS OF XYLOPHAGA MEXICANA. Length Height (mm) (mm) Location 4.8 4.5 Malibu Reef, California 5.0 4.2 holotype UD ES Malibu Reef, California 8.0 TS Santa Monica Reef, California 9.0 9.5 Hermosa Reef, California 10.5 9.0 Santa Monica Reef, California holotype of X. mexicana. Dall’s specimen was from much deeper water (141 fath- oms) than the Californian series (about 19 | fathoms) but it could have been advecti- tious at that depth. Xylophaga mexicana | Dall could, of course, be considered a no- men dubium but this would necessitate de- scribing the Californian specimens. Be- cause ihe valves of these specimens resem- _ ble the holotype of X. mexicana sufficiently well to carry the name, it seems best to — establish that taxon firmly on the basis of | this new material. The size of the beaks, the smooth mus- cle scars, and the narrow, rather deep um- bonal-ventral sulcus with a pronounced ridge at its posterior margin relate X. mex-_ icana to Xylophaga g alahoce Sowerby from Chile (Turner, 1955). Xylophaga mexicana | differs from X. globosa in having a much finer fringe on the lappets, in having a se- ries of irregular, opaque granules along the | sides of the siphons, and in having the; pos-- terior portion of the incurrent siphon pa- | pillose. In addition, the mesoplax of X. | mexicana is proportionately much smaller | than that of X. globosa, and the two plates diverge medially. In his original descrip-_ tion, Dall (1908) did not relate X. mexi- — cana to any other species in the genus. One year later (Dall, 1909), he gave the range of X. globosa Sowerby as Go Pan- | ama to Chile but cited no specific locali- ties. The Panamanian specimens on which | he based this range may well have been X. mexicana. | have not seen a specimen of | globosa from north of Chiloe Island, Chile. The valves of X. duplicata some- | what resemble those of X. mexicana; how- ever, the siphons of these two species place them in different groups. (See also Remarks under X. tipperi Turner n. sp. and X. japonica, the species to which X. mexicana is most closely related.) Through the kindness of John Fitch and Charles Turner of the California State Fisheries Laboratory, Terminal Island, California, I received 55 specimens of X. mexicana taken from test panels exposed on the “replication reef experiments” con- ducted by that laboratory. Old streetcars, automobile bodies, concrete shelters, and rocks were dumped offshore in several lo- calities to serve as settling areas for marine organisms and to make the area more at- tractive to fish. Within a short time, the test panels and wooden portions of the streetcars were attacked by teredinids and Xylophaga. Because all the specimens had been dissected from the wood, it is im- possible to say whether or not they pro- duced a chimney. Range. Living specimens known only from Santa Monica Bay, California, in about 35 m. Specimens Examined. MEXICO: Bocochibampo, Sinaloa (dead). UNITED STATES, CALIFORNIA: All from “replication reefs” off Hermosa, Malibu, Re- dondo, and Santa Monica (about 33°50'N, 118°30’W) in about 35 m. Xylophaga tipperi Turner new species Plate 29 Holotype. MCZ 316736; paratype, MCZ Sl6V3S7. Type Locality. 3.2 miles off Fort Lau- derdale, Florida (26°04’N, 80°04’W) in 152.4 m (500 ft) ina U.S. NOO test panel, submerged from October 1965 to October 1966. Distinctive Characters. Umbonal—ven- tral sulcus bounded by low rounded ridg- es. Mesoplax ear-shaped, slightly longer than wide, compressed, and with a sharp peripheral keel. Excurrent siphon truncat- ed; lappets on dorsal surface of incurrent siphon with a coarse fringe. Siphons with a single row of minute glasslike plaques XYLOPHAGAINAE ° Turner 259 embedded along the side and with knobby pustules at the posterior end. Description. Shell globose, reaching 9 mm in length and §.5 mm in height, thin, fragile, with prominent umbos and a dull, light brown periostracum. Pedal gape an- gle about 110°. Anterior slope sculptured with numerous close-set, denticulated ridges, there being 25 on the holotype. Umbonal-ventral sulcus slightly de- pressed, bounded by a threadlike rounded ridge anteriorly and a somewhat heavier one posteriorly. Posterior slope low and sculptured with distinct growth lines. Um- bonal reflection free for most of its length, the ventral edge of the mesoplax fitting be- neath it anteriorly. Inner surface of the valves smooth and glistening. Umbonal—ventral ridge very prominent, nearly smooth, and not greatly enlarged ventrally at the condyle. Chon- drophore and internal ligament well de- veloped. Posterior adductor muscle scar oval, tapering dorsally, only slightly im- pressed, and smooth. Pedal retractor scar kidney-shaped and located adjacent to the anterior margin of the posterior adductor scar, about midway dorsoventrally. Siphon- al retractor scar lightly impressed, located just posterior to the umbonal—ventral ridge at the level of the ventral margin of the beak. Ventral adductor scar usually not vis- ible but located near the ventral margin posterior to the umbonal—ventral ridge. Mesoplax large, ear-shaped, covering the umbos, longer than wide, compressed dorsoventrally, with a sharp peripheral keel, and a coiled early portion. Siphons united, excurrent siphon trun- cated, about one third the length of the incurrent siphon. Lappets extending from the truncation along the dorsal surface of the incurrent siphon with a coarse fringe. Incurrent siphon papillose posteriorly, with a single row of glasslike plaques im- bedded along the side at the juncture of the lappets and the excurrent siphon and with numerous small, broad papillae sur- rounding the inner rim of the aperture. 260 TABLE 17. MEASUREMENTS OF XYLOPHAGA TIPPERI. Length Height (mm) (mm) Location 2.0 2.0 off Fort Lauderdale, Florida 4.6 4.] off Fort Lauderdale, Florida UD 7.0 off Fort Lauderdale, Florida 8.2 Ts) off Fort Lauderdale, Florida 9.0 holotype Burrows reaching 20 mm in length. Fe- cal pellets not formed into a chimney. Measurements. See Table 17. Remarks. This species is closely related to X. mexicana Dall from California but differs in having a compressed, sharply keeled mesoplax; in having glasslike rather than white, chalky plaques imbedded in the siphons; and in having much more coarsely fringed lappets on the siphons. Xylophaga tipperi differs from Xylophaga dorsalis Turton in having a definitely sculptured mesoplax, an excurrent siphon one third to one half the length of the in- current (in X. dorsalis it is truncated at the posterior end of the valves), and in having glasslike plaques imbedded in the walls of the siphons. In addition, on the basis of the limited material now available, X. tip- peri apparently does not build a chimney. Xylophaga tipperi differs from X. depalmai Turner n. sp., also from off Fort Lauder- dale, in having a compressed, sharply keeled mesoplax, a much coarser fringe on the lappets, and in having glasslike plaques imbedded in a single line along the sides of the siphons. In addition, the posterior end of the incurrent siphon of X. tipperi is papillose. Xylophaga bayeri Turner n. sp., which also is found in Florida and the West Indies, differs from X. tipperi and all other species in Group 6 in having a broad, horizontal mesoplax (Plate 31, Fig. 6). Only eight specimens of X. tipperi Turn- er n. sp. were obtained. They were all from panel 505 and were collected along with X. depalmai Turner n. sp. The spec- imens, even the smallest, were consistent in having a broad, compressed mesoplax as Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 well as in the characteristics of the si- | phons. For data on the testing site, see in- formation under X. depalmai. Range. Known only from off Fort Lau-_ derdale, Florida, in 152.4 m. Specimens Examined. UNITED STATES, FLOR- IDA: 3.2 miles off Fort Lauderdale (26°04'N, 80°04'W) in 152.4 m (500 ft) in a test panel. Xylophaga bayeri’ Turner new species Plates 30, 31 Holotype. MCZ 316738; paratype, MCZ 316739. | } { i | | Type Locality. U.S. NOO test site, about | 3.2 miles off Fort Lauderdale, Florida | (26°04’N, 80°04'W) in 152.4 m (500 ft). From test panels submerged from October 1965 to October 1966. Distinctive Characters. Posterior margin — of the umbonal—ventral sulcus bounded by | a pronounced, sharp ridge. Mesoplax of © adult much broader than long, extending | laterally as wings and with a sharp periph- | eral margin. Excurrent siphon truncated, about one third the length of the incurrent siphon. Lappets extending along the dorsal surface of incurrent siphon finely fringed. | Description. Shell globose, reaching 8 mm in length, thin, fragile, with prominent umbos and a heavy, golden-brown perios- tracum covering the shell and the meso-_ plax. Pedal gape angle about 120°. Beaked | portion of anterior slope sculptured with | numerous, closely set, denticulated ridges, | there being 32 on the holotype. Posterior — portion of the anterior slope narrow, the | ridges very closely packed. Umbonal re-_ flection broad, closely appressed over the umbos, free anteriorly, and with a funnel- like pit beneath. Umbonal-—ventral sulcus moderately impressed, sculptured with — growth lines, and bounded posteriorly by _ a pronounced, sharp ridge. Posterior slope rather low and sculptured with well- marked growth lines. 7 Named for Frederick M. Bayer, Rosenstiel School | of Marine and Atmospheric Sciences, University of | Miami, Miami, Florida, who kindly loaned material — collected by the research vessels Pillsbury and Gerda. TABLE 18. MEASUREMENTS OF XYLOPHAGA BAYERI. Length Height (mm) (mm) Location 8.0 EO holotype 4.2 4.0 off Mona Island, Puerto Rico 5.4 4.9 Gerda, station 266 6.0 5.2 Gerda, station 266 6.0 5.8 Gerda, station 266 6.2 6.0 Gerda, station 266 6.8 6.0 Gerda, station 266 8.0 8.2 Gerda, station 266 Inner surface of valves smooth and glis- tening. Umbonal-ventral ridge very prom- inent and distinctly segmented but not greatly enlarged ventrally at the condyle. Chondrophore and internal ligament well developed. Posterior adductor muscle scar elongate oval in outline, tapering dorsally, and smooth. Pedal retractor scar broadly oval and contiguous with the anterior mar- gin of the posterior adductor. Ventral ad- ductor scar large, located adjacent to the groove margining the umbonal—ventral ridge posteriorly. Siphonal retractor scar lightly impressed and located just posterior to the umbonal-ventral ridge on a level with the ventral margin of the beak. An- terior adductor scar well marked and cov- ering most of the umbonal reflection. Mesoplax in young specimens com- posed of a large ventral plate with a small, triangular dorsal portion (Plate 31, Fig. 5). Dorsal plate of the adult is much wider than long, with a sharp perphery (Plate 31, Figs. 2, 3) and pronounced concentric sculpture. Siphons capable of complete retraction within the valves and of extension proba- bly not more than the length of the valves. Excurrent siphon truncated, about one third length of the incurrent siphon. Lap- pets extending from the truncation along the dorsal surface of the incurrent siphon, finely fringed. Periostracal sheath, if pres- ent, extremely thin and lacking calcareous or glasslike inclusions. Measurements. See Table 18. Remarks. Xylophaga bayeri Turner n. sp. is a distinctive species apparently not XYLOPHAGAINAE ° Turner 261 closely related to any other species in this genus. Its smooth muscle scars, ear-shaped mesoplax, truncated excurrent siphon, and fringed lappets place it in Group 6 and the lack of granular inclusions in the perios- tracal sheath of the siphons places it more closely to X. globosa and X. dorsalis than the others in the group. The mesoplax of X. bayeri is basically like that of X. dorsalis but is greatly extended laterally. Only two specimens were obtained from the panels placed off Fort Lauderdale, Florida, and unfortunately these were not sufficiently well preserved for detailed an- atomical work. The 14 specimens taken from wood dredged off Fowey Rocks were in situ, but all were dead and the soft parts had disintegrated, although the mesoplax was in place. All had typical valves but some had unusually wide dorsal plates (Plate 31, Figs. 2, 3). Four complete spec- imens and about 25 dead and disarticulat- ed specimens were extracted from the wood dredged off Mona Island. These were remarkably uniform in the characters of the shell and dorsal plates. Range. From off the coast of Florida probably throughout the Caribbean, in depths ranging from 150 to 365 m. Specimens Examined. UNITED STATES, FLOR- IDA: 3.2 miles off Fort Lauderdale (26°04'N, 80°04’W) in 152.4 m (500 ft) in test panel; Gerda, station 266, off Fowey Rocks (25°39'N, 79°58’W) in about 340 m. PUERTO RICO: Johnson—Smithsonian Expedition, station 37, midway between Mona and Desecho islands (18°11'55"N, 67°42'50’W) in about 365 m. Xylophaga japonica Taki and Habe Plate 32 Xylophaga japonica Taki and Habe, 1950, Illustrated Catalogue of Japanese Shells, No. 7, p. 45, text- figs. 6, 7 (Tosa Bay, Shikoku, Japan, in about 100 fathoms). Holotype, T. Habe Collection; paratype, MCZ 194822. Distinctive Characters. Posterior slope elongate in adult specimens. Umbonal— ventral sulcus well impressed, bounded by an inconspicuous ridge anteriorly and a high narrow ridge posteriorly. Posterior adductor muscle scar smooth. Mesoplax 262 TABLE 19. MEASUREMENTS OF XYLOPHAGA JAPONICA. Length Height (mm) (mm) Location 13.2 10.8 holotype (from Taki and Habe, 1950) 5.8 5.0 paratype Tosa Bay, Shikoku, Japan 3.8 3.0 off Clara Island, South Burma 2.0 1.8 off Java large, covering the upper one half of the ron area, extending over the umbos, ear-shaped, moder ately inflated, with a narrowly rounded periphery and distinct concentric sculpture (Plate 32, Figs. 2-5). Siphons capable of extending two to three times the length of the Hol, Excurrent si- phon about one third the length of the in- current, lappets extending along the dorsal surface of the fourm siphon finely fringed, opaque white granules imbedded along the sides of the siphons. Chimney of coarse fecal pellets lining the burrow. Measurements. See Table 19. Remarks. At the time Taki and Habe (1950) described this species, they did not have the soft parts and therefore could not describe the siphons. However, it is clear from their description and figures that the species belongs to the Hoe salis group (Group 6). Taki and Habe related their species to X. indica Smith, but noted its more elongate posterior area. It further differs from X. indica in its broad meso- plax for, according to Knudsen (1961), the mesoplax of X. indica is “oblong pear- shaped” and his figure [of it] shows it to be nearly three times as long as wide. Xy- lophaga indica is probably more closely re- lated to X. guineensis and X. depalmai n. sp., but because the siphons are unknown, this cannot be stated definitely. Xylophaga japonica appears to be most closely related to X. mexicana Dall but dif- fers in having a larger, more highly sculp- tured mesoplax that extends over the um- bos; in having the posterior slope more elongate; and in having the ridge bounding Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 the umbonal-ventral sulcus more pro- | nounced. Unfortunately, the material from off | Java and South Burma noted in the re- cords below was poorly preserved, but it | was possible to ascertain the basic char-_ acters of the siphons. The specimens from | Java had been kept in alcohol since they were collected by Mortensen in 1929. The | specimens from South Burma, dredged by the Anton Bruun and taken from a small | piece of wood and the husk of a nut, had | been allowed to dry. Range. From Tosa Bay, Japan, south and west to Burma and Java, in depths ranging _ from 183 to 384 m. Specimens Examined. JAPAN: Tosa Bay, Shikoku, in 183 m. JAVA: Danish Java—South African Expedi- tion, station 10, off SE tip of Java (08°36’S, 114°34’E) in 300 m. BURMA: Anton Bruun, station 23, about 77 miles W of Clara Island, South Burma (10°30’N, 96°35'E) in 384 m. : GENUS XYLOPHOLAS TURNER 1972 Xylopholas Turner, 1972, Basteria, 36(2): 97-99. Type Species. Xylopholas altenai Turner, original designation. i eneiied Characters. Valves and me- soplax typical for the genus Xylophaga. Animal long, not capable of retraction within the FAlhios. with a periostracal si- | phonal sheath posterior to the valves and a pair of lateral, chitonlike, siphonal plates | at the posterior end. Siphons short, ex-_ tending between the plates, the siphonal retractor muscles inserted on their inner surface. Gills and visceral mass contained | between the valves as in typical Xylophaga. | Wood-storing cecum large. Remarks. This genus Gutters from Xylo- phaga in having extended excurrent and incurrent annals: contained in a common sheath with siphonal plates at the posterior extremity. These plates probably arose in- | dependently but may be homologous with - the siphonoplax of other pholads G. e., Pho- | ladidea), which was carried posteriorly as the animal elongated. However, the si- | phonal retractor muscles of these species insert on the valves rather than on the si- phonoplax. The siphonal plates of Xylo- pholas probably function as do the pallets of the Teredinidae and the siphonoplax of other pholadids to close the end of the burrow. Embryological studies are needed to prove the affinities of the plates. From Xyloredo, Xylopholas differs in having siphonal plates, a periostracal sheath on the animal posterior to the valves, and in not lining the burrow with a chitonlike or calcareous tube. Range. To date the genus is known only from the type species found off the Lower Florida Keys in the western Atlantic and in the Gulf of Guinea in the eastern At- lantic in depths from 239 to 366 m. (Two specimens were dredged in 2,550 m but these may have been decntOns. ) Xylopholas altenai Turner 1972 Plates 33, 34 Xylopholas altenai Turner, 1972, Basteria, 36(2): 99— 103, figs. 1-12 (Gerda, station 66, about 13 miles SE of Fowey Rocks, Florida [25°25'N, 79°59’] in 366 m). Holotype, MCZ 279315. Distinctive Characters. Animal elon- gate, not capable of retraction within the valves, and with lateral, paddlelike siphon- al plates at the posterior end. Shell similar to that of Xylophaga. Mesoplax composed of two flat, elongate plates that are held in place by the periostracum extending be- tween the beaks dorsally. Posterior adduc- tor muscle scar large and with transverse forking impressions. Young carried on the ventral surface just posterior to the valves. Description. Shell globose, reaching 2.5 mm in length, thin, fragile, and with a rel- atively heavy, golden-brown periostracum covering the valves and mesoplax. Pedal gape angle about 90°. Beaked portion of the anterior slope recurved dorsally and sculptured with numerous strong, dentic- ulated ridges. Umbonal reflection narrow. Umbonal-ventral sulcus narrow and only slightly impressed. Disc and posterior slope sculptured with fine growth lines only. Inner surface of valves smooth and glis- tening. Umbonal-ventral ridge low, indis- XYLOPHAGAINAE ° Turner 963 tinct except near the ventral margin, slightly segmented, and with a small ven- tral Ronee. Chondrophore and internal ligament well developed. Posterior adduc- tor muscle scar lar ge, covering most of the posterior slope, elongate oval in outline and marked with transverse, forking im- pressions. Pedal retractor scar irregularly and broadly oval and located about mid- way on the anterior margin of the poste- rior adductor scar. Siphonal retractor mus- cles inserted on the siphonal plates and collar. Mesoplax small, not filling the gape be- tween the beaks, composed of two flat, elongate, subrectangular plates, somewhat pointed posteriorly, sculptured with fine transverse ridges, covering the dorsal sur- face of the anterior adductor muscle and held in place by the periostracum. Animal long, with a periostracal sheath covering the portion posterior to the valves and ee a pair of lateral, paddle-shaped, chitonlike plates at the posterior end (Plate 33, Fig. 4). The siphons extend be- tween the plates, and the siphonal retrac- tor muscles insert on the inner surface of them. Siphons separate, excurrent siphon longer than the incurrent siphon, the ap- ertures of both with fine cirri. Gills and visceral mass contained entire- ly between the valves, the portion of the animal extending beyond the valves com- posed of a dor eal excurrent and a ventral incurrent canal combined in a common muscular and periostracal sheath, with a chitinous collar and two lateral, paddle- shaped plates posteriorly. Measurements. See Table 20. Remarks. Isolated valves of this species would be difficult if not impossible to dis- tinguish from several species of Xylopha- ga; however, its reduced, flat mesoplax, elongate soft parts, and siphonal plates readily distinguish it from all other species in the Xylophagainae. Nothing is known of the biology of the species except that it has a large wood-storing cecum and, therefore, probably utilizes wood as food. The young are held within the burrow to the late ve- 264 TABLE 20. MEASUREMENTS OF XYLOPHOLAS ALTENAI. Length Height (mm) (mm) Location 1.8 1.8 holotype 1.0 1.0 Atlantique Sud, station 147 1.5 At Galathea, station 52 1.2 1.1 Gerda, station 66 1.9 1.5 Gerda, station 266 2.0 Rel Gerda, station 266 2.0 D5 Gerda, station 266 Obs O55 Gerda, station 266 liger stage when the foot has developed. They are attached to the ventral surface of the animal just posterior to the valves. The number of attached young ranged from two to eight and they averaged 0.28 mm in length. The hinge plate of the young is well developed, with two teeth and a sock- et in the right valve and a corresponding single tooth and two sockets in the left valve (Plate 33, Figs. 1, 2). About 60 specimens were taken at the three stations off Florida and 10. speci- mens were taken from the three stations in the Gulf of Guinea. The shells of many of the specimens were in poor condition and the valves had largely dissolved, pos- sibly a result of the wood being very acid. However, the characteristic siphonal plates readily identified the species. The two specimens taken from a coconut husk dredged off Sao Tomé in 2,550 m may have been advectitious; this species may not typically occur at that depth. Neither specimen carried young. The shelf in this area is very narrow and steep so that plant material could easily be carried into deep water. All other records are from depths of 239-366 m. Range. Known only from off the Florida Keys in the western Atlantic and the Gulf of Guinea in the eastern Atlantic in depths ranging from 239 to 2,550 m. Specimens Examined. Western Atlantic: UNITED STATES, FLORIDA: Gerda, station 266, off Fowey Rocks, about 16 miles SE of Miami (25°39’N 79°58'’W) in 340 m; Gerda, station 66, off Turtle Reef, about 13 miles SE of Fowey Rocks (25°25'N, Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 79°59'W) in 366 m; Gerda, station 220, about 30) miles S of Alligator Reef (24°25'N, 80°33.5'W) in 311 — m. Eastern Atlantic: GABON: Galathea, station 52, off Port Victoria, SAo Tomé Island (1°42’N, 7°51'E)! in 2,550 m (in coconut shell); Atlantique Sud, station. 147, about 45 miles N Port Gentil, Cape Lopez (0°S, | 8°58’'E) in 250 m; Atlantique Sud, station 154, about’ 35 miles NE of Port Gentil (0°15'S, 8°47’E) in 239 m. GENUS XYLOREDO TURNER 1972 Xyloredo Turner, 1972, Breviora, 397: 3 (type species, | Xyloredo nooi Turner, original designation); Turner, 1973, Science, 180: 1377-1379. | Distinctive Characters. Species in this. genus are characterized by having typical Xylophaga shells, which lack apophyses. and have a mesoplax composed of two flat plates, and by making a long teredolike burrow. The posterior two ane of the: burrow has a thin calcareous lining, marked with distinct growth lines and cov- ered with periostracum that extends to the | calcareous portion anterior as a band. The | part of the animal extending beyond the | valves into the calcareous tube is covered by a golden-brown periostracal sheath that | is continuous anteriorly with the covering of the valves and posteriorly with the per- iostracum of the tube. A fold of the mantle is attached to the growing end of the tube’ where both periostracum and calcium are added. In young specimens the tube may be composed entirely of periostracum. | Posterior to the valves, the combined in- current and excurrent pannel extend the. length of the tube and are attached lightly | to it at the base of the short separate si- | phons. Two dorsolateral ridges within the | incurrent canal appear to be ciliated and | possibly aid in water transport. Remarks. Members of this genus differs from Xylophaga and Xylopholas in making | a burrow that may reach more than 30. times the length of the shell and is lined with a calcareous tube. Several species of | Xylophaga make a burrow more than five times the length of the shells and form a | chimney composed of mucous- -cemented | fecal pellets at the posterior end of the burrows. These are not homologous with the caleareous tubes of Xyloredo but rath-_ er with the chimney of rock-boring pho- lads, as in Parapholas Conrad Carmen 1955: 123). The discovery of this teredolike genus in the Pholadidae requires a reexamination of the fossil teredinids, especially those re- corded as having ringed tubes. On the ba- sis of our present knowledge, it may be impossible to distinguish Xyloredo from teredinids in fossilized wood. However, if tubes are present, a microscopic analysis of their structure may aid in distinguishing between them because teredinid Hibee are amorphous, whereas tubes of Xyloredo have a definite structure with growth rings and periostracum. Certainly Xyloredo should be considered when examining drilled wood thought to have come from a deep water fossilized deposit. Although Xyloredo superficially resem- ble the Teredinidae, this is entirely con- vergent and does not in any way indicate relationship, nor does it suggest the evo- lution of the Teredinidae from the Xylo- phagainae. The latter lack apophyses and pallets, and have a mesoplax. In addition, none of the visceral mass or gills of the Xylophagainae extend beyond the valves posteriorly as they do in the Teredinidae. Range. To date three species of Xylo- redo are known, two in the western Atlan- tic and the other in the eastern Pacific. All occur at depths greater than 1,500 m. Xyloredo nooi Turner Plate 35 Xyloredo nooi Turner, 1972, Breviora, 397: 5-7, pls. 1, 2 (Tongue of the Ocean, about 4 miles off north- east tip of Andros Island, Bahama Islands [25°54'N, 77°49'W] in 1,737 m). Holotype, MCZ 279631; paratypes from the same and other panels exposed at the same locality, MCZ 279632, 279633, 279634, and 279635, and the Zoological Museum, Univer- sity of Copenhagen; Turner, 1973, Science, 180: 1377-1379. Distinctive Characters. Burrow long, teredinidlike, lined with a thin calcareous tube marked with growth rings and cov- ered with periostracum. Shell similar to Xylophaga, anterior slope narrow, umbo- nal—-ventral sulcus lightly impressed, pos- XYLOPHAGAINAE ° Turner 265 terior slope high and reflected dorsally. Posterior adductor muscle scar subellipti- cal, set high on the posterior slope, and divided into two distinct areas. Disc sep- arated from the posterior slope by a groove on the inner surface of the valves. Mesoplax small, the two flat triangular plates composed almost entirely of perios- tracum. Periostracal sheath between the valves and the tube smooth. Description. Shell globose, valves reach- ing 10 mm in length and 10.5 mm in height, thin, fragile; umbos prominent. Periostracum relatively thick, golden- brown, glistening, and covering entire valve. Pedal gape angle about 110°. Ante- rior slope sculptured with numerous den- ticulated ridges, there being 24 on the ho- lotype. Umbonal-ventral sulcus narrow, slightly impressed, and sculptured with fine, irregular growth lines. Posterior slope high, reflected near the dorsal margin, and sculptured with fine growth lines. Umbo- nal reflection rather wide, thin, adhering to the valves in the umbonal area, free an- teriorly. Inner surface of valves smooth and glis- tening. Umbonal—ventral ridge narrow, high, and segmented. Chondrophore and internal ligament prominent. Disc separat- ed from the posterior slope by a_pro- nounced narrow groove extending from the umbo to the posterior ventral margin. Posterior adductor muscle scar large, ellip- tical, and divided into two areas, the upper part marked with irregular impressions, the lower with regular chevron-shaped im- pressions. Anterior adductor scar covering most of the umbonal reflection. Pedal re- tractor scars elongate, the muscles insert- ing in the groove separating the disc from the posterior slope. Mesoplax small, flat, the two broadly tri- angular plates composed almost entirely of periostracum and located anterior to the umbos. Burrow long, teredinidlike, and lined with a thin calcareous tube that is sculp- tured with distinct growth rings and cov- ered with periostracum that extends as a 266 TABLE 21. MEASUREMENTS OF XYLOREDO NOOI. Length Height (mm) (mm) Location 5.0 Sy, IL paratype 6.7 6.9 paratype 7.2 7.8 paratype 9.5 9.8 paratype 9.5 10.0 holotype border at the anterior end. Tube in very young specimens composed entirely of periostracum. Between the valves and the anterior end of the tube the animal is cov- ered with a smooth periostracal sheath, which is continuous anteriorly with the covering of the valves and posteriorly with the tube. Siphons short, separate, and ap- parently with a few small cirri. Measurements. See Table 21. Remarks. Xyloredo nooi is related to both Xyloredo naceli Turner from the east- ern Pacific and X. ingolfia Turner from the North Atlantic. Xyloredo nooi differs from them in having a much thinner burrow lin- ing, a high, reflected posterior slope on the valves, and a proportionately smaller, more highly placed and divided posterior adduc- tor muscle scar. In addition, the periostra- cal sheath extending between the valves and the calearious tube is smooth. The larvae of X. nooi apparently do not rise very high in the water column, inas- much as a panel 25 ft off the bottom showed no trace of them, whereas the panel directly beneath it on the bottom was riddled. On the basis of the prodis- soconch still visible on some of the speci- mens, it would appear that the larvae are similar to those of Xylophaga. No evidence was found in the three riddled panels ex- amined that the young were brooded or held in the tubes. The burrows resemble those of teredi- nids and intertwine with each other but basically follow the grain of the wood. They may reach 200 mm in length and 15 mm in diameter at the anterior end. The largest tube with intact shells and animal remaining was 145 mm long; the valves Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 | were 9.5 mm in length and 10 mm in. height. The panels in the Tongue of the | Ocean were submerged for 34 months and | inasmuch as the larger burrows were emp- | ty or contained only fragments of shells, | the length of life may be about 2-2.5/ years. The calcareous lining of a burrow — 122 mm long extends to about 30 mm. from the anterior end, thus leaving room — for boring activities and changing the di- | rection of the burrow. Range. Known only from the type lo- | cality. Specimens Examined. BAHAMA ISLANDS: } Tongue of the Ocean, about 4 miles off NE tip of Andros Island (24°54'N, 77°49'W) in 1,737 m. Xyloredo ingolfia Turner Plates 36, 37 Xyloredo ingolfia Turner, 1972, Breviora, 397: 7-9, | pls. 3-5 (from wood dredged by the Ingolf Expe- dition at station 67, south of Eyrabakki, Iceland | [61°30'N, 22°30'W] in 975 fathoms [1,783 m]). Ho- — lotype, MCZ 279636; paratypes, MCZ 279637, and - the Zoological Museum, University of Copenha- | gen; Tumer, 1973 Science 180: 1377-1379. Description. Shell globose, valves reach- ing 2.5 mm in length and 2.0 mm in height, thin, fragile, with prominent um-_ bos; thin, glistening, almost colorless per- iostracum covering disc and posterior | slope. Beaked portion of anterior slope — wide, extending more than one half dis= | tance to ventral margin; sculptured with close-set and very finely denticulated ridg- | es. Posterior portion of anterior slope | about two thirds width of beak, sculptured with close-set ridges that extend to very | slightly impressed umbonal-ventral sulcus. Disc sculptured with well-marked growth | lines. Posterior slope small, low, and not | clearly demarcated on outer surface of valve. Umbonal reflection thick, narrow, short, and free except at posterior end. | Inner surface of valves smooth, slightly shiny to chalky (perhaps owing to long | preservation). Umbonal—ventral ridge wide, | flattened, often varying in width, irregularly | segmented, and not enlarged at ventral - condyle. Chondrophore and internal liga- TABLE 22. MEASUREMENTS OF XYLOREDO INGOLFIA. Length Height (mm) (mm) Location PAS) 2.3 holotype 2.5 2.0 paratype all 2.0 paratype 2.0 1.9 paratype 1.5 1.4 paratype 1.5 1.2 paratype ment large. Disc not clearly separated from posterior slope. Posterior adductor muscle scar large, slightly raised, elliptical, extend- ing nearly to ventral margin, with irregular, transverse impressions. Autener adductor muscle scar covering umbonal reflection. Siphonal retractor muscle scars not im- pressed. Pedal retractor muscle scar small, elongate to oval, and located just anterior to posterior adductor muscle scar. Meso- plax of two very small, narrow, subrectan- gular, flat, calcified plates lying on dorsal surface of anterior adductor muscle. Burrow 10-15 times length of valves; cal- careous tubular lining fives fourths length of burrow. Tube relatively heavy, fatinked with uniform, close-set, raised rings, and covered with light tan periostracum that ex- tends anteriorly as border. Portion of ani- mal between valves and tube covered by thin, irregularly ridged periostracal sheath. Siphons short; incurrent siphon slightly lon- ger than excurrent siphon. Protoconch large, medium golden-brown, and_ sculp- tured with fine, concentric ridges. Measurements. See Table 22 Remarks. This species differes from Xy- loredo naceli in having a less well-devel- oped posterior slope, a shallow, indistinct umbonal-ventral groove, a flattened um- bonal-ventral ridge, aml in having the valves longer than high. It differs from X. nooi in having valves longer than high, in having a isu rounded posterior slope, in lacking the distinct groove on the inner sur- face separating the valine from the posterior slope, and in having the plates of the me- soplax subrectangular and well calcified. XYLOPHAGAINAE ° Turner 267 Range. Known only from the type local- ity. Specimens Examined. ICELAND: Ingolf Expedi- tion, station 67, S of Eyrabakki (61°30'N, 22°30'W) in 1,783 m. Xyloredo naceli Turner Plate 38 Xyloredo naceli Turner, 1972, Breviora, 397: 9-11, pl. 6, figs. 1-5 (USNCEL STU I-4 about 30 miles S of San Miguel Island, off Port Hueneme, Santa Barbara Islands, California [33°46’N, 120°45’W] in 6,800 ft [2,072.6 m] from panels submerged from June odes to July 1965). Holotype, MCZ 279638; paratype, 279639. Distinctive Characters. Burrow teredi- nidlike, lined with a thin calcereous tube marked with growth rings and covered with periostracum. Shell Sailer to Xylo- phaga, with a narrow, slightly impressed umbonal—ventral sulcus. Posterior adduc- tor muscle scar elliptical, almost complete- ly covering the posterior slope and uni- formly marked with transverse impres- sions. Mesoplax small, the two flat, trian- gular plates composed largely of periostracum. Periostracal sheath covering the animal between the valves and the tube papillose. Description. Shell globose, reaching 1.5 mm in length, thin, fragile, white, aah al thin, pale yellow periostracum. Pedal gape angle about 100°. Anterior slope sculp- tured with 8-12 widely spaced, pro- nounced, denticulated ridges. Umbonal— ventral sulcus narrow and only slightly im- pressed. Discs and posterior slope sculp- tured with fine growth lines only. Inner surface of valves smooth and glis- tening. Umbonal—ventral ridge narrow, high, and indistinctly segmented. Chon- drophore and internal ligament well de- veloped. Posterior adductor muscle scar elliptical, extending from the dorsal nearly to the ventral margin of the posterior slope and regularly marked with transverse im- pressions. Pedal retractor scar not im- pressed. Ventral adductor only lightly im- pressed and located just posterior to the 268 TABLE 23. MEASUREMENTS OF XYLOREDO NACELI. Length Height (mm) (mm) Location 1.1 2 holotype 1.0 1.0 paratype 1.2 1.3 paratype 1.5 1.5 paratype umbonal-ventral ridge. Siphonal retractor muscles inserting on the tube. Mesoplax small, thin, set between and anterior to the umbos, the two triangular plates composed almost entirely of perios- tracum. Burrow of largest specimen about six times the length of the shell. Calcare- ous tube lining the burrow thin, distinctly marked with growth rings, and covered with periostracum that extends as a border anteriorly. Aperture of the burrow small, round, the white lining visible within. The portion of the animal between the valves and the calcareous tube covered with a pa- pillose periostracal sheath that is continous with the periostracum covering the valves anteriorly and the periostracal border of the tube posteriorly. Siphons short, of equal length, and apparently lacking cirri. Measurements. See Table 23. Remarks. This species is most closely re- lated to X. nooi Turner from the western Atlantic. Only eight specimens of X. naceli were found and they were all very small, but these appear to be sufficiently distinct to consider them members of a separate species. Xyloredo naceli differs from X. nooi in having a heavier tube that is cal- Ccareous, even in very young specimens; and a posterior adductor muscle scar that is proportionately larger and not divided into two areas. In addition, it lacks the pro- nounced internal groove separating the disc from the posterior slope, and the per- iostracal sheath covering the animal ante- rior to the tube is papillose. Nothing is known of the biology of the species except that at the sites where they were collected the temperature of the wa- ter was 2.1° C, the salinity was 34.52%o, the dissolved oxygen content was 1.26 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 ml/L, the pH was 7.84, and the hydrostatic pressure 3000 psi (Muraoka, 1966b). The embryonic valves still visible on some of the specimens suggest that the mature larvae are similar to those in Xy- lophaga. Xylophaga muraokai was the most common borer in the panels from which the X. naceli were taken. Range. Known only from the type lo- cality. Specimens Examined. UNITED STATES, CALI- FORNIA: USNCEL STU I-4 S of San Miguel Island, Santa Barbara Islands (33°46’N, 120°45’W) in 2,072 m. ACKNOWLEDGMENT This paper was prepared with the aid of funds received from the Department of the Navy, Biology Branch, Office of Naval Research, ONR grant NO0014-91-J-1402, Biological Studies on Marine Boring and Fouling Mollusks. LITERATURE CITED BARTSCH, P. 1921. A new classification of the ship- worms and descriptions of some new wood bor- ing mollusks. Proceedings of the Biological So- ciety of Washington, 34: 25-32. CULLINEY, J. L., AND R. D. TURNER. 1976. Larval development of the deep-water wood boring bi- valve Xylophaga atlantica Richards (Mollusca, Bivalvia, Pholadidae). Ophelia, 15(2): 149-161. DALL, W. H. 1886. Report on the Mollusca.—Part I. Brachiopoda and Pelecypoda. Reports on the re- sults of dredging—by the U.S. Coast Survey Steamer Blake. Bulletin of the Museum of Com- parative Zoology, 12(6): 171-318, 9 pls. . 1908. Reports on the dredging operations off | the west coast of Central America to the Gala- | pagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agas- siz, carried on by the U.S. Fish Commission steamer Albatross, during 1891, Lieut.-Dom- mander Z. L. Tanner, U.S.N., commanding. XXXVII. Reports on the scientific results of the expedition to the eastern tropical Pacific, in charge of Alexander Agassiz, by the U.S. Fish | Commission steamer Albatross, from October, | 1904, to March, 1905, Lieut.-Commander L. M. | Garrett, U.S.N., commanding. XIV. The Mollus- ca and the Brachiopoda. Bulletin of the Museum | of Comparative Zoology, 43(6): 205-487, pls. 1— _ 99 . 1909. Report on a collection of shells from | Peru, with a summary of the littoral marine Mol- lusk of the Peruvian zoological province. Pro- ceedings of the United States National Museum Bh OPA Ue DEPALMA, J. R. 1969. A study of deep ocean fouling, Straits of Florida and Tongue of the Ocean, 1961 to 1968. Informal Report IR 69-22. Washington, D.C.: U.S. Naval Oceanographic Office. 26 pp., 6 text-figs. Dons, C. 1929a. Zoologiske notiser. IV. Xylophaga praestans, ny for Norges fauna. Norske Viden- skabers Selskab Fordhandlinger, 1(57): 169-172, text-figs. 1-7. . 1929b. Zoologiske notiser. V. Xylophaga dor- salis i Norge. Norske Videnskabers Selskab For- handlinger, 1(65): 196-199, text-figs. 1-6. HOAGLAND, Kk. E. 1983. Morphological characters and their character states in the Pholadacea. Tryonia, 8: 1-51. HOAGLAND, K. E., AND R. D. TURNER. 1981. Evo- lution and adaptive radiation of the wood-boring bivalves (Pholadacea). Malacologia, 21(1-2): 111-148. KNUDSEN, J. 1961. The bathyal and abyssal Xylopha- ga (Pholadidae, Bibalvia). Galathea Report, 5: 163-209, text-figs. ]-41. MURAOKA, J. S. 1964. Deep-ocean biodeterioration of materials—part I. Four months at 5,640 feet. Technical Report R 329. Port Hueneme, Cali- fornia: U.S. Naval Civil Engineering Laboratory. 35 pp., 24 text-figs. . 1965. Deep-ocean biodeterioration of mate- rials—part II. Six months at 2,340 feet. Technical Report R 393. Port Hueneme, California: U.S. Naval Civil Engineering Laboratory. 42 pp., 5 text-figs. . 1966a. Deep-ocean biodeterioration of ma- terials—part III. Three years at 5,300 feet. Tech- nical Report R 428. Port Hueneme, California: U.S. Naval Civil Engineering Laboratory. 47 pp., 36 text-figs. . 1966b. Deep-ocean biodeterioration of ma- terials—part IV. One year at 6,800 feet. Techni- cal Report R 456. Port Hueneme, California: U.S. Naval Civil Engineering Laboratory. 45 pp., 31 text-figs. . 1966c. Deep-ocean biodeterioration of ma- terials—part V. Two years at 5,640 feet. Technical Report R 495. Port Hueneme, California: U.S. Naval Civil Engineering Laboratory. 46 pp., 36 text-figs. . 1967. Deep-ocean biodeterioration of mate- rials—part VI. One year at 2,370 feet. Technical XYLOPHAGAINAE ° Turner 269 Report R 525. Port Hueneme, California: U.S. Naval Civil Engineering Laboratory. 57 pp., 43 text-figs. PRASHAD, B. 1932. The Lamellibranchia of the Si- boga Expedition. Systematic part I. Pelecypoda (Exclusive of the Pectinidae). Vol. 53c. Siboga- Expeditie. Leiden: E. J. Brill. 353 pp. PURCHON, R. D. 1941. On the biology and relation- ships of the lamellibranch Xylophagadorsalis (Turton). Journal of the Marine Biological Asso- ciation of the United Kingdom, 25: 1—39, text- figs. 1-16. TAKI, I., AND T. HABE. 1945. Classification of Japa- nese Pholadacea. Japanese Journal of Malacolo- gy, 14: 108-123. . 1950. Xylophaginidae in Japan. No. 7, pp. 45-47. In T. Kuroda (ed.), Illustrated Catalogue of Japanese Shells. Vol. I, 1949-1953. Kyoto, Ja- an. tee. R. 1968. Ecological Aspects of Two Wood- Boring Molluscs from the Continental Terrace Off Oregon. Doctoral thesis. Corvalis, Oregon: Department of Oceanography, School of Sci- ence, Oregon State University. 137 pp., 50 text- figs. TURNER, R. D. 1954. The family Pholadidae in the western Atlantic and eastern Pacific. Part [— Pholadinae. Johnsonia, 3(33): 1-64, pls. 1-34. . 1955. The family Pholadidae in the western Atlantic and eastern Pacific part [I—Martesiinae, Jouannetiinae and Xylophagainae. Johnsonia, 3(34): 65-160, pls. 35-93. 1956. Notes on Xylophaga washingtona Bartsch and on the genus. Nautilus, 70: 10-12. . 1972. Xyloredo, a new teredinid-like abyssal wood-borer (Mollusca, Pholadidae, Xylophagai- nae). Breviora, 397: 1-19, pls. 1-6. . 1972. A new genus and species of deep water wood-boring bivalve (Mollusca, Pholadidae, Xy- lophagainae) Basteria, 36: 97-104, figs. 1-12. . 1973. Wood-boring bivalves, opportunistic species in the deep sea. Science, 180: 1377— 1379. TURNER, R. D., AND J. L. CULLINEY. 1971. Some anatomical and life history studies of wood-bor- ing bivalve systematics, pp. 65-66. In M. K. Ja- cobson (ed.), Annual Report for 1970. Seaford, New York: American Malacological Union. WATERBURY, J., C. B. CALLOWAY, AND R. D. TURN- ER. 1983. A cellulolytic nitrogen-fixing bacterium cultured from the gland of Deshayes in ship- worms (Bivalvia: Teredinidae). Science, 221: 1401-1403. Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 NS) aI i= Plate 1. Xylophaga concava Knudsen from Pillsbury, station 526. Figure 1. Dorsal view of apposed valves showing the concave posterior slope and erect mesoplax. Figure 2. Posterior end of | valve and siphons. Figure 3. Enlargement of the siphonal openings to show the six large cirri on the excurrent siphon and the | double row of small cirri on the incurrent siphon. XYLOPHAGAINAE ° Turner Daal Plate 2. Xylophaga gerda Turner n. sp. from Gerda, station 499. Figure 1. Lateral view of holotype showing the attachment of the posterior adductor muscle through the thin valve, the mesoplax that does not extend above the umbos, and the fecal cylinder in the excurrent canal. Figure 2. Dorsal view of the holotype showing the mesoplax. Figure 3. Enlargement of the posterior end of the siphons. Figure 4. Diagrammatic cross-section through the siphons and the fecal cylinder. Figure 5. Three-quarter view of holotype showing the inflated umbos and the simple curved plates of the mesoplax. Figure 6. A relatively smooth chimney composed largely of periostracum, with a thin coating of fecal material. Figure 7. A thick chimney, built in sections with “leaves” of periostracum extending at the anterior end of each section. i, Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 0.5mm Plate 3. Xylophaga gerda Turner n. sp. from Pillsbury, station 944. Figure 2. Inner view of left valve, showing posterior adductor muscle scar and low umbonal-— ! Figure 1. Outer view of right valve. ventral ridge. Figure 3. Enlargement of the hinge area of left valve to show the chondrophore and anterior adductor muscle — scar. Figure 4. Outer view of right valve of a young specimen. Figure 5. Inner view of left valve of young specimen showing | the prodissoconch, chondrophore, and muscle scars. i XYLOPHAGAINAE ° Turner 273 Plate 4. Xylophaga grevei Knudsen (Figs. 1-6 from Knudsen, 1961: 176-177). Figures 1-4. Holotype. Figure 1. Right side of entire specimen showing siphons and mesoplax. Figure 2. Dorsalview. Figure 3. Enlargement of the siphons showing the cirri around the incurrent aperture (35) and the excurrent aperture (6). Figure 4. Internal view of left valve of the holotype showing the broadly oval posterior adductor muscle scar set high on the posterior slope. Figures 5, 6. Ventral and dorsal view of the mesoplax. Figure 7. Small specimen carrying two young from Galathea, station 444. 274 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 5. Xylophaga clenchi Turner and Culliney. Figures 1—3. Side, dorsal, and anterior views of the holotype, showing the widely space ridges on the anterior slope of specimens | boring in soft wood and the position of young and the dorsal surface. Figures 4, 5. Inner and outer views of left valve of a. specimen from Atlantis I/, station 124, showing the muscle scar and large number of ridges on the anterior slope of a specimen — from hard wood. XYLOPHAGAINAE ° Turner 275 0.5mm Plate 6. Xylophaga clenchi Turner and Culliney. Figures 1, 2. Outer and inner views of the right valve of a large specimen with mesoplax in place, dredged by the Pillsbury, station 394. Figure 3. Lateral view of an entire specimen from the Tongue of the Ocean showing relative size of si- phons. Figure 4. Enlargement of the siphons, posterodorsal view, to show the incurrent and excurrent apertures and the cirri around them. 276 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 7. Xylophaga clenchi Turner and Culliney from /Ingo/f Expedition, station 67. Figure 1. Lateral view of right valve with mesoplax in place and young attached to dorsal surface. Figure 2. Dorsal view of| same specimen showing the prodissoconch. Figure 3. Lateral view of specimen with extended siphon. | ~l ~] XYLOPHAGAINAE ° Turner 2, Plate 8. Xylophaga clenchi Turner and Culliney. Figures 1, 2. From Pillsbury, station 238. Figure 1. Lateral view of right valve. Figure 2. Dorsal view of opposed valves showing prodissoconch, four young attached posterior to the umbos, and an atypical elongate, longitudinally folded mesoplax. Figures 3, 4. Specimen from Atlantis II, station 124. Figure 3. Anterior view of specimen with mesoplax bent at a right angle. Figure 4. Dorsal view of same specimen showing prodissoconch and mesoplax. 278 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 9. Xylophaga supplicata Taki and Habe from Tosa Bay, Shikoku, Japan. Figure 1. Anterior view of opposed valves with mesoplax in place. Figure 2. Lateral view of entire specimen from the right with mesoplax in place. Figure 3. Dorsal view of opposed valves tipped slightly forward to show the minute tubes at the posterior end of the mesoplax. Figure 4. Enlargement of mesoplax and umbonal area, looking down into the cavity formed by the incurving of the umbos. Figure 5. Inner view of left valve showing chondrophore with large tooth. Figure 6. Enlargement showing chondrophore with large tooth. XYLOPHAGAINAE ° Turner 279 Plate 10. Xylophaga whoi Turner n. sp. Figures 1-4. Specimen from Pillsbury, station 944. Figures 1, 2. Outer and inner views of the left valve. Figure 3. Frontal view of mesoplax with smallest tubes seen. Figure 4. Side view of left plate of mesoplax. Figures 5-12. Holotype. Figure 5. Outer view of right valve. Figure 6. Inner view of left valve. Figure 7. Dorsal view of left plate of mesoplax. Figure 8. Frontal view of mesoplax. Figures 9, 10. Outer surface of left and right plates of mesoplax with average-size tubes. Figures 11, 12. Inner surface of left and right plates of the mesoplax showing the curvature at the dorsal margin and the pore to the inner surface just below it. Figure 13. Enlargement of the hinge area of the left valve to show the chondrophore with a large tooth on it posterior dorsal margin, the umbonal reflection, and the inner surface of the mesoplax. 280 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 5mm Plate 11. Xylophaga whoi Turner n. sp. from Pillsbury, station 394. Figure 1. Anterior view of entire animal showing mesoplax in place, the anterior adductor muscle, the pedal opening of the mantle, the retracted foot, and young on the dorsal surface of the valves. Figure 2. Enlargement of the umbonal area, mesoplax, and attached young. XYLOPHAGAINAE ° Turner I8l1 Plate 12. Xylophaga profunda Turner n. sp. from U.S. Naval Oceanographic Office test site, Tongue of the Ocean, Andros Island, Bahama Islands. Figures 1-3. Holotype. Figure 1. Dorsal view showing round ridge posterior to the umbonal—ventral sulcus, the mesoplax, and attached young. Figure 2. Side view of left valve. Figure 3. Enlarged anterior view of dorsal area to show mesoplax. Figures 4, 5. Dorsal view of paratypes to show range of length—width relationships, concavity of posterior slope, as well as arrangement and number of young. Figure 6. Enlarged young. 282 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 2 5mm Plate 13. Xylophaga profunda Turner n. sp. from U.S. Naval Oceanographic Office test site Tongue of the Ocean, Andros Island, Bahama Islands. Figures 1, 2. Inner view of left valve of two specimens to show differences in intensity of muscle scar impressions, the riblike ridges extending from the umbonal-ventral ridge, the placement of the mesoplax, and the chondrophore. Figure 3. Enlargement of the hinge area of the left valve to show the chondrophore. Figures 4—6. Anterior, dorsal, and ventral view of mesoplax. XYLOPHAGAINAE ° Turner 283 Plate 14. Xylophaga abyssorum Dall. Figures 1-5. Specimens from U.S. Naval Oceanographic Office test site, Tongue of the Ocean, Andros Island, Bahama Is- lands. Figure 1. Outer view of right valve of immature specimen with mesoplax in place and showing ridge posterior to umbonal— ventral sulcus. Figure 2. Inner view of same valve, showing groove posterior to umbonal-ventral ridge, ligament, and charac- teristic muscle scar of young specimen. Figure 3. Dorsal view of same valve with mesoplax in place. Figures 4, 5. Mesoplax of young specimen with large ventral portion. Figure 6. Dorsal view of right valve of holotype of Xylophaga abyssorum for comparison with Figure 3. Figure 7. Dorsal view of young specimen from Gerda, station 266, with mesoplax developing lobes on dorsal portion. 284 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 ales 1mm Plate 15. Xylophaga abyssorum Dall. Figures 1-4. Specimens from Pillsbury, station 944. Figure 1. Inner view of right valve showing muscle scars, deep groove posterior to the umbonal-—ventral ridge, and the small chondrophore. Figure 2. Enlargement of hinge area of same speci- men. Figure 3. Hinge area of left valve. Figure 4. Dorsal view of umbonal area with mesoplax in place. This is the most elaborate mesoplax observed, having two tubes and a third developing. Figure 5. Dorsal view of specimen from Gerda, station 266, with two tubes formed on each plate of the mesoplax and with two young just posterior to the umbos. XYLOPHAGAINAE ° Turner 285 Plate 16. Xylophaga abyssorum Dall. Figures 1-5. Specimen from Pillsbury, station 944. Figure 1. Lateral view of right valve. Figure 2. Anterior view of opposed valves with mesoplax in place. Figure 3. Posterior view of opposed valves with mesoplax in place and showing the inflated umbos and ridge. Figure 4. Dorsal view of opposed valves. Figure 5. Enlarged view of mesoplax and umbonal area. 286 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 17. Xylophaga duplicata Knudsen from Galathea, station 745, Gulf of Panama (all from Knudsen, 1961: 175). Figure 1. Lateral view of left side of holotype, showing the mesoplax in place, standing off from the surface of the valves. Figure 2. Inner view of left valve of a paratype showing the small smooth posterior adductor muscle scar. Figure 3. Left side of | extended siphon of a paratype. Figure 4. Enlarged view of posterior end of the siphon. Figure 5. Dorsal view of the meso- plax. Figure 6. Ventral view of mesoplax. XYLOPHAGAINAE ° Turner 287 Plate 18. Xylophaga muraokai Turner n. sp. from U.S. Naval Civil Engineering Laboratory Test Site |. Figure 1. Lateral view of holotype showing siphons. Figure 2. Inner view of left valve showing muscle scar and simple chon- drophore. Figure 3. Dorsal view of the two plates of the mesoplax of a mature specimen showing large basal portion. Figure 4. Dorsal view of right plate of mesoplax. Figure 5. Ventral view of left plate of mesoplax. Figure 6. Lateral view of left plate of mesoplax. 288 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 1mm Plate 19. Xylophaga muraokai Turner n. sp. from U.S. Naval Civil Engineering Laboratory Test Site I. Figure 1. Anterior view of hinge area showing chondrophore and internal ligament. Figure 2. Posterior view of mesoplax fitting between the umbos. Figure 3. Dorsal view of young specimen with partially developed mesoplax. Figure 4. Mesoplax of young specimen. Figure 5. Enlargement of posterior end of siphons showing cirri around the siphonal apertures. XYLOPHAGAINAE ° Turner 289 3 1mm Plate 20. Xylophaga atlantica Richards from about 15 miles off Ipswich, Massachusetts, in 73 m. Figure 1. Entire specimen with siphons extended, showing relative lengths. Figure 2. Enlargement of the posterior end of the siphons, lateral view. Figure 3. Enlargement of the posterior end of siphons dorsal view showing the numerous small cirri surrounding the incurrent siphonal aperture and the larger, less numerous cirri of the excurrent siphon. 290 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 21. Xylophaga washingtona Bartsch. | Figures 1-3. Specimen from U.S. Naval Civil Engineering Laboratory Test Site |. Figure 1. Dorsal view of entire specimen: showing the relative length of the siphons. Figure 2. Dorsal view of siphons from just anterior to the truncation of the excurrent! siphon. Figure 3. Dorsal view of siphons in area of truncation to show contracted aperture of the excurrent siphon of a short! tube, and the short lateral lobes extending from the truncation. Figure 4. Specimen from about 40 miles W of Silver Point,| Oregon. Aperture of incurrent siphon showing cirri. | XYLOPHAGAINAE ° Turner 291 Plate 22. Xylophaga rikuzenica Taki and Habe from off Rikuzen, Honshu, Japan. Paratype, Museum of Comparative Zoology 194821. Figure 1. Inner view left valve showing posterior adductor muscle with herringbone markings. Figure 2. Outer view of left valve showing broad ventral sulcus. Figure 3. Anterior view of opposed valves showing mesoplax in place, the condyles, and chon- drophore. Figure 4. Dorsal view of opposed valves showing mesoplax in place, umbonal reflection, and broad, deep umbonal— ventral sulcus. Figures 5-7. Dorsal, lateral, and ventral views of the mesoplax showing the large ventral portion. bo bo 9 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 5mm 5mm Plate 23. Xylophaga depalmai Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida. Figure 1. Lateral view of holotype showing siphons with lateral periostracal sheath containing clusters of irregular glasslike granules. Figure 2. Dorsal view of holotype showing mesoplax that is only slightly coiled posteriorly. Figure 3. Dorsal view of specimen with slightly more coiled mesoplax. Figure 4. Siphons showing fringed lappets. Figure 5. Enlargement of fringe. Figure 6. Inner view of left valve showing muscle scars. Figure 7. Inner view of left valve showing prodisso- | conch. Figure 8. Enlargement of hinge area of left valve showing umbonal reflection and chondrophore. Figure 9. Enlargement - f of chondrophore. XYLOPHAGAINAE ° Turner 293 15 hear: 16 6 17 Plate 24. Xylophaga depalmai Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida. Figures 1-5. Specimen with the two plates of mesoplax fused. Figure 1. Outer view of right valve. Figure 2. Inner view of right valve showing lightly impressed muscle scars. Figure 3. Dorsal view of fused plates of mesoplax. Figure 4. Ventral view of mesoplax. Figure 5. Lateral view. Figures 6-8. Dorsal, ventral, and lateral view of mesoplax with plates partially fused ventrally. Figures 9, 10. Dorsal and ventral views of a mesoplax, which is broadened anteriorly and has a reduced ventral portion. Figures 11-13. Dorsal, ventral, and lateral views of an unusually broad mesoplax with unequal plates and reduced ventral portion. Figures 14, 15. Dorsal and ventral views of a broad mesoplax strongly coiled posteriorly, with a widened median area that separates the coils. Figures 16, 17. Dorsal and ventral views of a typical mesoplax with only the periostracal portion of the ventral portion fused. 294 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 25. Xylophaga guineensis Knudsen. Figures 1-3. Specimen from Pillsbury, station 260. Figure 1. Lateral view of entire animal showing foot, siphons, and perios- tracal sheath with fine whitish granules. Figure 2. Dorsal view showing siphons, prodissoconch, mesoplax, and loose perios- tracal sheath with granules. Figure 3. Anterior view showing foot, anterior adductor muscle, and mesoplax in place. Figures 4, 5. Specimen from Atlantique Sud, station 33. Figure 4. Inner view of right valve showing specimen with high flaring posterior slope, prodissoconch, posterior adductor muscle scar, and deep groove bounding the umbonal-ventral ridge. Figure 5. Outer view of right valve showing wide anterior slope and narrow, bladelike ridge posterior to the shallow umbonal-ventral sul- cus. Figures 6-9. Specimens from Atlantique Sud, station 146. Figure 6. Inner view of right valve of specimen with low, rounded posterior, small posterior adductor muscle scar set very high. Figure 7. Outer view of left valve with very closely set ridges on anterior slope. Figures 8, 9. Dorsal views to show variation in the mesoplax in place. XYLOPHAGAINAE ° Turner 295 2 Plate 26. Xylophaga guineensis Knudsen. Figure 1. Specimen from Atlantique Sud station 146. Umbonal area showing chondrophore, prodissoconch, and umbonal re- flection. Figures 2-4. Specimens from Atlantique Sud, station 33. Figure 2. Dorsal view of opposed plates of meso- plax. Figure 3. Ventral view of right plate of the mesoplax to show long cornucopialike shape. Figure 4. Dorsal view of right plate of mesoplax. 296 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 27. Xylophaga mexicana Dall. Figures 1, 2. Outer and inner views of left valve Figure 3. Lateral view of entire specimen showing inflated mesoplax and siphons. Figure Figures 5, 6. Outer and inner view of right valve of Figures 1-4. Specimens from “replication reef,” Santa Monica Bay, California. of specimen close to the holotype. 4. Dorsal view of entire specimen showing mesoplax and expanded foot. the holotype (from Turner, 1955, pl. 90). w Ke) ~l XYLOPHAGAINAE ° Turner INNS = SS Sreoes ——_ Plate 28. Xylophaga mexicana Dall from “replication reef,” Santa Monica Bay, California. Figure 1. Enlargement of the siphons to show the truncation of the excurrent siphon, the finely fringed lappets, granules em- bedded along the side of the siphon, and the papillose end of the incurrent siphon. Figures 2, 3. Ventral and dorsal view of mesoplax. Figures 4, 5. Inner and outer lateral views of mesoplax showing flange that fits down between the umbos. 98 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 EAN mameRe ere ee : % ia EEN Plate 29. Xylophaga tipperi Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida. Figure 1. Lateral view of holotype showing foot and extended siphons. Figure 2. Inner view of left valve to show smooth posterior adductor muscle scar. Figure 3. Dorsal view showing mesoplax in place. Figure 4. Enlargement of posterior end of incurrent siphon to show the fringed lappets and the single row of glasslike plaques along the side. Figure 5. Lateral and three-quarters view of glasslike plaques. Figures 6-9. Mesoplax. Figure 6. Dorsal view of right plate showing flat surface and faint sculp- ture. Figures 7, 8. Ventral view of right and left plates. Figure 9. Lateral view of right plate to show the compressed main portion and the posterior ventral flange that extends down between the umbos. XYLOPHAGAINAE ° Turner 299 Gi i ry igcs;) VAN WR eed < \ my \\ uy ( \\\ i j ( Wa ( ) A ae Ow Plate 30. Xylophaga bayeri Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida. Figure 1. Lateral view of holotype showing siphons and mesoplax in place. Figure 2. Dorsal view of holotype, showing the lateral extension of the mesoplax. Figure 3. Enlargement of the siphons showing the truncated excurrent siphon and the fringed lappets. Figure 4. Ventral view of the mesoplax. Figure 5. Dorsal view of the mesoplax. 300 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 31. Xylophaga bayeri Turner n. sp. Figures 1-3. Specimen from Gerda, station 266. Figure 1. Inner view of valves showing smooth muscle scar; narrow, high umbonal-ventral ridge, and broad, recurved umbonal reflection that adheres to the valve. Figure 2. Dorsal view of specimen showing broad umbonal reflection and the mesoplax. Figure 3. Enlargement of the mesoplax, dorsal view. Figures 4, 5. Specimen from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida. Figure 4. Lateral view of a young specimen with partially developed mesoplax. Figure 5. Dorsal view of a young specimen with partially developed mesoplax. XYLOPHAGAINAE ¢ Turner 301 Plate 32. Xylophaga japonica Taki and Habe. Figures 1-5. Paratype from Tosa Bay, Shikoku, Japan. Figure 1. Outer view of left valve showing thin ridge posterior to umbonal-ventral sulcus and elongate posterior slope. Figure 2. Dorsal view of mesoplax with distinct concentric sculp- ture. Figure 3. Ventral view of mesoplax showing flange that fits between umbos. Figure 4. Inner lateral view. Figure 5. Outer lateral view showing flange. Figures 6-11. Specimens from Anton Bruun, station 23. Figure 6. Outer view of left valve. Figure 7. Inner view of right valve showing chondrophore and smooth posterior adductor muscle scar. Figure 8. Outer view of mesoplax of a small specimen. Figure 9. Inner view of mesoplax of a small specimen. Figure 10. Outer lateral view of mesoplax of a small specimen. Figure 11. Inner lateral view of mesoplax of a small specimen. 302 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 SSE Yo Ki N LE pape ys \ Gs jist Plate 33. Xylopholas altenai Turner from Gerda, station 66. Figure 1. Lateral view of entire animal with the anterior adductor muscle relaxed so that the mesoplax is flattened and not visible, showing siphonal plates and young carried on ventral surface. Figure 2. Ventral view showing the mesoplax held in place by periostracum, the ventrally carried young, and the recurving of the siphonal plates. Figure 3. Dorsal view showing the mesoplax in place and the umbonal reflection. Figure 4. Enlargement of posterior end with the incurrent siphon projecting beyond the siphonal plate. Figure 5. Posterior end with left plate removed to show the muscle that extends into the cavity of the plate to which the siphonal retractor muscles attach. Figure 6. Lateral view of a very small specimen, contracted anteriorly so that the plates of the mesoplax are folded upward. Figure 7. Outer view of siphonal plate. Figure 8. Inner view of siphonal plate. Figures 1-3. Holotype. XYLOPHAGAINAE ¢ Turner 303 1mm . Plate 34. Xylopholas altenai Turner from Gerda, station 66. Figure 1. Outer view of right valve showing beaked portion of the anterior slope. Figure 2. Inner view of right valve showing the large posterior adductor muscle scar, pedal retractor scar, prodissoconch, and ligament. Figure 3. Inner view of upper part of left valve to showing the chondrophore and the mesoplax in its periostracal membrane. Figure 4. Dorsal view of the two plates of the mesoplax. 304. Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 3 | | | Plate 35. Xyloredo nooi Turner from U.S. Naval Oceanographic Office test site, Tongue of the Ocean, Bahama Islands. Figure 1. Inner view of left valve showing divided muscle scar. Figure 2. Outer view of left valve. Figure 3. Outer view of left! valve of holotype showing the flaring posterior dorsal margin. Figure 4. Inner view of right valve of holotype showing the divided posterior adductor muscle scar, the deep groove separating the disc from the posterior slope, and the umbonal reflection. Figure 5. Anterior view of opposed valves, showing the chondrophore and internal ligament. Figure 6. Dorsal view showing the thin | mesoplax. XYLOPHAGAINAE ° Turner 305 1mm Plate 36. Xyloredo ingolfia Turner from Ingolf Expedition, station 67. Figure 1. Lateral view of an entire animal, showing the periostracal sheath, the extended anal and siphonal canals, and the short siphons. Figure 2. Dorsal view of an entire animal, showing the periostracal sheath, the extended anal and siphonal canals, and the short siphons. Figure 3. Lateral view of a very young specimen showing the large prodissoconch and the produced beaked portion of the anterior slope. Figure 4. Lateral view of left valve with periostracal sheath attached. Figure 5. Dorsal view showing the produced, recurved beaks, the umbonal reflection, and the mesoplax with only the central portion of each plate calcified. Figure 6. Inner view of right valve with well-marked posterior adductor muscle scar. Figure 7. Inner view of left valve with lightly marked scar and large prodissoconch. 306 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 Plate 37. Xyloredo ingolfia Turner from Ingolf Expedition, station 67. | Figure 1. Inner view of left valve showing large prodissoconch, strong umbonal-—ventral ridge, reduced posterior slope, and lightly! impressed posterior adductor muscle scar. Figure 2. Outer view of left valve showing the produced beak and rounded low, posterior slope. XYLOPHAGAINAE ° Turner 307 Plate 38. Xyloredo naceli Turner from U.S. Naval Civil Engineering Laboratory Test Site | (submersible test unit I-4). Figure 1. Lateral view of holotype. Figure 2. Lateral view of specimen partially dissected from the wood showing the calcereous tube with the anterior periostracal margin and the papillose periostracal sheath covering the animal between the tube and the Figure 3. Anterior-lateral view of entire specimen showing foot and mesoplax. Figure 4. Inner view of left valve showing valves. Figure 5. Outer view of left valve. muscle scars and chondrophore. 308 INDEX Taxon names are printed in italics. Page numbers in bold refer to illustrations; those in italics refer t species descriptions; t indicates table. abyssorum, Xylophaga, 4, 12, 22, 23-24, 61-63 africana, Xylophaga, 4, 17-18 altenai, Xylopholas, 41-42, 80-81 atlantica, Xylophaga, 4, 27-28, 67 aurita, Xylophaga, 5, 31 bayeri n. sp., Xylophaga, 5, 12, 24, 38, 38-39, 77-78 bruuni, Xylophaga, 4, 19, 20, 21 clenchi, Xylophaga, 4, 13, 14, 16, 17-19, 28, 52-55 concava, Xylophaga, 4, 14-15, 15, 16, 26, 48 depalmai n. sp., Xylophaga, 5, 31-34, 35, 38, 40, 70- 71 variation of mesoplax in, 11-12, 32 distribution of Xylophagainae, 2 of Xylophaga species, 2, 6 (map) dorsalis, Xylophaga, 5, 12, 38, 39 dorsal plates. See mesoplax duplicata, Xylophaga, 4, 25, 26, 36, 64 erecta, Xylophaga, 4, 14 foliata, Xylophaga, 4, 19 food chain, 14 galatheae, Xylophaga, 4, 22 gerda n. sp., Xylophaga, 4, 14, 15-16, 49-50 globosa, Xylophaga, 5, 9, 36, 39 grevei, Xylophaga, 4, 16-17, 51 growth series, 12 guineensis, Xylophaga, 5, 32, 34-35, 40, 72-73 hadalis, Xylophaga, 4 indica, Xylophaga, 5, 40 ingolfia, Xyloredo, 44, 44-45, 84 japonica, Xylophaga, 5, 24, 37, 39-40, 79 Lignopholas, 3 lobata, Xylophaga, 4, 22, 24 Martesia, 3 mesoplax as character used in grouping, 3—5 variation in, 11-12 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 mexicana, Xylophaga, 5, 32, 35-37, 38, 40, 74-75 muraokai n. sp., Xylophaga, 4, 12, 25-27, 46, 65-66 murrayi, Xylophaga, 4, 18 naceli, Xyloredo, 44, 45, 45-46, 85 | nooi, Xyloredo, 13, 43-44, 46, 82 | obtusata, Xylophaga, 4, 20, 21 panamensis, Xylophaga, 4,18 Pholadidea, 40 | pholadids, 3, 41 | praestans, Xylophaga, By, Ie, Bul profunda, Xylophaga, 13, 14, 24 | profunda n. sp., Xylophaga, 13, 14, 21-23, 24, 59-60 | reproduction, 3 rikuzenica, Xylophaga, 5, 30-31, 69 siphons, 3-5 | supplicata, Xylophaga, 4, 18, 19-20, 20, 21, 22,56 Teredinidae, 41, 43 | teredinids, 2, 3, 43 teremachi, Xylophaga, 5 tipperi n. sp., Xylophaga, 5, 32, 37, 37-38, 76 tomlini, Xylophaga, 5 tubulata, Xylophaga, 4, 20, 21 turnerae, Xylophaga, 5 variation due to substrate, 6, 7t, 7-10, 10-11 in growth series, 12 in mesoplax, 11-12 washingtona, Xylophaga, 5, 27, 28, 28-30, 31, 68 variation due to substrate in, 6, 7t, 7-10, 10-11 whoi n. sp., Xylophaga, 4, 20-21, 22, 57-58 wolffi, Xylophaga, 4, 16, 18 Xylophaga, distribution of, 2, 6 groups in, 3-5 nomenclature of parts of, 4 Xylopholas, 2, 40-41, 42 Xyloredo Turner, 2, 41, 42-43 = aa = . = . y } R ull etin oF ae THE BOLAS SPIDERS OF THE GENUS MASTOPHORA ARANEAE: ARANEIDAE) HERBERT W. LEVI HERBERT W. LEVI HARVARD UNIVERSITY VOLUME 157, NUMBER 5 CAMBRIDGE, MASSACHUSETTS, U.S.A. 5 February 2003 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952— BULLETIN 1863— Memotrrs 1865-1938 Jounsonia, Department of Mollusks, 1941-1974 OcCASIONAL PAPERS ON Mo.Luusks, 1945— SPECIAL 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, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4. Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology 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 of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the 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 of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1947. (Complete sets only.) Proceedings of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2003. Plate 1. Upper left, Mastophora diablo new species (Argentina, photo J. Abalos). Upper right, M. stowei new species (Kentucky, photo K. F. Haynes and K. V. Yeargan). Middle left, MM. yeargani new species (Kentucky, photo K. F. Haynes and K. V. Yeargan). Lower left, M. alvareztoroi |barra and Jiménez new species (Texas, photo M. Stowe). Lower right, M. fasciata Reimoser (Costa Rica, photo W. Eberhard). THE BOLAS SPIDERS OF THE GENUS MASTOPHORA (ARANEAE: ARANEIDAE) HERBERT W. LEVI" ABSTRACT. Of 48 species of bolas spiders (Masto- phora) found in the Americas, 22 are new. Of that number, nine new species and six previously known species are North American. The North American Mastophora bisaccata is a group of species of similar appearance. We can expect additional finds of new species of these rare and specialized spiders. The spe- cies range from New Hampshire in the United States to central Argentina. No species are known from tropical Amazon or the northwestern states of the United States. The greatest abundance of species is in warm temperate areas of southeastern North America and southern Brazil and northern Argentina. Agatostichus is synonymized with Mastophora. With- in this paper, Ibarra and Jiménez describe a new spe- ‘cies from Chiapas and Texas. Evidence from palpal morphology indicates a relationship of Kaira with Taczanowskia and Mastophora, suggesting that insect attractants may have evolved only once. Epeiroides fasciolata, erroneously placed in Mas- tophora, is a Kaira, the male of Kaira altiventer. The related Asian genus Euglyptila is synonymized with Ordgarius. INTRODUCTION Female bolas spiders spend the day resting on leaves and branches, usually mimicking bird droppings and sometimes berries, snails, or leaf buds. At night, the bolas spider feeds on male moths attracted by the spider’s scent; the scent mimics the sex attractant of the female moth. This pheromone was first suggested by Hutch- inson (1903). Evidence of this pheromone was found by Eberhard (1977), and chem- ical analyses were conducted by Stowe et al. (1987) and Gemeno et al. (2000). An approaching male moth is caught with a silken thread bearing a viscid drop, the bo- ‘Museum of Comparative Zoology, Harvard Uni- versity, 26 Oxford Street, Cambridge, Massachusetts 02138-2902. Bull. Mus. Comp. las, hurled at the moth. Moths stick to the bolas, whereas detachable wing scales per- mit moths to escape from most orb webs. The unusual behavior of Mastophora, first observed by Hutchinson (1903), has at- tracted the attention of researchers, in- cluding taxonomists (Eberhard, 1981; Stowe, 1986; Stowe et al., 1987; Yeargan, 1988, 1994, 1997). Unlike most American orb weavers, the genus Mastophora has been revised in the past. That is, the de- scribed species were compared and illus- trated and keys were made. Mello-Leitao (1931) first reviewed all known species, and in the same year Canals (1931) revised all Argentinean species. Both authors re- lied on differences in the shape of the horns on the cephalothorax. The presence of horns is a character of the genus and is difficult to use for differentiating species. Genitalia were not illustrated until Gertsch (1955) revised the North American spe- cies. Unfortunately, Gertsch’s illustrations were poorly labeled. Those with legends indicating an internal view of the epigyn- um actually were the cleared posterior of the epigynum. Gertsch correctly reported that the diversity of egg sacs from Florida suggested overlooked species. Most of our knowledge of the biology of these spiders comes from a few species, mostly from North America. All late-instar and adult female bolas spiders spin a hor- izontal line composed of multiple threads, and then attach a bolas to it. The bolas consists of one, rarely several, balls of sticky glue drops on a line. A moth attract- ed by the spider's scent is caught by a swing of the leg holding the bolas; the ad- Zool., 157(5): 309-382, February, 2003 309 310 hesive is strong enough to hold moths. The moth is wrapped and usually the spider builds a new bolas and continues hunting before eating. After approximately 20 min- utes or more of hunting, when the bolas has not been used, the spider pulls it back and ingests the silk and glue and spins a new bolas. The bolas of spiders in the genus Mas- tophora is held with the first leg and swung at prey. Members of the Australa- sian genus Ordgarius and the African ge- nus Cladomelea use the second leg and whirl the bolas (Stowe, 1986; Yeargan, 1994; Leroy et-al., 1998). Only male moths are attracted. Stowe et al. (1987) showed that M. cornigera pro- duces several of the pheromone compo- nents produced by females of the moth prey species. Gemeno et al. (2000) showed that M. phrynosoma produces prey pher- omone components in proportions that represent an attractive blend. Attractants are released only while hunting (Gemeno et al., 2000). In one study, M. dizzydeani captured 2.2 moths per night, although a moth approached the spider as it hunted about once every 6 minutes (Eberhard, 1981). Different species of moths that mate at different times and that produce entirely different pheromones are caught at different times during the night. Lists of moth species captured: were reported by Stowe (1988), Yeargan (1994), and Stowe et al. (1995). Early instars of Mastophora of both sex- es and the minute adult males rest on the edges of leaves and feed mostly on male meriateccrn flies, primarily Psychodidae (moth midges), oven also are attracted by scent (Yeargan and Quate, 1996, 1997). The flies are captured with the first two pairs of legs, without the use of silk. The legs are armed with rows of strong setae (Figs. 5, 6). The spiderlings do not feed on each other (Stowe, 1986; Yeargan, 1994). In later instars, females lose these bristles and start to use a bolas. North American species have only 150— 250 eggs in each of one to five brown egg Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 sacs (Figs. 445-465). Each egg sac is the size of the female. Mastophora cornigera makes more egg sacs. Female Mastophora in the northeastern states die in autumn. More eggs have been reported from other species. Mastophora extraordinaria was found to produce 530 eggs (Bréthes, 1909) and M. dizzydeani produced 826 eggs (Eberhard, 1981). Oviposition of M. hutchinsoni (a North American species) takes place in fall and spiderlings emerge in May (Yeargan, 1988). This differs for M. cornigera. Glitch sizes for M. hutchinsoni ranged from 178 to 275. The sex ratio ap- proached one to one (an unusual excep- tion was observed in one egg case of M. phrynosoma, see below). Males and fe- males are similar in size at hatching but females grow to be much larger than males. Males mature in June, at about 1.7 mm total length, two months before fe- males become mature. The scent may come from the integu- ment (Lopez, 1998). The horns of the car- apace of Mastophora contain midgut di- verticula (Lopez et al., 1985). The silk glands were described by Lopez and Stowe (1985). The females rest in exposed places dur- ing the day with legs drawn in, often on a small pad of silk. All species are cryptic and uncommon, and difficult to find when present. The clustered egg sacs are sus- pended by strong threads on branches, and are noticed more often than the spi- ders, particularly in deciduous forests after leaves have fallen. When a spider is picked | up, it rolls in the hand rather than holding on, and when first disturbed may regur- gitate fluid that has a pungent odor (Ebem haved 1981). A summary of research was_ reported by Yeargan (1994). METHODS AND ACKNOWLEDGMENTS The collections of the following institu- tions and individuals were used. American Museum of Natural] History, New York (N. Platnick, | L. Sorkin) AMNH BMNH ECOTAR FCMU FMLT FSCA BSP INHS IRSNP MCN Natural History Museum, Lon- don, United Kingdom (P. Hill- yard, J. Margerison) California Academy of Scienc- es, San Francisco, California (C. Griswold, D. Ubick) Canadian National Collections, Ottawa, Canada (C. Dondale) Clemson University Arthropod Collection, Clemson, South Carolina (J. Moore, D. Carna- gey) Cornell University collection, kept in AMNH (N. Platnick, L. Sorkin) Denver Museum of Nature and Science, Denver, Colorado (P. Cushing) D. Ubick, San Francisco, Cali- fornia El Colegio de la Frontera Sur, Tapachula, Chiapas, Mexico (G. Ibarra) Facultad de Ciencias, Seccion Entomologia, | Montevideo, Uruguay (M. Simo) Fundacion Miguel Lillo, Tu- cuman, Argentina (S. Z. Turk, le v5 JE Jee rexchti)) Florida State Collection of Ar- thropods, Gainesville, Florida (G. B. Edwards) Instituto Butantan, Sao Paulo, Brazil (A. Brescovit) Illinois Natural History Survey, Urbana, Illinois (C. Favret) Institut Royal des Sciences Na- turelles de Belgique, Brussels, Belgium (L. Baert) J. Beatty, Carbondale, Illinois J. Kaspar, Oshkosh, Wisconsin J. Murphy, London, United Kingdom K. V. Yeargan, Lexington, Ken- tucky Museo Argentino de Ciencias Naturales, Buenos Aires, Ar- gentina (M. E. Galiano, C. L. Scioscia) Museu de Ciéncias Naturais, MCP MCZ MKS ML] MLP MNHN MNRJ MUSM MZAQ MZSP NHMW NMB NMP OSU QMB TAMU MASTOPHORA ® Levi 311 Fundagao Zoobotanica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil (E. H. Buckup, M. A. L. Marques) Museu de Ciéncias, Pontificia Universidade Catélica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil (A. A. Lise) Museum of Comparative Zo- ology, Cambridge, Massachu- setts M. K. Stowe, Gainesville, Flor- ida M. L. Jiménez, La Paz, Mexico Museo de Universidad Nacion- al, La Plata, Argentina (C. Ituarte, L. A. Pereira) Muséum National d’ Histoire Naturelle, Paris, France (C. Rollard) Museu Nacional, Rio de Janei- ro, Brazil (A. B. Kury) Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru (Diana Silva D.) Museu, Departamento de Zoologia da Escola Superior de Agricultura “Luis de Queiroz,” Piraciba, SA0 Paulo State, Bra- zil (G. J. de Moraes) Museu de Zoologia, Universi- dade de Sao Paulo, Sao Paulo, Sao Paulo, Brazil (E. M. Can- cello, R. Pinto da Rocha) Naturhistorisches Museum, Vi- enna, Austria (J. Gruber) Naturhistorisches Musuem, Basel, Switzerland (A. Hinggi) Natal Museum, Pietermaritz- burg, South Africa (D. A. Bar- raclough, C. Conway) Ohio State University, Marion, Ohio (R. A. Bradley) Queensland Museum, Bris- bane, Queensland, Australia (R. J. Raven) Texas A&M University, College 312 Station, Texas (A. Dean, E. Ril- ey) National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C. (J. Coddington, L. Lopardo) Zoological Museum of the Uni- versity of Copenhagen, Copen- hagen, Denmark (N. Scharff) USNM ZMUC Brazilian colleagues Erica H. Buckup and Arno Lise encouraged me to revise Mastophora. The revision was made pos- sible with the help of numerous South American colleagues with loan of collec- tions, including many old specimens, and advice. Especially helpful were C. Scioscia and the late M. Galiano of Buenos Aires, and the Brazilians E. H. Buckup, A. Bres- covit, R. Pinto da Rocha, G. J. de Moraes, and A. B. Kury. I also thank G. S. Oxford, K. V. Yeargan, M. K. Stowe, W. Eberhard, J. Leroy, and A. Leroy for help and advice and Eric J. Olson for translating difficult Spanish. L. Leibensperger was helpful in the laboratory. L. R. Levi and the editor, B. L. Clauson, polished the writing. W. Eberhard read the introduction and im- proved the wording. K. Yeargan and M. K. Stowe read the manuscript and suggested many improvements and corrections. Much appreciated are the comments of two anonymous readers of the manuscript. Gifts of Mastophora to the MCZ collec- tions were received from C. Hieber, G. Ibarra (from Chiapas, Mexico), J. M. Maes (from Nicaragua), M. K. Stowe, and K. V. Yeargan. G. Thue N. and M. L. Jiménez sent specimens of a new species with com- plete descriptions and finished illustra- tions. Mastophora alvareztoroi is de- scribed here with the names of Ibarra and Jimenez as authors. The project was started with the help of National Science Fooundation grant BMS 75-05719. The Wetmor ev Golles! Fund cov- ered publication costs in part. For examination of spiders, I followed the procedures described by Levi (1993a). For examination and illustration of the Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 small and difficult genitalia of Mastophora, additional methods were employed. After illustrating ventral and posterior views of the female, the epigynum was cut off from the body and the soft tissues were carefully removed (Fig. 2). The epigynum was placed on the dry end of a dish with white paraffin, and extra alcohol was re- moved with a bit of tissue. The epigynum was then placed in a conical pit in the par- affin (Fig. 3) containing two drops of Hoy- ers medium.? Placement was so that the anterior end faced the bottom; the flat posterior was parallel with the surface (Fig. 3). The viscous nature of Hoyer’s me- dium makes it an ideal clearing medium and keeps the epigynum in position, al- though presumably other clearing agents, such as oil of cloves or glycerol, could be used. The genitalia were carefully exam- ined and “ilies ated from at least one spec- imen of each species; several epigyna were cleared for abundant and variable species. Male spiders for which determination: was known were placed in a drop of tinc- ture of iodine and left overnight before ex- amining them the next morning in 80% alcohol. Tincture of iodine gives a good! stain, which may not be permanent, and it is easy to obtain in pharmacies. Tincture of iodine is isotonic with alcohol and does not expand the palpus or warp sclerites. — Gertsch (1955) in his revision did not’ interpret the two original descriptions by. Banks (1898) of Ordgarius obesus and O. corpulenta. The holotypes were placed in the California Academy of Science and were destroyed in the 1906 earthquake. The specimens, with others, were given to Banks from the Marx Collections after George Marx died. Banks (1898) pointeds out Le the time: > Hoyer’s medium is made by dissolving 15 g of gum arabic (clear flakes) in 25 ml of distilled water | at room temperature. Seventy-five grams of chloral hydrate is added, and the mixture is allowed to stand | for 1-2 days, until all solids have dissolved. Five mil-| liliters of glycerol is then added and the mixture is! filtered through glass wool and stored in a glass-stop-| pered bottle. Anyone familiar with Dr. Marx’s methods of work will not be surprised to learn that many of the spec- imens, when sent to me, bore no locality label whatever. Doubtless he knew where they came from, but left no clew [sic] that others might use. Some of the species were numbered, and by ex- amining several of his series of numbers it was pos- sible to find localities and his name for the species. However, Banks published illustrations of the two species that can be identified if one disregards the localities stated. DISCUSSION Determination of members of the genus Mastophora is the most difficult of araneid spiders. By using morphological charac- ters, 48 presumed species were separated, of which 22 species are new. Eleven of these new species are known from a single female. However, additional specimens were found of species earlier described from only a single specimen. All females are between 7 and 17 mm and most are approximately 12 mm total length. Most of the species that lack humps on the abdomen seem to be found in North America. A few species of Mas- tophora are slightly smaller and have dif- ferent Hibercles or horns on the carapace; these were placed in Agatostichus in the past, but males and females do not differ in other characters from Mastophora. Adult males of all species are approxi- mately 1.7 mm total length. Males of re- lated genera in Asia, Australia, and Africa are the same length. Males of the same species as females with abdominal humps coming from different egg sacs may have humps or not; females without humps may have males with or without humps. The males can be matched to females only by raising them from an egg sac from a de- termined female. Some males apparently emerge from the egg sac as mature adults (M. cornigera and perhaps M. gasteracan- thoides), But most take at leastn two instars to mature. The males of only a few species are known, because only ‘W. Eberhard, Mark Kk. Stowe, and Ken V. Yeargan have raised individuals. The males of most southern South American species remain MASTOPHORA ° Levi 3138 unknown. The palpi of different species are surprisingly similar and the determi- nation of males presents a challenge. Immatures coming out of the egg sac have visible median’ Raiborcles and horns on the carapace and are approxinately 1.2— 1.6 mm total length. They may have ab- dominal humps. Some egg sacs can be determined. Those of M. hutchinsoni are unique and are attached by their base (Figs. 453, 454), whereas all others are hanging. Some have a thick stalk (M. bisaccata; Fig. 446), extra long flaps (M. phrynosoma; Figs. 449, 450), or may lack flaps altogether (M. cor- nigera; Fig. 455). However, not enough detenumed egg sacs were available to make a key. The epigyna of females are much re- duced. Females are separated by their col- oration and shape of the abdomen, as well as by their genitalia. However, with so few specimens available of most species, gen- eralizations on appearance are a guess. The most common North American spe- cies, M. bisaccata, was found to be a group of species with similar abdominal mark- ings. Gertsch (1955) missed this because he relied on coloration and did not care- fully examine the genitalia of all speci- mens. Members of the M. bisaccata spe- cies group also differ slightly in average size, with M. bisaccata being the largest. Gonsch (1955) and Yeargan (1994) sus- pected M. bisaccata to be a group of spe- cies. The epigynum lacks a scape and has no ventral features permitting the palpus to be held in place (Figs. 12, 19, 26). A pos- terior edge (Figs. 12, 19, 26) may be pre- sent. Whatever diagnostic features exist are on the posterior face, which has two slits, a plate between, and a plate to the side (Figs. 13, 20, 27). The plates are usu- ally weakly sclerotized. The slits lead into a ventral, or sometimes dorsal, atrium (Fig. 4), which empties into the seminal receptacles (Fig. 4). The slits vary in di- rection in different species (Figs. 13, 27). The slits may be in depressions or may 314 have unique sculpturing along the edges, or a lip (Fig. 20). I considered these to be useful characters for determination of spe- cies. The palpi of males are lightly sclero- tized. A median apophysis, a radix, the em- bolus, and a terminal apophysis holding the embolus are present (Fig. 7). The con- ductor, a structure that arises from the te- gulum facing the embolus, is absent, al- though it seems to be present in Clado- melea species (Fig. 443). The different species of Ordgarius and Cladomelea show greater morphological differences from each other than do those of Mastophora. Different American species show some variation in genitalia and morphology in different specimens. Perhaps this is asso- ciated with the rarity of individuals and wide distribution of species. All books on venoms list M. gasteracan- thoides among the venomous American spiders ( Sedna 2000). These citations all come from Escomel (1918), who de- scribed the venom from specimens found in grapes in southern Peru. Farmers work- ing in grapes were bitten on the hands and legs. He considered the bites to be “cu- tanéo-hémolytique gangrénieux” and gave detailed description of the signs, which were necrotic skin lesions at the site of the bite. B. A. Houssay and J. J. Carbonnel checked the specimens at the time. In bor- rowed material, Escomel’s specimens were found in both the Buenos Aires and in the Paris museums. Examination showed them to be a new species of Mastophora (M. es- comeli), close to M. gasteracanthoides. No other evidence of human envenomation appeared with any labels on specimens of other species examined. Gertsch (1955) believed that the responsibility for the bites should be awarded to some other spi- der or arthropod. I suspect that M. esco- meli, unlike others, is more aggressive and more readily bites human skin than other species. No recent reports exist of ven- omous bites from M. escomeli. Mastophora escomeli may have been abundant in 1917 and then become rare again. Escomel Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 (1918) also found that extracts from eggs, injected into guinea pigs, were toxic. Relationship. Mastophora Holmberg, 1876, shares carapace outgrowths with Australasian Ordgarius Keyserling, 1886 (Figs. 422-433), and African Cladomelea Simon, 1895 (Figs. 434-444). Both Ord- garius and Cladomelea species handle the Boles with the second leg and, unlike Mas- tophora, which swings the bole in a pen- dulumlike motion, they whirl the bolas, Ordgarius when a moth approaches, ane Gindomeion for 15 minutes at a time when hunting, at approximately 150 rotations per minute (Leroy et al., 1998). The carapace outgrowths also are shared with the African Acantharachne Tullgren, 1910,> Madagascan Coelossia Si- mon, 1895, Madagascan Exechocentrus Si- mon, 1889, and immatures of Euglyptila Simon, 1908, from northern Vietnam (Tonkin). Euel yptila is synonymized below with O Ordgarius. The males of these and their habits are not known, but females were illustrated and described by Emerit (1980, 2000). The genus Agatostichus Si- mon, 1895, is synonymized below with — Mastophora, and Dicrostichus Simon, 1895, has been synonymized with Ordgar- ius by Davies (1988). The Mastophora, — Ordgarius, Dicrostichus, Cladomelea group is absent from European, Mediter- _ ranean, and central Asian faunas. When males are found, African Acan- tharachne and Coelossia should probably be synonymized with Cladomelea or Ord- garius. However, Exechocentrus differs in | having a long eye projection, a long me- dian tubercle, and a pair of long, posterior tubercles on the carapace (Emerit, 1978, 1980, 2000). Eberhard (1981), Stowe (1986), and | Yeargan (1994) studied the relationships with other genera. Scharff and Coddington ° Roewer (1942) and Platnick (2001) cited the ge- nus under the name Acantharanea Strand, 1929. The — name Acantharachne Tullgren, 1910, is not preoc- | cupied, as thought by Stand (Neave, 1939a: 9: Bon- | net, 1955: 124). constructed a cladogram (1998). The re- sults of Eberhard (1981), Stowe (1986), and Yeargan (1994) are summarized in Ta- ble 1; examination of these results showed a close relationship of Mastophora to Tac- zanowskia Keyserling, 1880. Members of the genus Mastophora seem related to genera lacking carapace tubercles (Tacza- nowskia, Celaenia, Kaira, Cyrtarachne, Poecilopachys, and Pasilobus; Table 1). Robinson and Robinson (1975) first sug- gested that the web of Pasilobus was in- termediate between orb webs and bolas. My own studies of genitalia of Taczanows- kia (Levi, 1997) showed that Taczanowskia is related to Celaenia and Kaira. The distal pocket of the epigynal scape of Tacza- nowskia correlates with the large hook on the median apophysis of the palpus of the male (Levi, 1997, fig. 19). Unequal claw lengths and armed femora, which are syn- apomorphies, relate Taczanowskia and Ce- laenia. The denticles next to a tooth on the side of the median apophysis of Tacza- nowskia also are found in Kaira species and Metepeira. Such median apophysis denticles are unique to several genera and I consider such a row of denticles as a syn- apomorphy of Kaira, Metepeira, and Tac- zanowskia. One of the synapomorphies of most genera allied to Araneus is a spine or tooth on the median apophysis. In con- trast, males that have a paramedian apoph- ysis in the palpus rarely have a tooth or spine on the median apophysis (e.g., Al- paida, Eriophora, Ocrepeira, Acacesia, Cyrtophora, and many others). The shape of the median apophysis of the palpus of Pasilobus also probably is derived from a median apophysis similar to that of Kaira. The presence of a spine on the median apophysis of all these genera with carapace outgrowths indicates a distant relationship with Araneus. All these genera have a ter- minal apophysis (Fig. 7) in the palpus, which was erroneously labeled as a con- ductor by some authors. The evidence from the study of genitalia thus shows that attraction of insects in Kaira and Masto- phora most likely evolved only once, not MASTOPHORA ® Levi 315 twice as thought previously (Stowe, 1986; Table 2). The homology of secreting glands of the insect attractant in Masto- phora and Kaira remains uncertain. TAXONOMIC SECTION Mastophora Holmberg Mastophora Holmberg, 1876: 112. Type species M. extraordinaria Holmberg by monotypy. The gender of the name is feminine. Neave, 1940: 55. It is not preoccupied as claimed by Bonnet, 1957: 1995. Mello-Leitao, 1931: 65. Canals 1931: 17. Roewer, 1942: 900. Gertsch, 1955: 223. Platnick, 2001. Heterocephala Holmberg, 1876: 143. Type species H. conifera Holmberg by monotypy. The gender of the name is feminine. Neave, 1939b: 634. It is not pre- occupied. Glyptocranium Simon, 1895: 885. Type species G. cornigerum Hentz designated by Simon. Neave, 1939b: 484. The gender of the name is neuter. Bonnet, 1957: 1995. First synonymized by Bréthes, 1909. Agatostichus Simon, 1895: 885. Type species A. leu- cacantha Simon by original designation and mon- otypy. Neave, 1939a: 86. Gertsch, 1955: 250. NEW SYNONYMY. Agathostichus:—Simon, 1895: 473. Roewer, 1942: 900. Bonnet, 1955: 181. Platnick, 1998, 2001. Note. Simon spelled the generic name Agatostichus with and without “h,” but the first revisor of the genus, Gertsch (1955), spelled the genus without “h.” Subsequent users must follow the first revisor (Inter- national Code of Zoological Nomenclature, art. 24.2 |International Commission on Zoological Nomenclature, 1999]). The ge- nus is synonymized here because the type species M. leucacantha has long median tubercles on the carapace (Fig. 316). The second species described, M. leucabulba Gertsch, lacks these but has a tubercle be- tween the posterior median eyes and is relatively small (Fig. 288). Gertsch thought small size was diagnostic for the genus, but this is not the case for the third species, M. alvarextoroi, and also not for the type species. Tubercles also are found between eyes in M. corpulenta (Figs. 336, 337). A revision of all species of the genus was made by Mello-Leitaéo (1931) and a revi- sion of Argentinian species was made by Canals (1931). Both authors had shared in- formation. I could not find dates of pub- Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 316 ‘UOHRAIOSGO OU “YURI ‘JUESqR IayRIRYO *— squosoid ToyORIeypo “4 % jeorroyds = ‘IQAOD YJOS sso] osn syjoul 9 DIWY pey uortsod -IPOUL SRO pure podeys-dorp sununy se oures qo ou INU} :Sse] osn syjOUuL 9 DIYSNOUDZIDI, Pee uontsod -Ipoul smeypo pur 2! podeys-dorp sununy se oures qoa ou INU} :ssay] osm syjout p — sary prpoyodAsd_ 9 pwuavjay) podeys sqOUL 07 syons At -o]purds posodxa ze qoa ur yyIS AYONS syjout ‘ds auyoviwphy podeys syOUL 07 syousS -oypurds posodxe iL ais qoM Ur YES AYOHS syoUr [peurs ATFensn shyovdopaa0g SqJOUL OF SyoNS pesodxa ail i qo ut ypIs AyoNS sqour snqopsvd WUDUWLAAYD pearesqo you podeys-doxp posodxa zi G Bo] ppt sejoq stout 9 Sof [peurs pajaulopvy]g podeys op posodxa 10 peatesqo you -urds 10 doxrp — soavoy ur JeotqQe1 a G Ba] popAryr sejoq syQoUuL SMADBPAC aL pedeys-dorp — pasodxo Aqjensn al ite [ Se] Sunys sejoq syjout 9 — sary prpoyodsd 9 paoydojsvjy UO eUSINBOY eceus ard qe YES Pus synpy jo poo SoANVUy snuosy org 38q Suysoy MITES Jo poo *, HOIAVHAD ONICGTING SLI SI SV “SAHOVdOTIOZOd AO LVHL OL YVTILNIS AWAA SI’ ‘dS ANHOVYVINAD AO PAM AH] “IVOLLNACI ATHVAN AYV SAAM AHL “ANTICUN AHL WOUH ONVH AFHL LVHL OS SAVAYHL ONINNIdS SQdOTISVd HO SGNH AHL LY SLNIO! YVWAHS-MOT AHL SYVAYA ANO AI ‘AHM SAFOTISVG V OL UVIINIS SI GVAYHL TVLNOZIYOH HIONIS V WOH ONIONVH STIVE TVWYUAATS GVH HOIHM ARM INVACAZZIC “WAH, ..(GL61 NOSNIGOY GNV NOSNIAOY NI Gh Old ‘O'A) GVAYHL TVLNOZINOH V MOTH ATIVOLLYYA SSH'T WO AYOW ONIONVH WHHM SLAYANNIdS WIAHL WOW MIIS AYOLLS T1Nd ATAALLOV TTY,, ‘GALVLS (TS6T) GUVHYAIY “AM AHL ANV HOIAVHAA ONTTIOd NTIS ONIGHUVOAY (NOLLVOINQWWOO TYNOSUAd ‘MOUNT ‘FE6L “NVOUVAA ‘O86 “AMOLS ‘G61 “NOSNICAOY ANV NOSNIPOY -1O0G “Ty LA VLIHSVAIW ‘S/6L “LIMANY “TS6l ‘GUVHYAIY “SL6T “ANXIO WOUd VLVG) VUYHNAS GHLVIAY GANV VYOHdOLSVPY AO NOILVLIOUNOAY UNV “HAdVHS OVS OOF “HOV Id ONILSAY “(TS6T ‘GUVHYAAY AAS) AAA ANV YOIAVHAA ONITIAd NTIS “SHLOW OL AUS AO SSHANIMOLLS “SLTNGV GNV SHYALYNWI JO GO00d FAL OT TAY L 317 MASTOPHORA ® Levi ‘Juasqe rayoRreyo ‘— Suasard rozORIBYS ‘+ x sapropoxt a JUOUIeTY outds aqo] T LF SUIS o1eq SW auyovivjihny y00} Syont| podeys-1ossep pourds omy oqo] 0 BG UPIPOUL [RAO SPS F advos yyos SW DID DAWY yQ00} smvypo yenboun DIDS a poedeys-1ossep — ‘aurds oqo] I SE URIPOUL [RAO SPS G advos Yos “Mutsy peyeNMonuep —s PLYSMOUDZODI, smevyjo penboun pypavoxa At. podeys-1assep outds = 0 0G azejd sor1ajsod prey aqoy prey ‘areq “mmutey poyeynonuep pwuavjag psourdsiq ai podeys-1ossep outds = il 9g UBIPOUI [BAO ‘SPS Z arRq eSIW— shyovdopoa0g snowuofng ai podeys-1ossep autds 2qo] 0 Fe SUIS Z arRq SW snqopisvd MUDUWAAYD ae qUOWUe ourds 2qo] 0 FI SUIS Z areq SI ou pajautopvj yg snoyuspu ae podeys-1ossep autds = 0 ZI SUIS Z a1eq SIN ou SMUDBPIC a podeys-1ossep autds ae T/O CT SUIS S a1eq SIN ou pioydoysvy stsAydody snjoquiay stsAydody aE) TPA unudsidy unuAsidy WORVOY IPO, sowedg [PUuluLtoy, sndieg u da snd [ed JovdeIRT) jo IOLIQSOg jo ToJUuoA, Boy pur snus) sndeg 6% <2 * S66 “IV LA AOYA'T *L66L “AAU “SS6L ‘SHIAVG “LL61 “LIMAWHA ‘SL6T ‘“ANATO AA SNOILVULSATIL GNV LXHL AO d1TAaH AHL HLIM VLVG NMO AW WOU) SHTVIN AWVAC NI OSTVY GNV VYOHdAOLYAZ PUB VADONAOUW “AdOIDYYV NI LINASAV LAG “AVGIANVYY AO SHTVW NI LNASHUd ATTIVASN ‘HLOOL ALIGNA NV MOV ‘TTY ‘SISAHdOdV NVIGANVUVd V SVH ANON “XICGVY V HAVH TIV AO Id1Vd AHL, SISKHdOdV TVNINYAL AO AZIS UNV AONASAYd *(SQOLNANVIA YO GadVHS-HADOVA) SATOANA ‘(SA TOLLNAG AO MOW ANV “SHNIdS NIHL OAL “ANIdS HALIM) SISAHdOdVY NVIGAW ‘(ALINATIOS V HO “APOT V “LNASAV) SAdTVd NI YHOLONGNOO AO AdVHS GNV AONASAYd AHL ‘AVLAS UVITALVd TVdTVd JO YAAINOAN ‘AOVdVYVO AIVNAA JO HLGIM AOVdVYVO ATVIN AO AOVLINAOUAd *(ASIMYAHLO GNV “ALVId NVIGHIN TVAO HLIM “SLITS OML) WOANADIdH HO YHOIMALSOd *(AdVOS GAZLLOYATIOSNA NV ‘dIT HOINALSOd KINO HLIM ‘AYVE) WONADIdH HO WALNAA (MVIO TVSHUVL ONOT “YONA GHNYV ‘TSW AVLASOYOVW AO MOVT) ATVNAA LIAGV AO NOLLVOIAIGOW OAT ONIGNTONI ‘VHANAD GALVIAY GNV VYOHdOLSVW HOA VLVC IVOIDOTOHdYOW 'G ATAVL 318 lication of the two 1931 articles, but Mel- lo-Leitao’s must have been published first, because Canals cites the page numbers of Mello-Leitao. Gertsch (1955) revised the North American species, and first illustrat- ed the genitalia of the species of his re- gion. Diagnosis. Mastophora differs from oth- er araneid genera by having the carapace with tubercles and having horns (Fig. 8), and lacking macrosetae on legs (Figs. 5, 6, 421). Early instars and males have rows of setae on distal articles of first and second legs (Figs. 5, 6). Mastophora differs from Cladomelea and Ordgarius, which may have horns (Fig. 423), by using the first pair of legs to Thence the bole: Clado- melea and Ordgarius use the second pairs. Description. Female. Carapace color variable, often uniform orange to strongly marked. Legs usually not Ramded! Apdo men often donee anteriorly, lighter pos- teriorly, venter often with a edian white square or rectangle (black in M. hutchin- soni and some M. phrynosoma). Carapace about as wide as long. Eyes small, sub- equal. Anterior eta eyes slightly larger, lateral eyes smallest. Median ocular tr ap- ezoid always longer than wide, widest at posterior median eyes, rarely almost square. Lateral eyes and medians usually on a bulge. Clypeus higher than two an- terior mcd eye Giammcters: Carapace very high, with median tubercles, a pair of horns, usually biforked, and laterally with medium to less large tubercles (Figs. 8, 9), sometimes with short white setae. Chelic- erae with two or three teeth on the ante- rior margin, one on the posterior margin (M. bisaccata has two or three teeth; Fig. 1). Legs without macrosetae, early instars with row of setae on legs as in mature males (Fig. 5). Length of first patella and tibia about 1.1 to 1.8 times width of cara- pace. Second leg longer than first, third shortest. Abdomen wider than long usually with a pair of humps, and a pair of scler- otized discs between humps and anterior margin (Fig. 10), sometimes with addition- al tubercles or scattered clumps of setae Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 or setose. Some species (M. alvareztoroi, M. felis, and M. haywardi) have carapace and abdomen hirsute (Figs. 107-109). The tubercles on the carapace and on horns are not perfectly symmetrical and show indi- vidual differences. The abdominal humps of some species (M. gasteracanthoides) are of variable length. Male. Mature after only two instars or emerging as adult from egg sac (M. cor- nigera), all about 1.7 mm total length. Dirty orange color, sometimes with white spots on carapace or abdomen. Carapace as wide as long with eyes on bulges as in female. Carapace with two median tuber- cles and minute horns. Legs without ma- crosetae but with row of soft setae on first two pairs of legs, slightly shorter than in females (Figs. 5 6). Total length of first patella and Ge about 1.1 fines width of carapace. Abdomen with or without humps, often in the same species, regard- less of the presence of humps on the fe- male abdomen. Female genitalia. The epigynum has plates on the ventral side, and differs only slightly from other species in posterior view (Figs. 2, 13, 20, 27) but has lost all copulatory structures. The posterior has two slits with shadows of atria. Male genitalia. Palpus of male has a pointed median apophysis, an embolus with a simple terminal apophysis (Fig. 7), but no distinct conductor. One group of Mastophora is distinct: the species close to M. gasteracanthoides (corpulenta, rabida, escomeli, obtusa, felis, holmbergi, reimoseri, satan, and diablo). Their carapace is high with vertical sides (Fig. 408); the sides OE the thorax have tu- bercles (Fig. 409), with short, white setae between; ane the abdomen may have high, tube-shaped horns (Fig. 413). The Albee men of all is dark and species can be sep- arated only by studying the genitalia. They have been referred to in Spanish as the cat’s head spiders (arafia cabeza de gato) because of their resemblance to the head of a cat, as can be seen in Figure 421. ‘S4 VA Roe ONO UE MEXICO A NX “—} BELIZE Uw HON. GUATEMALA-S 4 = / 5 SICA a COSTA RIGA - Wz PANAMA +3 ap we JAMAICA MASTOPHORA ® Levi 319 rare TRINIDAD f n= ~ VENEZUELA SOn aN’ A SURINAM oy €oLomeia = / NS fe >- ECUADOR GALAPAGOS Map 1. Relationship. Relationships with other genera are discussed above. Natural History. Mastophora all are un- common and difficult to find. Females rest on branches or leaves of trees, usually 1.5— 3 m high. M. K. Stowe (personal com- munication) has seen egg cases 10 m up and thinks that M. bisaccata seeks out branches high up in trees. They often are found on trees and bushes in orchards, gardens, or along fences. M. K. Stowe (personal communication) reported that most species in Florida are found in for- ests. Sk : NS 2227 YRUGUAY Approximate number of species in various American regions. Distribution. Mastophora are only found in America (Map 1). Misplaced Species. Mastophora fasciol- ata erroneously placed in Mastophora (Levi, 1991: 180) is a Kaira. See below. KEY TO FEMALE MASTOPHORA IL Abdomen without humps (Figs. 10, 24, 88, 130); swellings may be visible in pronler (igs ys) eee ee eee 2 = Abdomen with distinct humps or tuber- cles (Figs. 137, 145, 298, 331, 401) _. 15 (1). INoxrthyAmericas(Mapi2) e= serene eee 3 = South America (Mapsi3 44) oa 11 3(2) Abdomen subtriangular, with anterior 320 (2): Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 lateral swellings (Fig. 88); posterior of epigynum with lateral and ventral lips (Fig. 91); eastern United States (Map SH] B) )) eae ee elt fee sui isa) SUE Ny phrynosoma Abdomen without anterior lateral swell- ings (Figs. 10, 17, 24); epigynum with- out posterior lips (except apalachicola) BE aoe te ete ne eer Pe Ne Ne ay Se ee 4 Posterior of epigynum with slits framed by lateral lips (Fig. 20); southeastern United States (Map 2C) ___ apalachicola Slits without lips (Figs. 13, 27) 5 Slits short, their length distant from ven- tral margin (Fig. 80); eastern United States (Map 2A) yeargani Slits approaching close to ventral margin Geass 7), Ge 6 Dorsum of abdomen with bisaccata pat- tern, anterior orange to gray with spaces and lines (Figs. 42, 53, 65) Abdomen marked otherwise (Figs. 10, 24) Slits approaching each other ventrally, atria their diameter or less apart (Figs. 56, 57); eastern United States (Map 2B) bisaccata Slits parallel or separating ventrally (Eig SrA ai S)) ee a eer 8 Slits parallel (Fig. 45), atria visible in ventral view (Figs. 44); Florida (Map 2G) alachua Atria slightly separating ventrally (Figs. 68-70), atria not visible ventrally (Figs. 67); eastern United States (Map 2G) stowei Atria separating from each other ven- trally (Figs. 34, 35); legs ringed; Vir- ginia to Florida (Map 2A) timuqua Atria approaching each other (Figs. 13, Oye lNeratoey Se ee ee 10 Abdomen black on anterior and _ sides (Figs. 24, 25); atria anterior of seminal receptacles, seminal receptacles their diameter distant from ventral border (Fig. 28); southern Florida (Map 2A) ey nicenee ie, tek Na SE wee eee felda Abdomen with only anterior median area black (Fig. 10); atria at level with large seminal receptacles, seminal re- ceptacles less than their diameter dis- tant from ventral border (Fig. 14); lmlloyato eV (IM eho) XC) satsuma Abdomen hirsute (Figs. 109, 110), slits and atria separating ventrally (Figs. 112, 113); Tucuman, Argentina (Map 3C) haywardi Abdomen with few setae (Figs. 102, 116): atria otherwise (105, 119) 12(11). Abdomen with many black spots and dark lines (Figs. 130, 131): southern Brazil (Map 3B) carpogastra 16(15). 17(16). 18(17). Abdomen marked otherwise (Figs. 102, 116, 123) . Abdomen with light longitudinal lines (Figs. 102, 103); atria approaching each other ventrally (Fig. 105); Santa Catarina, southern Brazil (Map 3C) — a catarina Abdomen marked otherwise (Figs. 116, ]ED 23) ea A ee NU Leelee ccaeelt ae ae eee es 14 . Abdomen marked with pair of black rings, open anteriorly (Fig. 116); Sao Paulo, Brazil (Map 3C) _.._ corumbatai Abdomen with two black discs (Fig. 123); Lara, Venezuela (Map 3A) __.... lara Abdomen with many dorsal or lateral humps (Figs. 267, 298, 300, 310, 331, RSIS) WR Mer ete Tania tee Tae ee ee URE Sa 16 Abdomen with one pair of humps (Figs. 155, 171), rarely median area of ab- domen swollen (Fig. 325) 19 Humps anterior on each side of abdo- men (Fig. 267); Santiago del Estero, Acgentunas (Mayor Si) ee eae abalosi Humps or tubercles dorsally (Figs. 298, MOL SSM Brae Nike cle aes aaa 17 Carapace with short tubercles (Figs. 329, 330); abdomen with numerous small dorsal tubercles (Figs. 331, 332): northern Argentina to Buenos Aires Province (Mapi3G@)ss sans conifera Carapace with large tubercles (Figs. 297, 3()9))gesie 2 ane Can eT UBS aera pseu eaeee 18 Carapace with median tubercles spine- shaped (Fig. 309); Panama (Map 2F) LS Sn SEED eh in TS soberiana Carapace with median tubercles cone- shaped (Fig. 297); Texas to Chiapas (Map 2F) alvareztoroi ; Carapace median tubercles same size or longer than horns (Figs. 288, 316, 323) AS Sata et lichen AA te et hed i la ln 5 EN 20 Carapace with median tubercles smaller thanshoms)(Higs: 1365.38)l)) )iemseene 22, . Median tubercles spine-shaped, very long (Fig. 316); Bahia to Rio de Ja- neiro, Brazil (Map 3G) _....... leucacantha Median tubercles otherwise (Figs. 288, SOS) es OE NR EE AE Sea ee 21 . Median tubercles cone-shaped (Fig. 323); Rio Grande do Sul, Brazil (Map 3G) brescoviti Carapace tubercles large, rounded on dark carapace (Fig. 288); Texas, Mex- Itcfo) (UN eto) PAN) Se leucabulba . Carapace with sides vertical (Figs. 366, 380), sides with tubercles (Figs. 367, 381); Mexico to South America (Map 721) nie ee eel ae TEs me ETE Sune ta 39 Carapace with sides slanting (Figs. 135, 169): sides with few or no tubercles (HigstllS65ell7(0;"Miapse2 3) ea=eeaenes 23 MASTOPHORA ° Levi apalachicolag@ satsuma YW stowei @ A felda g@ timuqua @ yeargani archeri o hutchinsoni @ alachuaa YW cornigera @ seminole A vaquera SP. Ss Qq Se ; —> @ alvareztoroi wv corpulenta ; A fasciata au @ Jeucabulba ° §) /\ @ soberiana ‘y iG Map 2. Distribution of Mastophora species of North and Central America. 321 322) Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 catarina @ corumbatai V hayward; 4& @ dizzydeani @ fasciata BRAZIL - ~~~ - A BRAZIL~ { BRAZIL_ | LZ - Tbe =e) NN MS «\ Woe cw KN Q abalosi Vv comma Vv melloleitaoi @ yacare A ypiranga cranion A longiceps w } pesqueiro @ pickeli A. piras extraordinaria F brescoviti V conifera @ leucacantha @ Map 3. Distribution of Mastophora species of South America. 34(33). . North America and Cuba (Map 2) _....... 94 Central and South America (Map 3) . Cuba (Map 2G); abdomen with anterior, lateral swellings (Figs. 145, 146) ee ee eee ee vaquera Continental North America; abdomen without lateral swellings (Figs. 137, Ay Sy) Re nem Aik ey otic se Ree eee 25 . Posterior of epigynum with slits and atria approaching each other ventrally (Figs. 159, 160); northeastern United Steltes (Mizyo 2B) ee hutchinsoni Slits and atria otherwise (Figs. 140, 174, ARS SS) prgetrrere mee eccrine a Maas oases EA 26 . Posterior of epigynum with slits framed by a lip on each side (Fig. 188); south- eastern United States (Map 2E) — archeri Slits without lips (Figs. 140, 174) if . Slits parallel and approach ventral bor- der (Fig. 174); eastern United States to California and Honduras (Map 2G) po ie Eee SESE Ebon wed Tecate ok Aaah ae cornigera Slits short, their length apart from ven- tral border (Fig. 140); southern Flori- (6 (Fie TT cy cee ARE Se aR ae seminole . Central America and northern South Niraveraver) (we) leteinby =e 29 Southern South America, from Pernam- buco State, Brazil, in the north 30 . Humps of abdomen broad swellings (Fig. 196); slits on posterior of epigyn- um approaching each other (Fig. 199); Costa Rica to Venezuela (Map 3A) — oe a ee fasciata Humps of abdomen narrow (Fig. 207); slits separating ventrally (Fig. 210); Colombia to Peru (Map 3A) — dizzydeani . Humps of abdomen extended to a point (Figs. 271, 272); Santiago del Estero, Argentina (Map 3E) comma BOERS Sa pe Vary a aL oe pg ee ee Hee Ve et 31 . Each side of abdomen with a pair of nar- row dark-framed longitudinal, white marks (Figs. 239, 240); humps small (Fig. 240); Rio Grande do Sul, Brazil (Nia c3ID)) i lee eee pesqueiro Abdomen without such marks _ 32 . Posterior of epigynum with a lip on each side between slits and ventral margin (Fig. 263); Uruguay (Map 3E) _.. yacare Posterior of epigynum without such lips (ale OG) ae ee 33 . Epigynal slits appear forked (Figs. 235, 236); carapace horns unusually thick and laid back (Figs. 230, 231); Sao Paulo, Brazil (Map 3D) _......... longiceps Epigynal slits simple (Figs. 249, 256); horns small (Figs. 245, 252) __...-... 34 Epigynal slits separating ventrally (Figs. 228, 249) 35(34). 36(34). 45(44). MASTOPHORA ® Levi 323 Epigynal slits and atria approaching each other ventrally (Figs. 256, 286) Abdomen with a pair of dorsal stippled dark spots (Fig. 225); Pernambuco State, Brazil (Map 3D) cranion Anterior of abdomen black (Fig. 246); Minas Gerais, Sao Paulo states, Brazil GAY C0) S6)) D)) te eee lee eee Zales piras Anterior of abdomen light, light area bordered posteriorly by dark (Fig. 283); epigynum as in Fig. 286; south- erm Brazil, northern Argentina (Map 3 Fg ete LA trac de el extraordinaria Anterior of abdomen black (Figs. 218, 253, 276) . Atria their diameter or less apart (Figs. 256, 257); Rio de Janeiro to Santa Ca- tarina, Brazil (Map 3E) _...... ypiranga Atria more than their diameter apart ; (Figs. 221, 222; 279; 280) 38 . Atria with median knobs facing each oth- er (Fig. 280); Parana to Buenos Aires, Argentina (Map 3E) melloleitaoi Atria without knobs (Fig. 222); Pernam- buco; Brazil (Map 3D) = pickeli . Central America (Map 4A); posterior of epigynum with dark patch dorsally in each of pair of depressions (Fig. 341) corpulenta OUnNerraelo ns 40 . Galapagos (Map 4A); abdomen with only tiny humps (Fig. 346); epigynum slits each with a loop ventrally (Figs. 348, 349) rabida Other regions; abdomen with larger humps . Peru (Map 4A); posterior of epigynum, between slits, with median area having a bulge on each side (Fig. 355) escomeli Otherre gions yeast arsme een wen Lt 42 . Chile (Map 4B); posterior of epigynum with a dark spot dorsally in each ad- jacent depression (Fig. 415) Eh a ee Na gasteracanthoides Brazil to Argentina 2). Abdomen with humps placed on swollen area (Figs. 364, 365); Pernambuco State, Brazil (Map 4C) obtusa Humps not placed on swollen area (Figs. 368, 369) . Dark patch (atria) placed dorsal or mid- dle of seminal receptacles (Figs. 392, 404) Atria absent or placed ventrally of sem- inal receptacles (Figs. 372, 379, 386) tee fy dhe Dalal Uae ead dicen seat k eS 46 Posterior of epigynum with dark patch (atria) dorsal within a depression (Figs. 392, 394, 395); Pernambuco, Brazil, to central Argentina (Map 4B) satan 324 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 pura?” : CB) canaGua B corpulenta @ escomeli w ‘abida wv felis & holmbergi QO obtusa @ ‘eimoseri diablo D Map 4. Distribution of Mastophora species of the M. gasteracanthoides group. = Atria lateral, placed outside of depres- = Slits without such bend (Figs. 371, 378) sionu(Higs 404) mortherneicentralAve? Tay) | eee to ae ee ee eee AT gentinan(Mapw4 1D) ese) ser diablo 47(46). Posterior of epigynum with slits separat- 46(44). Posterior of epigynum with slits with a ing ventrally and with lateral lip (Figs. ventral bend (Fig. 385); Paraguay 371, 372); Rio de Janeiro, Sao Paulo, (Map AG) 18s e ete eer oe ee reimoseri sievalll (UMleie) 4G) ee felis Slits parallel (Fig. 378); Paraguay, San- tiago del Estero (Map 4C) __. KEY TO KNOWN NORTH AMERICAN MALE MASTOPHORA Space surrounded by median apophysis, in ectal view of palpus, longer than wide (Figs. 50, 62, 74, 164, 168) Space surrounded by median apophysis wider than long (Figs. 39, 85, 96, 99, 179, 182); terminal apophysis usually shorter than embolus (Figs. 37, 83, 94) Terminal apophysis shorter than embo- lus (Fig. 48); median apophysis of pal- pus very short (Figs. 48-50); Florida (Mian? CG) aes eee ee Terminal apophysis as long or longer than embolus (Figs. 7, 60) Base of embolus large (Figs. 60, 61); eastern United States (Map 2B) ——— holmbergi alachua ae ee Se ea SI a bisaccata Base of embolus small (Figs. 72, 73) Base of median apophysis rounded (Figs. 162, 166); in ectal view, narrow part of median apophysis shorter than base (Figs. 164, 168); northeastern United States (Map 2E) _.... Base of median apophysis angular (Fig. 72, 73); in ectal view, narrow part of median apophysis as long or longer than width of base (Fig. 74); eastern Wnited States! (Map. 2©) = es Terminal apophysis almost as long as embolus (Figs. 94, 177, 180) Terminal apophysis about half length or less of embolus (Figs. 83, 191) ........... Base of median apophysis longer than wide (Figs. 177, 180); southern United States to California and Central Amer- jean (Mawr 2 Chon. sn: SA eee Base of median apophysis short (Figs. 94, 98); eastern United States (Map 2D) Base of median apophysis large, touch- ing embolus (Figs. 83, 84); eastern United States (Map 2A) Base of median apophysis small (Figs. 37, 191) Length of narrow part of median apoph- ysis as wide as base in ectal view (Fig. 193); Gulf Coast, Kansas (Map 2E) — 4 hutchinsoni stowei cornigera phrynosoma yeargant Ba nee ae wR AEN ce pe Ae RP archeri Length of narrow part of median apoph- ysis longer than width of base (Fig. 39); Virginia to North Carolina (Map CIN) eviee Sree e ine Mier ikea Ae! timuqua MASTOPHORA ° Levi 325 Kaira altiventer (O. P.-Cambridge), new combination Epeiroides fasciolata O. P.-Cambridge, 1889: 15, pl. 8, fig. 5, 3d. Male from Bugaba, Panama, in BMNH, examined; now lost. Keyserling, 1893: 309, pl. 16, fig. 228, d. Male from Guatemala. Kaira altiventer O. P.-Cambridge, 1889: 56, pl. 3, fig. 13, 2. Female from Veragua |Veraguas Prov.], Pan- ama, in BMNH, examined. Levi, 1993b, 213, figs. 3-22, 2, 6. NEW SYNONYM Y. Aranea fasciolata: —F. P.-Cambridge, 1904: 519, pl. 51, fig. 5, d. Claims that Keyserling’s specimen is lost and probably was misidentified according to F. P.-Cambridge, 1904. Note. Epeiroides fasciolata is a Kaira. My unpublished illustration of the holo- type of E. fasciolata shows the distinct large median apophysis tooth at the base of the flagella, the characteristic curved, long, soft conductor, and the drop-shaped, sclerotized terminal apophysis of Kaira al- tiventer (Levi, 1993b, figs. 20, 21). I examined the type in 1967, when vis- iting the BMNH, and made a drawing of the palpus of the male, thinking errone- ously that the species is a Mastophora, But the palpus is not that of Mastophora. Nei- ther O. P.-Cambridge, F. P.-Cambridge, or Keyserling showed carapace tubercles. I overlooked the species when revising Kaira. Since 1967, the holotype has been misplaced and cannot be found. Mastophora satsuma new species Figures 8-14; Map 2C Holotype. Female holotype from Riverview, 11 mi. [17.6 km] SE of Tampa, on Highway 301, Hills- borough Co., Florida, on satsuma, Citrus nobilis (tangerine tree), 23 Aug. 1966 (E. R. Simmons), in FSCA. The specific name is a noun in apposition after the tree on which the holotype was collected. Description. Female holotype. Carapace orange-brown. Chelicerae, labium, endites light brown. Sternum grayish orange. Cox- ae and distal leg articles brown. Abdomen dorsum whitish with dark gray frame hav- ing a lobe extending posteriorly to midline (Fig. 10); venter gray with white square. Carapace, with few tubercles (Figs. 8, 9) and short white setae. Abdomen without humps (Fig. 10). Total length 9.6 mm. 326 Carapace teat mm long, 4.0 wide in tho- racic region, 2.4 wide a lateral eyes. First femur 4 1 mm, patella and tibia 5.2, meta- tarsus 3.5, tarsus 1.0. Second patella and tibia 4.0 mm, third 2.3, fourth 3.6. Length of first patella and tibia 1.1 times width of carapace. Males are not known. Variation. The epigynum is asymmetri- cal: the left slit is more curved than the right one and the left seminal receptacles are larger than the right ones (Fig. 14). Both ccminal receptacles are oval. Diagnosis. Mastophora satsuma is dis- tinguished from M. felda (Figs. 22-28) by being smaller, by differences in dorsal pat- tern (Fig. 10), by having larger seminal re- ceptacles (Fig. 14), ariel Alee. by the larger depression in ras midline of the epigynum (Fig. 14). Distribution. Central Florida (Map 2C). Specimens Examined. No other specimens have been found. Mastophora apalachicola new species Figures 15-21; Map 2C Holotype. Female holotype from ravine, Bristol, Cal- houn Co., Florida, 29 Dec. 1939 (A. F. Archer), in AMNH. The specific name is a noun in apposition after the name of the river at the locality. Description. Female holotype. Carapace contrastingly marked, sides dark brown, dorsum light brown anteriorly, pair of forks lightest brown (Figs. 15, 16). Chelic- erae yellow- white with a ae patch on sides. Labium, endites dark brown. Ster- num anterior light, posterior dark brown. Coxae dusky brown, fourth darkest. Distal leg articles yellow-white, femora and pa- tellae with brown bands. Abdomen white (Fig. 17), dorsum with a pair of black spots, venter with white square containing three pairs of black dots. Carapace with few tubercles, with very large forked horns and with short white setae on sides (Figs. 15, 16). Median eyes on bulge, lateral eyes on bulges. Abdomen without humps and with large distinct dorsal pair of discs (Fig. 17). Total length 8.8 mm. Carapace 3.5 mm long, 3.4 wide in thoracic region, 2.2 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 wide at lateral eyes. First femur 3.3 mm, patella and tibia 4.4, metatarsus 3.2, tarsus 1.0. Second patella and tibia 3.3 mm, third 1.8, fourth 3.0. Length of first patella and tibia 1.3 times width of carapace. Males are not known. Variation. Total length of females 8.8 to 9.0 mm. The specimen from Levy County has the atria larger and more spherical than those of the holotype (Fig. 21); the one from Hamilton Co. has the atria larger and the seminal receptacles much larger. The illustrations were made from the ho- lotype. Diagnosis. Mastophora apalachicola is distinguished from others by the contrast- ing carapace coloration (Figs. 15, 16), the abdomen lacking humps, ‘lacking dorsal color pattern, and having large Boreal discs (Fig. 17). The horns (Fig. 15) are larger than those of M. timuqua (Fig. 29) and M. satsuma (Fig. 8). The epigynum, unlike that of similar species, has a lip on each side; the slits are in a slight depression (Fig. 20). The epigynal slits and atria are almost parallel (Figs. 20, 21). Distribution. South Carolina to north- em Florida (Map 2C). Paratypes. SOUTH CAROLINA Anderson Co.: Simpson Agric. Exp. Station, 16 Aug. 1974, 1 imm. (R. Paigler, CUAC). FLORIDA Hamilton Co.: nr. White Springs, Big Shoals State Forest, 25 Nov. 1991, 12 (M. K. Stowe 2116, FSCA). Levy Co.: Manatee Springs State Park, 10 Nov. 1992, 12 (M. K. Stowe 2114, MCZ). Mastophora felda new species Figures 22—28; Map 2A Holotype. Female holotype from near Felda, Hendry Co., Florida, in orange grove, 8 March 1993 (D. Smith), in FSCA. The specific name is a noun in apposition after the type locality. Description. Female holotype. Carapace dark orange-brown. Chelicerae dusky brown. Labium, endites dusky brown. Sternum brownish orange. Coxae orange- brown, lighter than Stemi and legs. Dis- tal leg spetales dark orange- esate Abdo- men anterior, sides, and enter gray (Fig. 24), center and posterior whitish; venter MASTOPHORA ® Levi 327 seminal receptacle now atrium satsuma terminal apophysis radix embolus tegulum conductor area median apophysis Figures 1-7. Mastophora. 1-4, female. 1, M. bisaccata, left tip of chelicera and fang from posterior. 2, epigynum, diagram- matical. 3, dish with paraffin to examine epigyna. 4, M. diablo, epigynum cleared, in posterior view, showing ducts. 5-7, male. 5, M. gasteracanthoides. 6, M. bisaccata. 7, M. gasteracanthoides left palpus without cymbium, mesal view. Figures 8-14. M. satsuma new species, female. 8, 9, carapace and chelicerae. 8, frontal. 9, lateral. 10, 11, carapace and abdomen. 10, dorsal. 11, lateral. 12-14, epigynum. 12, ventral. 13, posterior. 14, posterior, cleared. Figures 15-21. MM. apalachicola new species, female. 15, 16, carapace and chelicerae. 15, frontal. 16, lateral. 17, 18, carapace and abdomen. 17, dorsal. 18, lateral. 19-21, epigynum. 19, ventral. 20, posterior. 21, posterior, cleared. Figures 22-28. WM. felda new species, female. 22, 23, carapace and chelicerae. 22, frontal. 23, lateral. 24, 25, carapace and abdomen. 24, dorsal. 25, lateral. 26-28, epigynum. 26, ventral. 27, posterior. 28, posterior, cleared. Scale lines. 1.0 mm; genitalia, 0.1 mm. 328 with median white square. Carapace with few tubercles and with short, white setae (Figs. 22, 23). Median eyes on bulge, lat- eral eyes on bulges. Abdomen without humps (Fig. 24). Total length 12.3 mm. Carapace 5.3 mm long, 5.4 wide in tho- racic region, 3.0 wide at lateral eyes. First femur 5.2 mm, patella and tibia 6.7, meta- tarsus 4.6, tarsus 1.2. Second patella and tibia 5.1 mm, third 2.7, fourth 4.8. Length of first patella and tibia 1.2 times width of carapace. Males are not known. Variation. The holotype has only seven eyes; it lacks the left posterior median eye. Diagnosis. Mastophora is distinguished from M. bisaccata by the different color- ation of the abdomen (Fig. 24), by the epi- gynum having only a thin rim in ventral view (Fig. 26), and by the small dorsal knobs in the depression on the posterior of the epigynum (Fig. 27). The atria bend toward each other (Fig. 28), but are far- ther apart than those of M. bisaccata (Fig. Be Distribution. (Map 2A). South-central Florida Specimens Examined. No other specimens have been found. Mastophora timuqua new species Figures 29-39, 445; Map 2A Holotype. Female holotype from Devil’s Millhopper State Park, Gainesville, Alachua Co., Florida, 19 Nov. 1983 (M. K. Stowe 107A), in MCZ. The spe- cific name is a noun in apposition after an extinct, northern Florida Indian tribe. Description. Female holotype. Carapace light brown, with sides and eye areas dark- er brown and white mark in center (Figs. 29, 30). Chelicerae, labium, endites brown. Sternum light brown. Coxae lighter than sternum, digs leg articles with dark brown rings. omens gray with anterior darker (Fig. 31), venter with indistinct white square. Thorax with short white se- tae, a distinct narrow line around margin; lacking large tubercles on sides of thoracic region (Figs. 29, 30). Median eyes on a bulge, lateral eyes on bulges. Abdomen Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 without humps. Total length 8.5 mm. Car- apace 3.5 mm long, 3.4 wide in thoracic region, 1.8 wide at lates al eyes. First femur 2.9 mm, patella and tibia 4.1, metatarsus 2.7, tarsus 0.8. Second patella and tibia 3.3 mim, third 1.8, fourth 2.7. Length of first patella and tibia 1.2 times width of cara- pace. Male allotype. Carapace orange with white median patch. Sternum orange. Coxae, legs lighter orange. Abdomen dusky orange. Abdomen with two adjacent humps. Total length 1.6 mm. Carapace 0.88 mm long, 0.79 wide in thoracic re- gion, 0.53 wide at lateral eyes. First femur 0.78 mm, patella and tibia 0.78, metatarsus 0.45, tarsus 0.28. Second patella and tibia 0.69 mm, third 0.40, fourth 0.55. Length of first patella and tibia same as width of carapace. Note. Males were raised from egg sac of M. timuqua. Variation. The illustrations were made from the female holotype and male allo- type. Diagnosis. Mastophora timuqua is dis- tinguished from both M. bisaccata and M. pisgah by lacking dorsal abdominal pattern (Fig. 31), by having ringed legs, by having ies posterior of the epigynum with atria separated, and by having a ridge in the midline (Fig. 34). The male has a wide space enclosed by the median apophysis (Fig. 39) and differs from the male of M. cornigera by the prox- imal position of the radix (Fig. Sas The egg sac is shown in Figure 445. Distribution. North C@xolina to north- ern Florida (Map 2A). Paratypes. NORTH CAROLINA Moore Co.: reared from egg sac in spring 1941, 3d (J. Perry, M. K. Stowe 2113, “MCZ). FLORIDA Alachua Co.: Deve ils Millhopper State Park, reared spring 1992, ¢ al- lotype, 11 paratypes (M. K. Stowe 2101, MCZ, AMNH); spring 1992, raised 3d (M. K. Stowe 2106, FSCA). Levy Co.: Manatee Springs State Park, 5 Nov. 1987, 12 (M. K. Stowe 2111, AMNH). Hillsborough Co.: Pinecrest Alderman Ford County Park, 9 May 1988, egg sac (M. K. Stowe 21050, FSCA). MASTOPHORA ° Levi 329 timuqua Figures 29-39. Mastophora timuqua new species. 29-35, female. 29, 30, carapace and chelicerae. 29, frontal. 30, lateral. 31, 32, carapace and abdomen. 31, dorsal. 32, lateral. 33-35, epigynum. 33, ventral. 34, posterior. 35, posterior, cleared. 36-39, male left palpus, stained. 36, apical. 37, mesal. 38, ventral. 39, ectal. Figures 40-50. WM. alachua new species. 40-46, female. 40, 41, carapace and chelicerae. 40, frontal. 41, lateral. 42, 43, carapace and abdomen. 42, dorsal. 43, lateral. 44-46, epigynum. 44, ventral. 45, posterior. 46, posterior, cleared. 47-51, male left palpus, stained. 47, apical. 48, mesal. 49, ventral. 50, ectal. Scale lines. 1.0 mm; genitalia, 0.1 mm. 330 Mastophora alachua new species Figures 40-50; Map 2G Holotype. Female holotype from Devil’s Millhopper State Park, Gainesville, Alachua Co., Florida, 19 Nov. 1983 (Mark K. Stowe 107B), in MCZ. The species is named after the type locality. Description. Female holotype. Carapace orange. Chelicerae, labium, endites or- ange. Sternum orange. Coxae and distal leg articles orange. Abdomen orange with a dorsal, anterior dusky area containing some bare patches, and a transverse gray line posteriorly (Fig. 42); venter with white square. Carapace granular rather than tu- bercular, sides of carapace with short white setae; no tubercles on lateral thorac- ic region (Fig. 41). Abdomen subtriangu- lar, without humps (Fig. 43). Total length 8.0 mm. Carapace 3.4 mm long, Sv ide in thoracic region, 2.2 wide at ieteral eyes. First femur 3.5 mm, patella and tibia 4.5, metatarsus 3.2, tarsus 1.0. Second patella and tibia 3.5 mm, third 1.8, fourth 2.9. Length of first patella and tibia 1.3 times width of carapace. Male allotype. Carapace dusky orange with median white patch. Sternum, legs, abdomen dusky orange. Abdomen oh pair of adjacent humps. Total length 1.6 mm. Carapace 0.78 mm long, 0.67 wide in thoracic region, 0.52 wide Ae lateral eyes. First fom 0.67 mm, patella and tibia 0.78, metatarsus 0.42, tarsus 0.11. Second patella and tibia 0.65 mm, third 0.38, fourth 0.54. Length of first patella and tib- ia 1.1 times width of carapace. Note. The male was raised from an egg sac. Variation. Total length of females 8.0— 9.2 mm, males 1.6—1.7. The illustrations were made from female holotype, correct- ed with the paratypes. The male illustrated was the only one available, the allotype, whose palpus was expanded, and thus the median apophysis (Figs. 47-50) may not be at the same angle as in the contracted palpus. Diagnosis. Mastophora alachua is distin- guished by the abdomen, slightly triangu- Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 lar in shape and having markings like those of M. bisaccata (Fig. 42), and by the epi- gynum, in ventral view showing two dark areas, the atria, on the posterior margin, and a narrow double margin (Fig. 45), and on the posterior, parallel slits, slightly clos- er ventrally, and between, next to each other, two shallow U-shaped shadows (Fig. 45). Distribution. AE) Paratypes. FLORIDA Alachua Co.: Devil's Mill- hopper State Park, 21 Nov. 1983, 12 (M. K. Stowe 106, MCZ); no date, prob. 1984, allotype d (M. K. Su 2102, MCZ); Gainesville, 1 Nov. 1990, 12 (M . Stowe 2115, FSCA). Northern Florida (Map Mastophora bisaccata (Emerton) Figures 51-62, 446; Map 2B Cyrtarachne bisaccata Emerton, 1884: 325, pl. 34, fig. 11, °, pl. 38, fig. 12, egg sac. Female holotypes from beech tree, New Haven, Connecticut, in MCZ, examined. C. multilineata Atkinson, 1888: 546. Two syntypes presumably from near Chapel Hill, North Carolina, lost. First synonymized by Banks (1910). Ordgarius bisaccata: -—Keyserling, 1892: 42, pl. 2, fig. 35, 2. McCook, 1894: 198, pl. 12) figs. 2: 3, oe Glyptocranium bisaccatum:—Bonnet, 1957: 1996. Ordgarius obesus Banks, 1898: 250, pl. 15, fig. 9, °. Two female syntypes from La Chuparosa [Chupar- rosa, San Luis Potosi], Mexico, in CAS, destroyed. NEW SYNONYMY. Mastophora_ bisaccata:—Mello-Leitaéo, 1931: 71. Roewer, 1942: 900. Kaston, 1948: 232, figs. 737— 740. Gertsch, 1955: 242, pls. 3-5, pl. 6, figs. 1, 4; text figs. 19-23, 35, 43, 44, 2, ¢d. Platnick, 1997: 513. Platnick, 2001. Note. Atkinson did not tell how Cyrtar- achne multilineata differs from bisaccata, although he mentioned the latter species. The large size, total length 11 and 13 mm, abdomen 13 and 15 mm wide, suggest ne had M. bisaccata. The name was first syn- onymized by Banks (1910). Ordgarius obesus differs from bisaccata, according to Banks, by being larger in size and having the cephalothorax truncate. The illustration shows the dorsal abdomi- nal pattern of M. bisaccata. The size is within the range of M. bisaccata and the carapace, unlike most Mastophora, is trun- cate. There is no doubt that this was M. MASTOPHORA ° Levi 331 Se Wo stowei bisaccata Figures 51-62. Mastophora bisaccata (Emerton). 51-58, female. 51, 52, carapace and chelicerae. 51, frontal. 52, lateral. 53, 54, carapace and abdomen. 53, dorsal with male. 54, lateral. 55-58, epigynum. 55, ventral. 56, posterior. 57, 58. posterior, cleared. 57, (Virginia). 58, (Florida). 59-62, male left palpus, stained. 59, apical. 60, mesal. 61, ventral. 62, ectal. Figures 63-74. M. stowei new species. 63-70, female. 63, 64, carapace and chelicerae. 63, frontal. 64, lateral. 65, 66, carapace and abdomen. 65, dorsal, with male. 66, lateral. 67-70, epigynum. 67, ventral. 68, posterior. 69, 70, posterior, cleared. 69, (Florida). 70, (North Carolina). 71-74, male left palpus, stained. 71, apical. 72, mesal. 73, ventral. 74, ectal. Scale lines. 1.0 mm; genitalia, 0.1 mm. 332 bisaccata. The locality of this specimen is doubtful, because it came from the Marx collection (see Banks, 1898). Description. Female from Rhode _Is- land. Carapace orange-brown. Sternum dark orange-yellow. Legs orange yellow, first femur darker ventrally. Abdomen white, anterior dorsally gray with charac- teristic pattern (Fig. 53), venter with white square. Carapace with very small tuber- cles, and small eye projections (Figs. 51— 53). Abdomen without humps (Fig. 53). Total length 14.2 mm. Carapace 5.2 mm long, 5.4 wide in thoracic region, 2.8 wide at lateral eyes. First femur 4.4 mm, patella and tibia 6.3, metatarsus 4.3, tarsus 1.2. Second patella and tibia 4.7 mm, third 2.7, fourth 4.6. Length of first patella and tibia 1.2 times width of carapace. Male from Arkansas. Carapace orange- brown with white line in middle, branch- ing posteriorly into tubercles (Fig. 6). Ster- num white. Lateral eyes smaller than me- dian. Legs colorless yellowish. Abdomen white. Abdomen with slight, indistinct tu- bercles (Fig. 6). Total length 1.8 mm. Car- apace 0.88 mm long, 0.81 wide in thoracic region, 0.55 wide at lateral eyes. First fe- mur 0.92 mm, patella and tibia 0.93, meta- tarsus 0.48, tarsus 0.27. Second patella and tibia 0.79 mm, third 0.47, fourth 0.59. Length of first patella and tibia 1.1 times width of carapace. Note. Examined males include males raised from egg sacs by K. Yeargan and males collected in the same locality as fe- males. Variation. The holotype has the epigynal slits farther apart than in the specimen il- lustrated. Total length of females 9.0-15.3 mm. Males may have humps on the ab- domen. The illustrations were made from the female holotype and males from Ar- kansas and Ohio. An egg sac collected with Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 female at Bushnell, Florida, lacked flaps and was smooth. Diagnosis. The female of M. bisaccata is distinguished from the similar M. alachua, M. stowei, and M. yeargani by being larg- er, and having the atria approaching each other in posterior view of the epigynum (Figs. 56-58). The male has a narrow space encircled by the median apophysis in ectal view (Fig. 62), and the median apophysis has a longer base (Fig. 60) than in M. hutchinsoni. The egg sac has a heavier stalk than in other Mastophora species (Fig. 446). Natural History. Mastophora bisaccata has been collected from bittersweet in Connecticut, field of cemetery in Illinois, and on dogwood in Florida. Females rest under leaves, and sometimes neighboring leaves are stitched together; the females may look like leaf galls in Florida (M. Stowe, personal correspondence); in North Carolina they may resemble tree snails (Atkinson, 1888). Distribution. Eastern United States (Map 2B). Specimens Examined. CONNECTICUT Litchfield Co.: Kent, on bittersweet, Sept. 1937, 12 (AMNH). NEW YORK Nassau Co.: Long Island: Sea Cliff, 1d (MCZ). NEW JERSEY Elizabeth Co.: Roselle Park, 25 Sep. 1910, 12 (AMNH). Middlesex Co.: New Brunswick, July 1930, 12 (AMNH). PENNSYLVA- NIA Westmoreland Co.: 4.8 km S Rector, 13 Sep. 1966, 12 (B. Vogel, DMNS). OHIO Jackson Co.: Oak Hill, 12 (R. A. Reller, OSU). Logan Co.: Can- twell Cliffs, 8 Sep. 1935, 12 (OSU). Butler Co.: Bachelor Woods, Oxford, 11 July 1998, 1d (D. M. Golden, OSU). DISTRICT OF COLUMBIA Wash- ington, Sept., 1d (Fox, CUC, AMNH); summer 1935, 12 (H. E. Ewing, USNM). VIRGINIA Falls Church, 12, 1¢ (MCZ). Powhatan Co.: Powhatan, Sep. 1984, 12 (A. Moreton, MCZ); 1985, 12 (A. Moreton, MKS). KENTUCKY Jessamine Co.: imm., 73 raised spring 1995 (K. V. Yeargan, KVY). SOUTH CAROLINA Oconee Co.: Clemson College, 1° (MCZ). Lexington Co.: Batesburg, 12 (MCZ). GEORGIA Fulton Co.: Atlanta, 2 Aug. 1937, 12 (F. => Figures 75-85. Mastophora yeargani new species. 75-81, female. 75, 76, carapace and chelicerae. 75, frontal. 76, lateral. 77, 78, carapace and abdomen. 77, dorsal. 78, lateral. 79-81, epigynum. 79, ventral. 80, posterior. 81, posterior, cleared. 82-85, male left palpus, stained. 82, apical. 83, mesal. 84, ventral. 85, ectal. MASTOPHORA °® Levi 333 Figures 86-99. MM. phrynosoma Gertsch. 86-92, female. 86, 87, carapace and chelicerae. 86, frontal. 87, lateral. 88, 89, carapace and abdomen. 88, dorsal, with male. 89, lateral. 90-92, epigynum. 90, ventral. 91, posterior. 92, posterior, cleared. 93-99, male left palpus, stained. 93-96, (Kentucky). 97-99, (Florida). 93, apical. 94, 97, mesal. 95, 98, ventral. 96, 99, ectal. Scale lines. 1.0 mm; genitalia, 0.1 mm. 334 W. Fattig, AMNH). FLORIDA Alachua Co.: Devil's Millhopper State Park, July, Aug. 1978-1981, 52 (M K. Stowe, MKS); June, July 1981, OMEN Ke Stowe, MKS): 19 Nov. 1983, 12; Gainesville, 26 July 1980, penult. (C. Hieber, MCZ); High Springs, Oct. 1935, 12 (H. H. Simpson, USNM). Indian River Co.: Se- bastian, Apr. 1944, 12 (G. Nelson, MCZ). Lake Co.: Umatilla, Sep. 1954, 12 (M. W. Tyler, AMNH). Sum- ter Co.: Bushnell, 24 Oct. 1979, 12 (W. Edwards, FSCA). ILLINOIS Franklin Co.: Zeigler, 10 May 1928, 12 (J. K. Carlovic, MCZ). Jackson Co.: Little Grand Canyon, S. Murphysboro, 5 Sep. 1971, 1d (N. Magnuson, JAB). Williamson Co.: Canterville, field of cemetery, 14 Oct. 1978, 12 (R. Reith, JAB). MIS- SOURI St. Louis Co.: 17 Feb. 1940, 1d paratypes of M. archeri (W. M. Gordon, AMNH). ARKANSAS Carroll Co.: Berryville, July 1942, 1d (C. Wilton). MISSISSIPPI Harrison Co.: Gulfport, 12 (AMNH). Mastophora stowei new species Plate 1; Figures 63-74, 447; Map 2C Holotype. Female holotype from American Entomo- logical Institute, Gainesville, Alachua Co., Florida, 29°3 6.0'N, 82°22.0'W, 7 Dec. 1987 (M. Stowe 07001), in MCZ. The species is named after the collector, Mark Stowe, who has contributed much to our knowledge of Mastophora. Description. Female holotype. Carapace orange-brown, dusky in eye region (Fig. 65). Chelicerae light orange, labium, en- dites dusky orange. Sternum orange. Legs light orange, dusky dorsally. Abdomen dor- sum whitish with gray pattern anteriorly, having distinct white spots and_ streaks (Fig. 65); venter with white square. Cara- pace with indistinct tubercles; horns al- most rectangular in anterior view (Figs. 63, 64). Abdomen without humps and slightly pointed posteriorly (Fig. 65). Total length 8.0 mm. Carapace 3.5 mm long, 3.4 wide in thoracic region, 2.1 wide at lateral eyes. First femur 3.4 mm, patella and tibia 4.5, metatarsus 2.8, tarsus 1.0. Second patella eiael milo) S45 saonen, anixel BOs amon B2. Length of first patella and tibia 1.3 times width of carapace. Male allotype. Carapace gray-orange Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 with a white median patch. Sternum or- ange. Legs orange, dusky dorsally. Abdo- men dusky orange, with humps. Total length 1.7 mm. Carapace 0.80 mm long, 0.68 wide in thoracic region, 0.48 wide at lateral eyes. First femur 0.67 mm, patella and tibia 0.78, metatarsus 0.41, tarsus 0.29. Second patella and tibia 0.65 mm, third 0.39, fourth 0.53. Length of first patella and tibia 1.1 times width of carapace. Note. Males were raised from the egg sac of the holotype. Variation. Total length of females 6.3— 10.5 mm. Males may lack humps. The il- lustrations were made from the female ho- lotype, except Figure 70 from a North Carolina specimen, and the male from the allotype. Diagnosis. Mastophora stowei is distin- guished from M. bisaccata by being small- er in size and having the atria of the epi- gynum ventrally departing from each other Fi igs. 68—70). The egg sac is shown in Fig- ure 447. Natural History. Females have been collected from trees along farm fences in Kentucky, and on a carpet of silk on a ma- ple leaf in Virginia. Dicom Widespread in the eastern United States (Map 2C). Paratypes. CONNECTICUT Hartford Co.: Rain- bow nr. Windsor, 9 Aug. 1939, 12 (A. de Caprio, USNM). OHIO Logan Co.: Old Man’s Cave, 12 Sep. 1924, 15 (OSU). VIRGINIA Arlington Co.: Arling- ton) 7 Septyl953) 121k Vv; Ker Omiein AMNH). IL- LINOIS Pope Co.: Dixon Spring State Park, 7 Sept. 1974, 12 (J. A. Beatty, JAB). KENTUCKY Fayette Co.: Lexington, Oct. 1997, 1°, egg sac (K. V. Yeargan, KVY); 6 Aug. 1998, 14d (K. Yeargan, KVY):; Cold Stream farm fence, 16 Oct. 1998, 32 (K. Yeargan, KVY). Jessamine Co.: 3 Aug. 1995, 75 (K. Yeargan, KVY). NORTH CAROLINA Haywood Co.: Canton, 12 (Holden, MCZ). GEORGIA 12 (MNHWN 210). FLORIDA Alachua Co.: Gainesville, male allotype and 1 penultimate paratype from egg sac of holotype, => Figures 100-106. Mastophora catarina new species, female. 100, 101, carapace and chelicerae. 100, frontal. 101, lateral. 102, 103, carapace and abdomen. 102, dorsal. 103, lateral. 104-106, epigynum. 104, ventral. 105, posterior. 106, posterior, cleared. Figures 107-113. MM. haywardi Birabén, female. 107, 108, carapace and chelicerae. 107, frontal. 108, lateral. 109, 110, carapace and abdomen. 109, dorsal. 110, lateral. 111-113, epigynum. 111, ventral. 112, posterior. 113, posterior, cleared. MASTOPHORA ° Levi 335 Figures 114-120. MM. corumbatai new species, female. 114, 115, carapace and chelicerae. 114, frontal. 115, lateral. 116, 117, carapace and abdomen. 116, dorsal. 117, lateral. 118-120, epigynum. 118, ventral. 119, posterior. 120, posterior, cleared. Figures 121-127. M. laranew species, female. 121, 122, carapace and chelicerae. 121, frontal. 122, lateral. 123, 124, carapace and abdomen. 123, dorsal. 124, lateral. 125-127, epigynum. 125, ventral. 126, posterior. 127, posterior, cleared. Scale lines. 1.0 mm; genitalia, 0.1 mm. 336 hatched 27 Apr. to 15 May 1988, preserved 18 July 1988 (M. Stowe 20181, 20183, MCZ, AMNH, FSCA). TEXAS Harrison Co.: Marshall, 5 July 1991, ES X(Se1G-aWellsos ke): Mastophora yeargani new species Plate 1; Figures 75-85; 448; Map 2A Holotype. Female holotype from Coldstream Farm fence, Lexington, Kentucky, 26 Oct. 1998, male al- lotype and 11 male and 4 female paratypes emerged 24 May 1999, preserved in fall (K. V. Yeargan), in MCZ. The species has been named after the collector, who has contributed much to our knowledge of Mastophora ecology. Description. Female holotype. Carapace light brown. Chelicerae, labium, endites yellow. Sternum yellow underlain by white pigment granules. Coxae and distal leg ar- ticles yellow. Abdomen anterior of dorsum gray with white marks anteriorly (Fig. 77), posterior white, venter whitish with white square. Carapace shiny. Abdomen with pair of very slight dorsal swellings. Total length 10.0 mm. Carapace 4.2 mm long, ALS) ride in thoracic region, 2.3 wide at lat- eral eyes. First femur 3.9 mm, patella and tibia 5.4, metatarsus 3.8, tarsus 1.2. Sec- ond patella and tibia 4.0 mm, third 2.4, fourth 3.7. Length of first patella and tibia 1.3 times width of carapace. Male allotype. Carapace brown with white triangle in center. Chelicerae, labi- um, endites orange. Sternum orange. Cox- ae and distal leg articles orange. Abdomen whitish with pair of humps. Total length 1.7 mm. Carapace 0.78 mm long, 0.78 wide in thoracic region, 0.52 wide at lat- eral eyes. First femur 0.87 mm, patella and tibia 0.88, metatarsus 0.52, tarsus 0.34. Second patella and tibia 0.75 mm, third 0.39, fourth 0.60. Length of first patella and tibia 1.1 times width of carapace. Note. Males have been raised from egg oo sac of female M. yeargani (by K. Yeargan). Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 Variation. Total length of females 10.0— 11.5 mm. The illustrations were made from the holotype and allotype. Diagnosis. Mastophora yeargani is dis- tinguished from M. bisaccata and M. stow- ei, which have similar abdominal markings, by the short slits of the epigynum, which are almost parallel and their length distant from the ventral margin (Figs. 80, 81). The female also lacks duskiness on the dorsum of the femora and gray pigment on the clypeus, both of which are present in M. stowei. The male differs from M. bisaccata (Fig. 62) by having a longer median apophysis (Fig. 85) and om M. cornigera (Figs. 177 1 79) by having only a short terminal apophysis in the palpus (Figs. 82, 83), and by having the median apophysis Ol anait= ferent shape, and the embolus wider (Figs. Soma): Distribution. From New York to Ken- tucky (Fig. 448). Paratypes. NEW YORK nr. New York City, on Amalanchier sp., 12 (AMNH). KENTUCKY Mercer Co.: Feb. 1995 egg sacs, 29 Sept. 1995, 7 imm., 4d (K. V. Yeargan, KVY). Garrard Co.: Feb. 1995, egg sacs, Sept. 1995, 6 imm., 16d (K. V. Yeargan, KVY). Mastophora phrynosoma Gertsch Figures 86-99, 449, 450; Map 2D Mastophora phrynosoma Gertsch, 1955: 245; pl. 6, fig. 5, text figs. 24-27, 31, 2. Female holotype from Burlington, North Carolina, in AMNH, examined. Brignoli, 1983: 273. Platnick, 2001. Description. Female holotype. Carapace orange-brown. Sternum orange-brown. Legs. orange-brown, indistinctly ringed. AGdewen anter iorly gray, posteriorly white (Fig. 88), venter with white square. Cara- pace with tubercles very small (Figs. 86, 87). Median eyes on bulge, lateral eyes on bulges. Abdomen subtriangular with a = Figures 128-134 Mastophora carpogastra Mello-Leitao, female. 128, 129, carapace and chelicerae. 128, frontal. 129, lateral. 130, 131, carapace and abdomen. 130, dorsal. 131, lateral. 132-134, epigynum. 132, ventral. 133, posterior. 134, posterior, cleared. Figures 135-141. M. seminole new species, female. 135, 136, carapace and chelicerae. 135, frontal. 136, lateral. 137, 138, carapace and abdomen. 137, dorsal. 138, lateral. 139-141, epigynum. 139, ventral. 140, posterior. 141, posterior, cleared. MASTOPHORA ® Levi 337 7 nor 5 ee ~ SF carpogastra Figures 142-152. M. vaquera Gertsch, female. 142, 143, carapace and chelicerae. 142, frontal. 143, lateral. 144, left first femur and patella, mesal. 145, 146, carapace and abdomen. 145, dorsal. 146, lateral. 147-152, epigynum. 147, 150 ventral. 148, 151, posterior. 149, 152, posterior, cleared. 147-149, (holotype from Matanzas). 150-152, (Santiago). Scale lines. 1.0 mm; genitalia, 0.1 mm. 338 swelling on each anterior lateral side and indistinct sclerotized discs (Fig. 88). Total length 12.3 mm. Carapace 4.6 mm long, 4.4 wide in thoracic region, 2.7 wide at lat- eral eyes. First femur 4.6 mm, patella and tibia 6.5, metatarsus 4.7, tarsus 1.3. Sec- ond patella and tibia 4.5 mm, third 2.4, fourth 4.1. Length of first patella and tibia 1.5 times width of carapace. Male from Kentucky. Carapace orange- brown, median triangle enclosing horns and two tubercles lighter. Sternum orange. Legs orange-brown. Abdomen both sides orange-white. Carapace slightly rugose, with two median tubercles and four pos- terior horns. Abdomen with small dorsal humps. Total length 1.8 mm. Carapace 0.81 mm long, 0. 80 wide in thoracic re- gion, 0.55 wide at lateral eyes. First femur 0. 91 mm, patella and tibia 0.93, metatarsus 0.46, tarsus 0.31. Second patella and tibia 0.78 mm, third 0.45, fourth 0.60. Length of first patella and tibia 1.2 times width of carapace. Note. Males have been raised by M. K. Stowe and K. V. Yeargan from egg sacs that were determined to ies from M. phryno- soma. Variation. Total length of females 8.3— 12.3 mm, males 1.5—1.7. The lateral swell- ings of a specimen from Falls Church are less distinct. The carapace of a female from Missouri is blackish brown, sternum and coxae black, legs ringed, aval abdom- inal venter black. The venter of several in- dividuals is black. The illustrations were made from the female holotype and from the reared male. Diagnosis. Mastophora phrynosoma is distinguished by the subtriangular shape of the Aidomien. the small, dorsal sclerotized discs (Fig. 88), and by the posterior of the epigynum having a lip surrounding the slits on three sides (Figs. 90-92). The palpus of the male has the space enclosed by the median apophysis wider than long and the prong straight (Figs. 96, 99), and has the base of the median apophysis almost as wide as long (Figs. 94, 98). It differs from that of M. archeri by Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 having a longer terminal apophysis (Figs. 93, 94), and wider radix (Figs. 95, 98). The egg sac is distinct, having longer flaps than in other species (Figs. 449, 450). Natural History. A female was collected on elm bush, 1.5 m high, in Texas; in Mis- souri, a female was collected hanging on a silk strand on canebrake (Arundinaria gi- gantea) at night at 0200 h. Males were col- lected crawling on a table in the labora- tory, and in low branches of hawthorn in Ohio. Immature spiderlings attract psy- chodid flies (Psychoda phalaenoides) (Yeargan and Quate, 1996). The female al- ways rests on the upper leaf surfaces, which often accumulate a silk pad that may be visible under the spider. The spi- ders look like bird droppings in Florida (M. Stowe, personal communication). The moths captured in Kentucky and Florida, observed by M. K. Stowe and K. V. Year- gan, were reported by Yeargan (1994). An egg sac raised by K. V. Yeargan pro- duced only male spiderlings. A suggestion has been made that the sex chromosome system might be different from that of oth- er spiders (G. Oxford, in letter). Perhaps this skewed sex ratio was due to the mor- tality of females. Distribution. (Map 2D). Eastern United States Specimens Examined. CONNECTICUT New Ha- ven Co.: Mount Carmel, 4 Sept. 1946, 12 (K. M. Somerman, INHS 4748). Meriden Co.: South Meri- den, Oct. 1945, 12 (H. L. Johnson, USNM). NEW YORK New York City, on Prunus, 12 (AMNH). OHIO Cuyahoga Co.: Sagmore Picnic, Sagmore Hills, Buckeye Trail area, 16 Sept. 1999, 12 (K. Brad- ley, OSU). MARYLAND Anne Arundel Co.: Annap- olis, 21 Sep. 1941, 12 (M. H. Muma, AMNH). How- ard Co.; Colombia, Snowden River Parkway, 26 Sep. 1994, 12 (M. Harden, USNM). Montgomery Co.: Cabin John, 10 Nov. 1943, egg sacs (I. N. Hoffman, USNM). VIRGINIA Falls Church, 22 (N. Banks, MCZ). KENTUCKY Fayette Co.: Lexington, Raven Run, 1989, 19? (K. V. Yeargan, KVY); 17 May 1999, egg sacs with only dd (K. V. Yeargan, KVY, MCZ). GEORGIA Fulton Co.: Atlanta, 21 Aug. 1944, 12 (F. W. Fattig, AMNH). FLORIDA Alachua Co.: Gaines- ville, Apr. 1988, 1¢ (M. K. Stowe, MKS); Devil's Millhopper State Park, 15 July 1980, 12 (M. K. Stowe, MKS); spring 1992, 2d (M. K. Stowe, MKS): spring 1994, 8d (M. K. Stowe, MKS). ALABAMA Monroe Co.: Randon’s Creek, 19 Oct. 1941, 12 (A. F. Archer, AMNH). INDIANA 12 (A. Petrunkevitch, NHMW). ILLINOIS Jackson Co.: Carbondale, in woods at night, 4 Aug. 1967, 12 (J. M. Nelson, AMNH). MISSOURI Wayne Co.: Markham Spring, Mark Twain National Forest, 14 Oct. 2000, 12 (E L. Quinter, AMNH). St. Louis Co.: 17 July 1940, 13 paratype of M. archeri (W. M. Gordon, AMNH). TEXAS Walker Co.: Huntsville State Park, on elm bush, 27 Sep. 1987, 12 (W. R. Martin, TAMU). Mastophora catarina new species Figures 100-106; Map 3C Holotype. Female from Pinhal, Est. Santa Catarina, Brazil, Dec. 1948 to Jan. 1949 (A. Maller), in AMNH. The specific name is a noun in apposition after the locality. Description. Female holotype. Carapace dark orange-brown (Fig. 102). Sternum dark orange. Chelicerae dark in front, or- ange on sides. Endites, labium, sternum, coxae orange; coxae lighter than sternum. Legs dark brown. Abdomen gray-brown, dorsally with three pairs of white longitu- dinal lines (Fig. 102); venter with a white square. Eyes distinct. Lateral eyes 0.8 di- ameter of median eyes. Abdomen without humps (Fig. 102). Carapace, legs, and ab- domen with some long white hair, Total length 7.2 mm. Carapace 3.3 mm long, 3.2 wie in thoracic region, 1.8 wide at tae al eyes. First femur 3.5 mm, patella and tibia 4.4, metatarsus 3.0, tarsus 1.0. Second pa- tella and tibia 3.3 mm, third 1.7, fourth 2.7. Length of first patella and tibia 1.4 times width of carapace. Males are not known. Diagnosis. Mastophora catarina is sep- arated from others by the abdomen lack- ing humps and having six white lines (Fig. 102), by the epigynum having the atria ap- proaching each other (Figs. 105, 106), and by the paired notches at the edge of the epigynum (Fig. 106). Distribution. Santa Catarina State, Bra- zil (Map 3C). Specimens Examined. No other specimens have been found. Mastophora haywardi Birabén Figures 107-113; Map 3C Mastophora haywardi Birabén, 1946: 327, figs. 1-3, 2. Female holotype, from Tucuman, Argentina, in MASTOPHORA ® Levi 339 MLP. examined. 2001. Brignoli, 1983: 274. Platnick, Description. Female holotype in poor condition. Carapace brown, with a lighter square area in thoracic region (Fig. 109), and with long white setae. Stemi lighter, Legs frown Abdomen brown (the Thole. type has an injury, a large transverse gash anteriorly), posterior and: sides darker gray than anterior (Fig. 109), with some long white setae and indistinct humps ate only from side; venter light. Total length 11.0 mm. Carapace 5.5 mm long, 4.7 wide in thoracic region, 3.1 wide in cephalic re- gion. First formate 4.7 mm, patella and tibia 7.6, metatarsus 5.8, tarsus 1.5. Second pa- tella and tibia 5.2 mm, third 3.0 (from Bir- abén, 1946), fourth 4.5. Length of first pa- tella and tibia 1.3 times width of carapace. Males are not known. Diagnosis. This species is separated from others by being the only setose spe- cies without distinct humps (Fig. 109), and by having the slits and the atria in the pos- terior view of the epigynum ventrally sep- arated (Figs. 112, 113). Distribution. Known only from Tucu- man, Argentina (Map 3C). Specimens Examined. No other specimens have been collected. Mastophora corumbatai new species Figures 114-120; Map 3C Holotype. Female holotype from Corumbataf, Est. Sao Paulo [40 km N Rio Claro], Brazil, 15 July 1935 (Syilvio Bariau), in IBSP no. 1203A. The spe- cific name is a noun in apposition after the locality. Description. Female holotype. Carapace red-brown with white rim (Fig. 116). Che- licerae, labium, endites, sternum, coxae, proximal ends of femora orange. Distal leg articles brown. Abdomen light brown, with colorless butterfly-shaped light area dor- sally and pair of large dark, open rings (Fig. 116); venter light brown with white square. Carapace with various sized, drop- let-shaped tubercles, sides with long white setae, some curled (Figs. 114, 115). Ab- domen without humps, with scattered 340 long, white setae (Fig. 116). Total length 14.0 mm. Carapace 5.6 mm long, 5.5 wide in thoracic region, 3.0 wide at lateral eyes. First femur 5.5 mm, patella and tibia 7.5, metatarsus 4.9, tarsus 1.6. Second patella and tibia 5.3 mm, third 3.2, fourth 4.8. Length of first patella and tibia 1.4 times width of carapace. Males are not known. Diagnosis. Mastophora corumbatai_ is distinguished by carapace tubercles that look like oil droplets (Figs. 114, 115), by lack of humps and distinctive color pattern on abdomen (Fig. 116), and by the epi- gynum with its posterior median plate pro- jecting, almost scapelike in ventral view (Figs. 118-120). Distribution. zil (Map 3C). Specimens Examined. No other specimens have been collected. Santa Catarina State, Bra- Mastophora lara new species Figures 121-127; Map 3A Holotype. Female holotype from Hato Arriba, 1,400 m, Lara, Venezuela, May 1970 (J. M. Osorio), in FSCA. The specific name is a noun in apposition after the locality. Note. The type locality Hato Arriba is probably at or near Quebrada Arriba, 1,600 m, 10°14'N, 70°32’W, close to the border with Falc6n, Zulia, and Lara, 52 km W Carora. Description. Female holotype. Carapace dark brown, lighter behind, in center and in area on sides. Chelicerae, labium, en- dites orange-brown. Sternum orange- brown. Coxae and distal leg articles or- ange-brown, distally darker. Abdomen gray with anterior dorsal pattern of spots and two large brown discs, each dissected by a lateral light line (Fig. 123); venter light brown aah white longitudinal rectangle. Carapace with tubercles and _ scattered long white setae (Figs. 121, 122). Legs aan some long white setae. Abdomen without humps, blunt behind, each side slightly swollen, long white setae on each side anteriorly (Fig. 123). Total length 11.5 mm. Carapace 5. 0 mm long, 5.0 wide in Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 thoracic region, 2.7 wide at lateral eyes. First femur 5.2 mm, patella and tibia 6.7, metatarsus 4.5, tarsus 1.3. Second patella aul wwlore, SJ waren, dette! G0), toorda 2h), Length of first patella and tibia 1.3 times width of carapace. Males are not known. Diagnosis. Mastophora lara differs by the high almost conical carapace (Figs. 121, 122), by the pattern on the abdomen (Fig. 123), and by the posterior of the epi- gynum having a pair of depressions (Figs. IQ). Distribution. This species is known only from the type locality in west-central Ve- nezuela (Map 3A). Specimens Examined. No other specimens have been collected. Mastophora carpogastra Mello-Leitao Figures 128-134, 451; Map 3B Mastophora carpogastra Mello-Leitao, 1925: 460. Two female syntypes from Rio de Janeiro, Brazil, in MNE 672, examined. Mello-Leitaio, 1931: 72, na Gh, IS, De Ghee an fagoides Vellard, 1926: 327, figs., 2, egg sac. Female holotype from Butantan, Sao Pau- lo, Brazil, in IBSP, lost. First synonymized with car- pogastra by Mello-Leitaéo (1931). Mastophora carpogastera:—Roewer, 1942: 900. Glyptocranium carpogastrum:—Bonnet, 1957: 1996. Mastophora carpogaster:—Platnick, 1993: 447. Plat- nick, 2001. Note. Roewer (1942) changed the name carpogastera and listed fagoides as a syn- onym of M. corpulenta Banks. The name change carpogaster of Platnick (1993, 2001) is not needed because previous re- visors (Mello-Leitao, 1931; Gertsch, 1955) kept the original spelling. Description. Female from Sao Paulo. Carapace reddish brown. Chelicerae, la- bium, endites brown. Sternum light brown. Legs brown. Abdomen white with symmetrical brownish black patches and less distinct lines dorsally (Fig. 130); ven- ter with a median white square containing eight black spots, sides white with black spots. Carapace glossy above, sides with long white setae, some setae on clypeus (Fig. 129). Legs with long white setae. Ab- domen oval with scattered setae and barely visible pair of dorsal tubercles (Fig. 131). Total length 16.5 mm. Carapace 6.4 mm long, 6.0 wide in thoracic region, 3.3 wide behind posterior lateral eyes. First femur 5.3 mm, patella and tibia 7.6, metatarsus 5.7, tarsus 1.6. Second patella and tibia 5.5 mim, third 3.3, fourth 5.2. Length of first patella and tibia 1.3 times width of cara- pace. Males are not known. Variation. Mello-Leitao (1925) de- scribed the species as orange to raspberry- red when alive. Total length of females 11.2-18.0 mm. The female holotype of M. carpogastra is 20 mm total length. The il- lustrations were made of specimens from Sao Paulo. Diagnosis. The spots and black lines and lack of humps of the abdomen (Fig. 130) distinguish the species from all oilers: In the cleared epigynum, the atria separate ventrally (Figs. 133, 134). The egg sac elke flaps and has a short stalk (Fig. 451). Natural History. The spider mimics a berry. According to Vellard (1926) the spe- cies has a preference for orange trees. It makes three to five egg sacs, ach a little sphere with four white spots below the midline, 10 mm in diameter, hanging on a stalk about 3 mm long. The venom is not active (for mammals?). From the collec- tions available, M. carpogastra seems lo- cally more common than other Mastopho- ra species. Distribution. Southeastern Brazil, from Rio de Janeiro State to Rio Grande do Sul (Map 3B). Specimens Examined. BRAZIL Bahra [?Bahia], 1° (MNHN 18669). Rio de Janeiro: Rio de Janeiro, 2° (NHMW); Aug. 1937, 1 imm. (Mello-Leitao, MACN 515). Sdo Paulo: Agua da Figueira Maracai, 9 Feb. 1967, 12 (G. Brisolla, IBSP 2064); Barueri, Apr. 1963, 1°, 5 egg sacs (K. Lenko, MZSP 3069); 16 Mar. 1966, 1 imm. (K. Lenko, MZSP 5265); Diadema, June 1986, 12 (R. Snignani, IBSP 4992); Embu, Sept. 1982, 12, egg sac (A. L. Prestes, IBSP 3501); Jandira, May 1980, 1? (C. Luiz, IBSP 963); Osasco, 25 Mar. 1974, 12 (F. Ramirez, IBSP 2721); Pacaem- bu, Nov. 1942, 12 (Braudas, MZSP 357): Perdizes, MASTOPHORA ° Levi 341 26 Aug. 1951, 12 (H. Camargo, MZSP 7496); Ribei- rao Pires, May 1975, 12 (F. B. Lopes, IBSP 3591); Rio Claro, July 1941, 1° (P. Pereira, MZSP 4552); 4 May 1942, 1 imm. (Clareteano, MZSP 4402); Saco- ma, 7 Sep. 1943, 1 (J. Lima, MZSP 4403); Sao Pau- lo, July 1921, 12 (MZSP 8068); Feb. 1928, 1° GE Lima, MZSP 8070); 31 Jan. 1934, 12 (M. Oliveira, IBSP 1920); 26 Bebe 1936, 12 (S. Remetente, IBSP 3598): 3 Oct. 1951, 12 (R. Vieira, IBSP 586); 17 Mar. 1955, 12, 2 egg sacs (J. Navas, IBSP 1176); June 1960, 22 (J. London, IBSP 1552); June 1960, 1° (L Zodiygansky, IBSP 1516); 21 IDS 1960, 12 (W. Ge drade, IBSP 1618), 10 July 1962, 12 (R. R. Guidug- lin, IBSP 1775); 21 Dec. 1961, 12 (F. V. Boas, IBSP 1620); 18 July 1962, 12 (E. Botelhiat IBSP 1776): Apr. 1963, 12 (Merck Co., IBSP 1834); 2 Aug. foeee 1¢@ (S. Remetente, IBSP 1949): 16 Dec. 1971, 19 a Rafael de Simone, IBSP 309); June 1975, 12 (J. S Gomes, IBSP 3594): Feb. 1976, 12 (M. Uchiyama, IBSP 3597): July 1975, 12 (G. P. Treu, IBSP 3595); Feb. 1976, 12 (E. I. Yamane, IBSP 17789): Feb. 1982, 12, egg sac (D. Zammataro, IBSP 3025); 26 eb. 1982, 1, egg sac (M. C. Franco, IBSP 14279); 2 Jan. 1986, 1° Gn Steiner, IBSP 8471); 21 Mar. Aa 12 (D. R. Bizzachi, IBSP 14424): 1 Feb. 1991, (J. Batista, IBSP 14398): Oct. 1992, 12 (C. M Nerici, IBSP 5827); 7 Oct. 1996, 12 (A. Fallatti, IBSP 14192); 11 Aug. 1997, 12 (S. M. Carnelho, IBSP 14012); 18 Mar. 1998, 12 (A. Pastore, IBSP 16208): Brooklin, Feb. 1962, 12 (L. Travassos, MZSP 4347); Hato Museu Paulista, Feb. 1951, 12 (C. Ra- bello, MZSP 6609); Ipiranga, Nov. 1906, Oct. 1912, 2@ (Ihering, L. M. Torre, MZSP 3047); Mar. 1924 (J. Lima, MZSP 3048); 15 Mar. 1961, 12, 5 egg sacs (Almeida and Cautero, MZSP 4359); 4 May 1961, 19, 2 egg sacs (N. C. Oliveira, MZSP 4340); 23 Jan. 1984, 1 imm. (C. R. F. Brandéo, MZSP 435); Magi das Cruzes, Ranch das Carmelitas, June 1976, 12 (C. Torrus, S. Filho, MZSP 11433). Santa Catarina: Ca- ceador, 1982, 12 (D. Lorenzato, IBSP 3539). Rio Grande do Sul: Ee Alegre, 12, 1 egg sac (P. Buck, MNRJ 1831); 7 July 1986. 12 (S. Oresco, MCN 15236): S. Leopoldo, 24 May 1964, 12 (C. Valle, MZSP 4233): 14 Oct. 1965, 12 (C. Valle, MZSP 5422). Mastophora seminole new species Figures 135-141; Map 2G Holotype. Female holotype from Hollendale, Bro- ward Co., Florida, 11 June 1987 (W. Birch), in FSCA. The specific name is a noun in apposition after the name of the local Indian tribe. Description. Female holotype. Carapace olive-brown. Chelicerae, labium, endites olive-brown. Sternum orange-olive. Coxae and distal leg articles olive, distally darkest. Abdomen anteriorly gray, posteriorly light gray with dark pattern on each side (Fig. 342 137); in anterior view, dark area forming a triangle as wide as humps above, pointed to carapace below, each side white. Venter olive gray with white longitudinal rectan- gle. Carapace glossy and with tubercles (Figs. 135, 136). Abdomen with a pair of dorsal humps and triangular (Figs. 137, 138). Total length 11.6 mm. Carapace 4.4 mm long, 4.3 wide in thoracic region, 2.7 wide at posterior lateral eyes. First femur 3.8 mm, patella and tibia 5.2, metatarsus 3.7, tarsus 1.2. Second patella and tibia 3.8 mim, third 2.2, fourth 3.3. Length of first patella and tibia 1.2 times width of cara- pace. Males are not known. Diagnosis. Mastophora seminole has a humped, somewhat triangular abdomen (Figs. 137); the epigynum is as in the humpless M. yeargani, with the slits dor- sally at some distance from the ventral borders of large shallow depressions (Figs. 140, 141). Distribution. Southern Florida (Map 2G). Specimens Examined. No other specimens have been collected. Mastophora vaquera Gertsch Figures 142-152, 452: Map 2G Mastophora vaquera Gertsch, 1955: 240: figs. 15-18, 2. Female holotype from Torriente, Matanzas, Cuba, in AMNH, examined. Brignoli, 1983: 274. Platnick, 2001. Description. Female holotype. Carapace orange-brown, bald dorsally with some white setae on sides. Sternum, legs or- ange-brown. Median eyes on bulge, ister al eyes on bulges. Abdomen light orange- brown with darker area anteriorly beeecen swellings (Fig. 145); venter with white square. Abdomen with humps and an an- terior, lateral swelling on each side (Figs. 145, 146). First femur with distal, anteri- orly small tubercles (Fig. 144). Total length 10.5 mm. Carapace. 3.6 mm long, B35) paide in thoracic region, 2.3 wide at lat- eral eyes. First femur 3 3 .2 mm, patella and tibia 4.7, metatarsus 3.2, tarsus 1.1. Sec- ond patella and tibia 3.4 mm, third 2.0, Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 fourth 3.1. Length of first patella and tibia 1.3 times width of carapace. Males are not known. Variation. Total length of females 8.5— 10.5 mm. The tubercles on the femur of the Cuabitas specimens are smaller, less distinct. Although the shape of the abdo- men and their coloration are similar, the epigyna of the two specimens differ (Figs. 147-152). The illustrations (Figs. 142— 149) were made from the female holotype. Diagnosis. Mastophora vaquera is dis- tinguished from others by the shape of the abdomen, humps, and two pairs of swell- ings, and by the dorsal oval depressions of ae opening slits on the posterior of the epigynum (Figs. 148, 151). The egg sac has only minute flaps (Fig. 452). Distribution. Cuba (Map 2G). Specimens Examined. CUBA Santiago: Cuabitas, Oriente [20°04’N, 75°48’W], 20 Aug. 1949, 12 (AMNH). Holquin: Bafies, 1-3 Aug. 1955, egg sac (A. F. Archer, AMNH). Mastophora hutchinsoni Gertsch Figures 153-168, 453, 454; Map 2E Cyrtarachne cornigera:—McCook, 1890: 98, 99, fig. 81, egg sacs. Kaston, 1948: 231, figs. 741, 742, 2039 (misidentification). Mastophora hutchinsoni Gertsch, 1955: 236, pl. 6, fig. 3, text figs. 10-14, 39, 47, 48, 2, 5. Female holo- type, from Somers [Westchester Co.], New York State, in AMNH, not examined. Brignoli, 1983: 273. Yeargan, 1988: 524. Gemeno et al., 2000: 1235. Haynes et al., 1996: 76. Platnick, 2001. Description. Female from Virginia. Car- apace brown, sides and posterior much darker than median and cephalic region, short setae on sides and no rim. Abdomen with transverse black band with character- istic light lines (Fig. 155); venter black in- cluding behind spinnerets and anterior of pedicel, with four white spots on each side between epigynum and spinnerets (Figs. 157). Abdomen with two humps (Figs. 155, 156). Total length 7.3 mm. Carapace 3.3 mm long, 2.7 wide in thoracic region, 2.3 wide at lateral eyes. First femur 2.5 mm, patella and tibia 3.6, metatarsus 2.8, tarsus 0.8. Second patella and tibia 2.7 MASTOPHORA ® Levi 343 2 cen S 3 Figures 153-168. Mastophora hutchinsoni Gertsch. 153-160, female. 153, 154, carapace and chelicerae. 153, frontal. 154, lateral. 155, 156, carapace and abdomen. 155, dorsal, with male. 156, lateral. 157, abdomen, ventral. 158-160, epigynum. 158, ventral. 159, posterior. 160, posterior, cleared. 161-168, male left palpus, stained. 161-164, (New Hampshire). 165-168, (Ken- tucky). 161, 165, apical. 162, 166, mesal. 163, 167, ventral. 164, 168, ectal. Figures 169-182. M. cornigera (Hentz). 169-175, female. 169, 170, carapace and chelicerae. 169, frontal. 170, lateral. 171, 172, carapace and abdomen. 171, dorsal, with male. 172, lateral. 173-175, epigynum. 173, ventral. 174, posterior. 175, posterior, cleared. 176-182, male left palpus, stained. 176-179, (California). 180-182, (Texas). 176, apical. 177, 180, mesal. 178, 181, ventral. 179, 182, ectal. Scale lines. 1.0 mm; genitalia, 0.1 mm. 344 mm, third 1.6, fourth 2.4. Length of first patella and tibia 1.3 times width of cara- pace. Male from New Hampshire. Prosoma beige with two median tubercles and forks. Abdomen whitish with paired humps. To- tal length 1.7 mm. Carapace 0.86 mm long, 0.71 wide in thoracic region, 0.53 wide at lateral eyes. First femur 0.80 mm, patella and tibia 0.91, metatarsus 0.41, tar- sus 0.27. Second patella and tibia 0.80 min, third 0.42, fourth 0.63. Length of first patella and tibia 1.3 times the width of car- apace. Note. The male was matched with fe- males because of its northerly collecting site. They have also been collected at sites for females in Kentucky and South Caro- lina. Variation. Total length of females 6.2— 10.4 mm. The males may lack humps. The illustrations were made from a female from Rhode Island, the male from south- ern New Hampshire, and a male from Kentucky. Diagnosis. The humps of the abdomen (Fig. 155) and the black median coloration of the venter, sometimes containing two longitudinal white patches (Fig. 157), sep- arate the species from M. bisaccata. Un- like M. cornigera, which has humps, the slits on the posterior of the epigynum ap- proach each other ventrally (Figs. 158, 160), as they do in M. bisaccata. The palpus (Figs. 161-168) is similar to that of M. bisaccata (Figs. 59-62), with a narrow space surrounded by the median apophysis in ectal view (Figs. 164, 168) but differs in the shape of the round base of the median apophysis (Figs. 162, 166). The egg sacs are unique compar ed with those of aliver species. They are attached by the broad base with the funnel facing away from the attachment (Figs. 153, 154). Natural History. In Ohio, females were found in a house yard on a variety of low branches of a variety of trees, including crab apple, redbud, hawthorn, burr oak (R. Bradley, in letter). Spiders were col- lected primarily from hackberry (Celtis oc- Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 cidentalis L.) and wild cherry (Prunus ser- otina Ehrgard) in Kentucky (Yeargan, 1988), on an abandoned apple tree in Vir- ginia, in a peach orchard in South Caroli- na, and in an apple orchard in Illinois. Prey caught in Kentucky were reported by Yeargan and Quate (1996). Although the egg sacs look like hazel- nuts or a Small broken branch, the species may be found by searching for the egg sacs in autumn and winter Ate the leaves have fallen. Distribution. Northeastern United States from New Hampshire and South Carolina to Minnesota (Map 2E). Specimens Examined. NEW HAMPSHIRE Hills- borough Co.: Ponemah [in Amherst], Aug. 1912, 1d (E. B. Bryant, MCZ). MASSACHUSETTS 1 (San- born, MCZ). Middlesex Co.: Pepperell, 15 July 1978, limm. ¢, 1.5 mm (F. J. Murphy, JM). RHODE IS- LAND Providence, 12 (N. Banks, MCZ). CON- NECTICUT Hartford Co.: Rocky Hill, 23 Oct. 1940, 12 (A. Morgan, USNM). NEW JERSEY Burlington Co.: Riverton, Sept. 1926, 2 imm., 1d (R. J. Sim, OSU). Hunterdon Co.: White House Station, Sept. 1917, 12 (J. J. Brochon, USNM). Morris Co.: Dover, 8 Dec. 1950, egg sacs (USNM). OHIO Delaware Co.: 5 km W of Deen Dam, 40.37°N, 83.10°W, Oct. 2001, 22, egg sacs (R. Bradley, OSU). DISTRICT OF COLOMBIA Washington, yard, 18 Oct. 1920, 12 (USNM). VIRGINIA Falls Church, 52, 2 imm. (N. Banks, MCZ). Augusta Co.: Augusta, 14 June 1976, 13; 6 Oct. 1976, 12 ( (J. P. McCaffrey, CNC). SOUTH CAROLINA Anderson Co.: Simpson Agric. Exp. Sta- tion; LOT July 197385 di 13 eAug 19708 ho SON Seps 1978, 12 (G. Lee, CUAC). KENTUCKY Fayette Co.: Lexington, 7 Sep. 1990, 12 (K. V. Yeargan, MCZ); 20 Sep. 1990, 19 (K. V. Yeargan, KVY); 2 Oct. 1999, 1° (K. V. Yeargan, MCZ). Clark Co.: Feb. 1996, egg sac, 43 (K. V. Yeargan, KVY). TENNESSEE Ashburn, 30 mi. N. Nashville, 17 July 1933, 1d (W. Ivie, AMNH). MICHIGAN Livingston Co.: E. S. George Reserve, 22 July 1951, 1 imm. (H. K. Wallace, FSCA). IN- DIANA Putnam Co.: Greencastle, 12 (N. Banks, MCZ). ILLINOIS Champaign Co.: Univ. Illinois ap- ple orchard, Sept. 1993, 12 (S. D. Gaimari, INHS). Jackson Co.: 8 km S Carbondale on avocado plant, Oct. 1976, 12 (JAB). MINNESOTA Hennepin Co.: Minneapolis, 1 Nov. 1931, 12 (W. J. Gertsch, AMNH). Mastophora cornigera (Hentz) Figures 169-182, 455; Map 2G Epeira cornigera Hentz, 1850: 20, pl. 3, fig. 8, &. Immature female holotype from Alabama, de- stroyed. Hentz, 1875: 123, pl. 14, fig. 8, @. Cyrtarachne bicurvata Becker, 1879: 77, pl. 2, figs. 16-19, 2. Female holotype from peach tree, Don- aldsonville, Louisiana, in IRSNB, examined. First synonymized by Marx (1890). Cyrtarachne cornigera:—Keyserling, 1880: 300, pl. 4, fig 4502 MeCocle 1890: 98. fig. “80 (in part). Ontlauntns cornigerus:—Marx, 1890: 541. McCook, 1894: 197, pl. 12, fig. I, @. Glyptocranium cornigerum:—Simon, 1895: 882, 885. Bonnet, 1957: 1996. Mastophora cornigera:—Mello-Leitao, 1931: 70, figs. 9, 20. Gertsch, 1955: 233, pl. 6, fig. 2, text figs. 1-5, 37, 41, 42, 2, 3. Platnick, 1997: 513. Platnick, 2001. Description. Female from Alabama. Carapace evenly colored orange. Chelic- erae, endites, labium orange. Sternum or- ange. Legs orange, darker above. Abdo- men with black caps on humps, anterior gray with light lines, posterior light (Fig. 171); venter with a white square. Carapace granular with many small tubercles and dark spots on sides (Figs. 169, 170). Ab- domen with pair of humps (Fig. 171). To- tal length 12.0 mm. Carapace 5.6 mm long, 4.6 wide in thoracic region, 3.0 wide at lateral eyes. First femur 4.2 mm, patella and tibia 6.0, metatarsus 4.0, tarsus 1.2. Second patella and tibia 4.7 mm, third 2.6, fourth 4.3. Length of first patella and tibia 1.3 times width of carapace. Male from California. Carapace beige with median dorsal white band including median horns, no tubercles. Sternum, legs golden yellow. Abdomen white, anteriorly dusky; venter dark yellow. Carapace gran- ulate. Total length 1.7 mm. Carapace 0.88 mm long, 0.79 wide in thoracic region, 0.52 wide behind posterior lateral eyes. First femur 0.66 mm, patella and tibia 0.79, metatarsus 0.40, tarsus 0.28. Second patella and tibia 0.68 mm, third 0.41, fourth 0.54. Length of first patella and tib- ia 1.1 times width of carapace. Note. Males came from California, an area from which only one species of Mas- tophora, M. cornigera, is known. Variation. Total length of females §.8— 14.0 mm, males 1.6—1.7. The illustrations were made from the female from Alabama (Figs. 169-175) and males from California MASTOPHORA ° Levi 345 (Figs. 176-179) and Texas (Figs. 180-182). Adult males may lack humps. Diagnosis. Unlike most other North American species, M. cornigera has dis- tinct humps on the abdomen, often with a black cap or slightly sclerotized (Fig. 173), and the epigynum differs from that of M. hutchinsoni (Figs. 159, 160) by having al- most parallel slits, only slightly converging ventrally, on the posterior face of the epi- gynum (Figs. 174, 175), and by lacking the lip surrounding the slits as in M. archeri (Fig. 188). The male differs from other North American species by having the space within the curl of the median apophysis in ectal view wider than long (Figs. 179, 182), and from M. yeargani by the base of the embolus, which, in mesal view, is longer than wide (Figs. 177, 180). The egg sac has small flaps or none and a relatively wide stalk (Fig. 455). Natural History. Unlike other North American species, M. cornigera is active in California all year. Also, unlike other North American species, the males emerge from the egg sac as mature indi- viduals. This species was found on cycad leaf in full sun in San Diego, and on Jat- ropha curcas euphorbia in Nicaragua. Distribution. From Kentucky and Ten- nesse west to California and south to Cen- tral America (Map 2G). Specimens Examined. KENTUCKY Fayette Co.: Univ. Kentucky Maine Chance farm, Sept. 1996, 1° (K. V. Yeargan, KVY). TENNESSEE Ashburn, 30 mi. N Nashville, 17 July 1933, 12 (W. Ivie, AMNH). AL- ABAMA Mobile Co.: Mobile, 12 (N. Banks, MCZ):; 1932, 12 (H. PB. Loding, MCZ). LOUISIANA East Baton Rouge Par.: Baton Rouge, Apr. 1916, 12 (New- ell, MCZ). Orleans Par: New Orleans, 1918, 12 (H E. Hubert, USNM); 1 Oct. 1935, 12 (T. E. Snyder, MCZ); 26 Sep. 1936, 12 (J. N. Cowanloch, USNM). TEXAS Travis Co.: Shellberg Tract, 30°25'N, 97°52’W, 18-19 Apr. 1994, 3d (Dunlap et al., TAMU). Galveston Co.: Texas City, 1921?, 2° (S. W. Bilsing, MCZ). San Patricio Co.: Welder Wildlife Refuge, 11.8 km NE Sinton, 17 Oct. 1967, 1d (C Parrish, CAS). Hidalgo Co.: Edinburg, Oct. 1934, 39, 1d; 1935, 12 (S. Minleie AMNH): 7 Dec. 1935, 14 (M. Welch, AMNH), 2 imm., 16 (S. Mulaik, AMNH): 18 km SE Pharr, Santa Ana Wildlife Ref- uge, 1 Oct. 1977, 12 (O. Ahrenholtz, AMNH); 20 346 Deer 983n ll 2a(M Stowe, MKS); 27 June 1984, 12 (M. K. Stowe, Anes ee 10 Apr. 1986, egg sac, 70 imm., 596 (D. A. Dean, TAMU). Cam- eron Co.: Brownsville, 9 Apr. 1986, hatched 2 June 1986, 1 egg sac, 64 imm., 62d (D. A. Dean, TAMU); Harlingen, Feb. 1980, 12, 64 imm., 63d (C. W. Ag- new et al., TAMU); E of Harlingen, 3 Jan. 1936, 12 (L. I. Davis, M. Stegmeier, AMNH). ARIZONA Mar- icopa Co.: Phoenix, Apr. 1941, 12 (AMNH). CALI- FORNIA Contra Costa Co.: Walnut Creek, Sydney Drive, July 1992, 12, 1d (T. Trosin, J. Fraser, CAS). Santa Clara Co.: Palo Alto, 1914, 12 (H. Heath, MCZ). Santa Barbara Co.: Santa Barbara, 5 Oct. 1948, 12 (H. Shantz, AMNH). Los Angeles Co.: Claremont, 12 (N. Banks, MCZ); Glendale, 1°, many 6 (C. E. Hutchinson, AMNH, MCZ); Los An- geles, 12 (MNHN, 3059): 10 Oct. 1942, 5d Qe EL: Branch, AMNH); East Los Angeles, 1943, 12 (C. Cowles, AMNH); Malibu, Nov. 1968, 12, 3 egg sacs (USNM); Westwood Village, Aug.—Oct. 1942, 12 (P. Verrity et al., FSCA). Orange Co.: Santa Ana, 4d (R K. Bishop, USNM); San Juan Capistrano, 25 Sept. 1952, 1?, 7 egg sacs (R. E. Ryckman, AMNH). San Bernardino Co.: San Bernardino, 1880, 12 (J. B. Par- ish, MCZ). San Diego Co.: San Diego, 22 (USNM), 4 Oct. 1974, 1° (D. Bishop, USNM); San Diego, Vista, 8 June 1989, | imm. (J. W. Schott, MCZ); Chu- la Vista, 5 Dec. 1981, 12 (H. V. Weems, FSDA): 10 mi. NE Ramona, 22 July 1982, 19 (J. Halstead, DU); Lakeside, 12 (C. Kingery, USNM); Feb. May 1968, egg sacs, dd (C. Kingery, USNM). MEXICO Baja California Sur: 44 km W La Paz, 0.2 km S km 44, on Highway 1, 31 Dec. 1978, 12 (D. Weissman, R. Love et al., CAS). NICARAGUA Managua: Mateare, 12 Sep. 1995, 12 (C. Grimm, M. Maes, MCZ). Mastophora archeri Gertsch Figures 183-193, 456; Map 2E Mastophora archeri Gertsch, 1955: 239: figs. 6-9, 36, 45, 46, 2, d. Female holotype from Fruitland Park, Florida, in AMNH, examined. Brignoli, 1983: 273. Platnick, 2001. Description. Female holotype. Carapace orange-brown. Sternum dark orange. Legs orange-brown, indistinctly ringed. Abdo- men ‘anteriorly gray, posteriorly \ white; ven- ter with white square. Carapace with short tubercles (Figs. 183, 184). Median eyes on bulge, later al eyes on bulges. Abdomen mn small humps (Fig. 185). Total length 11.5 mm. Carapace 4.2 mm long, 4.3 wide in thoracic region, 2.7 wide at lateral eyes. First femur 3.8 mm, patella and tibia 5.6, [metatarsus 3.5, tarsus 1.5, after Gertsch, 1955]. Second patella and tibia 4.1 mm, Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 third 2.3, fourth 3.7. Length of first patella and tibia 1.3 times width of carapace. Male allotype. Carapace orange, darkest on sides, a median white line and median of forked tubercles white. Sternum or- ange. Legs orange. Abdomen orange- white without marks. Carapace with two small asymmetrical tubercles in addition to forked tubercles. Abdomen with indistinct humps. Total length 1.7 mm. Carapace 0.86 mm long, 0.78 wide in thoracic re- gion, 0.53 wide at lateral eyes. First femur 0.79 mm, patella and tibia 0.87, metatarsus 0.44, tarsus 0.33. Second patella and tibia 0.74 mm, third 0.54, fourth 0.71. Length of first patella and tibia 1.1 times width of carapace. Note. A male from Gainesville, Florida, was raised from the egg sac. The match of the male allotype is uncertain. Variation. Total length of females 9.4— 14.8 mm. The illustrations were made from the female holotype (Figs. 183-189) and the raised male from Gainesville (Figs. 190-193). Diagnosis. Mastophora archeri is distin- guished from M. cornigera by the smaller tubercles on the carapace (Figs. 183, 184) and left and right lip on the posterior of the epigynum (Figs. 188, 189), and from M. hutchinsoni by having a white square on the venter and by the sculpturing of the epigynum. The palpus of the male has the space surrounded by the median apophysis wid- er than long (Fig. 193) and differs from M. phrynosoma and M. hutchinsoni by having only a minute terminal apophysis (Figs. 190, 191). It differs from M. phry- nosoma by the narrow radix (Fig. 191). The egg sac has small flaps and a rela- tively long stalk of median thickness (Fig. 456). Natural History. Collected from Myrica in hammock woods in Alabama. Distribution. Southern United States from South Carolina, Florida, and Ala- bama to Kansas (Map 2E). Paratypes. ALABAMA Baldwin Co.: Lagoon, 29 MASTOPHORA ® Levi 347 Figures 183-193. Mastophora archeri Gertsch. 183-189, female. 183, 184, carapace and chelicerae. 183, frontal. 184, lateral. 185, 186, carapace and abdomen. 185, dorsal, with male. 186, lateral. 187-189, epigynum. 187, ventral. 188, posterior. 189, posterior, cleared. 190-193, male left palpus, stained. 190, apical. 191, mesal. 192, ventral. 193, ectal. Figures 194-204. WM. fasciata Reimoser. 194-200, female. 194, 195, carapace and chelicerae. 194, frontal. 195, lateral. 196, 197, carapace and abdomen. 196, dorsal, with male. 197, lateral. 198-200, epigynum. 198, ventral. 199, posterior. 200, posterior, cleared. 201—204, male left palpus, stained. 201, apical. 202, mesal. 203, ventral. 204, ectal. Scale lines. 1.0 mm; genitalia, 0.1 mm. 345 Sep. 1949, 1d allotype (A. F. Archer). KANSAS Douglas Co.: 22 Sept. 1948, 12 (R. H. Beaner, AMNBE). Specimens Examined. SOUTH CAROLINA Charleston, Oct. 1941, 12 (E. B. Chamberlain, USNM). FLORIDA Alachua Co.: Devil’s Millhopper State Park, reared spring 1992, 4d (M. K. Stowe, MKS). Hillsborough Co.: Thonotosassa, 11 Dec. 1969, 1? (D. A. Vaughn, FSCA). ALABAMA Mobile Co.: Mobile, 12 Nov. 1939, 12 (A. C. Cole, AMNH); Mt. Vernon, Oct. 1941, 1? (H. P. Léding, AMNH). Mastophora fasciata Reimoser Plate 1; Figures 194-204, 457; Maps 2F, 3A Mastophora fasciata Reimoser, 1940: 356, fig. 8, &. Female holotype, from Orosi, Proy. Cartago, cen- tral plain [12 km SE Cartago], Costa Rica, in NMW, examined. Roewer, 1942: 900. Platnick, 2001. Mastophora pickeli occidentalis Schenkel, 1953: 29. Female holotype from Poz6n [Falcén], Venezuela, in NMB, examined. Brignoli, 1983: 274. Platnick, 2001. Note. Mastophora p. occidentalis has the same small lobe on the posterior margin of the epigynum, and the same shadows on the posterior face as in M. fasciata. They also share the very broad humps of the abdomen and the posterior light band. Description. Female from Costa Rica. Carapace light orange-brown. Sternum brown. Legs lighter brown. Abdomen white with Tblaek and gray marks (Fig. 196); venter light brown, center barely lighter than sides. (Reimoser described a posterior, transverse, yellow-red band, which has disappeared from the holotype, but is light in other specimens.) Humps broad (Fig. 196). Holotype total length ILL x5) Lacon, Carapace 4.7 mm long, 5.0 wile in thoracic region, 3.0 wide at lateral eyes. First femur 4.3 mm, patella and tibia 6.3, metatarsus 4.4, tarsus 1.2. Second patella andistibial 4:6. mms) thing) 247. stountia 4.0) Length of first patella and tibia 1.3 times width of carapace. Bulletin Museum of Comparative Zoology ; Vol. 157, No.5 Male from Costa Rica. Carapace yellow- brown with white, central mark covering median tubercles. Legs light yellow- brown. Abdomen dorsally white, ventrally yellow-brown. Eyes without pigment. Car- apace rugose with posterior median forked tubercles. Abdomen subtriangular without humps. Palpal patella with one macroseta. Total length 1.6 mm. Carapace 0.74 mm long, 0.65 wide in thoracic region, 0.46 wide behind posterior lateral eyes. First femur 0.55 mm, patella and tibia 0.65, metatarsus 0.35, tarsus 0.26. Second pa- tella and tibia 0.59 mm, third 0.28, fourth 0.48. First patella and tibia as long as width of carapace. Note. Males were collected with females at San Antonio de Escazi. Variation. Total length of females 11.5— 14.5 mm. Both males and shriveled fe- males have a triangular abdomen, whereas that of a well-fed female is more rounded. The illustrations were made from the fe- male holotype (Figs. 194-200) with a spec- imen from Puntarenas Province, Costa Rica. Diagnosis. The female is distinguished from other species by the many small car- apace tubercles (Figs. 194, 195), by the broad humps of the abdomen (Fig. 196), and by the ventrally converging slits of the epigynum (Figs. 199, 200). The male has a median apophysis that is longer than that of other species (Figs. 202: 04) and more rounded than that of M. leucabulba (Figs. 293, 295) and M. al- vareztoroi (Fig. 306, 307). The median apophysis is almost as long as the diameter of the bulb (Fig. 204). The egg sac lacks flaps (Fig. 457). Natural History. Males are mature when they leave the egg sac. Distribution. The species is known from Costa Rica and Venezuela (Maps 2F, 3A). —>) Figures 205-215. Mastophora dizzydeani Eberhard. 205-211, female. 205, 206, carapace and chelicerae. 205, frontal. 206, lateral. 207, 208, carapace and abdomen. 207, dorsal, with male. 208, lateral. 209-211, epigynum. 209, ventral. 210, posterior. 211, posterior, cleared. 212-215, male left palpus, stained. 212, apical. 213, mesal. 214, ventral. 215, ectal. MASTOPHORA ® Levi 349 pickeli Wye jy? Maa ig i) CAIpr Ay yr, ) Uf) WE? We PG, d i yy 04 Figures 216-222. M. pickeli Mello-Leitao, female. 216, 217, carapace and chelicerae. 216, frontal, 217, lateral. 218, 219, carapace and abdomen. 218, dorsal. 219, lateral. 220-222, epigynum. 220, ventral. 221, posterior. 222, posterior, cleared. Figures 223-229. M. cranion Mello-Leitao, female. 223, 224, carapace and chelicerae. 223, frontal. 224, lateral. 225, 226, carapace and abdomen. 225, dorsal. 226, lateral. 227-229, epigynum. 227, ventral. 228, posterior. 229, posterior, cleared. Scale lines. 1.0 mm; genitalia, 0.1 mm. 350 Specimens Examined. COSTA RICA San José: San José, 12 (Tristan, MCZ); San Antonio de Escazt, 1,350 m, May 1980, 12 (W. Eberhard $31, MCZ): 7 Sept. 1980, imm., 2, many 6 (W. Eberhard SJ1—26, MCZ): 11 Oct. 1980, 12 (W. Eberhard $31, MCZ). Puntarenas: Parrita, 30 m, 12 Jan. 1987, 32, 2d (W. Eberhard FN9—40ff, MCZ). Mastophora dizzydeani Eberhard Figures 205-215; Map 3A Mastophora dizzydeani Eberhard, 1981: 144, figs. 1-9, 2, d. Female holotype from field of Melendez campus of the Universidad del Valle on the south- ern edge of Cali, Colombia, in MCZ, examined. Platnick, 1989: 340. Platnick, 2001. Description. Female holotype. Carapace dark brown. Sternum orange-brown. Legs brown. Abdomen white anteriorly, dorsally with transverse gray band, posteriorly dusky white (Figs. 207, 208); venter with a median white square. Carapace heavily sclerotized, grooves on sides, many tuber- cles flat and wide (Figs. 205, 206). Abdo- men subtriangular with pair of dorsal humps (Fig. 207). Total length 13.3 mm. Carapace 5.7 mm long, 5.3 wide in tho- racic region, 3.3 wide at lateral eyes. First femur 4.8 mm, patella and tibia 6.7, meta- tarsus 4.7, tarsus 1.5. Second patella and tibia 5.0 mm, third 2.8, fourth 4.4. Length of first patella and tibia 1.2 times width of carapace. Male allotype from eastern edge of Lago Calima. Yellowish white with longitudinal white thoracic mark, and dorsum of ab- domen white with a couple of indistinct gray patches anteriorly. Height of clypeus about 1.3 diameters of anterior median eye. Abdomen with a pair of humps. Pal- pal patella with one weak macroseta. Total length 1.6 mm. Carapace 0.75 mm long, 0.74 wide in thoracic region, 0.55 wide at lateral eyes. First femur 0.71 mm, patella and tibia 0.72, metatarsus 0.41, tarsus 0.26. Second patella and tibia 0.65 mm, third 0.36, fourth 0.51. Length of first patella and tibia about same length as width of carapace. Note. Males and females have been col- lected together at Cali, Colombia. Variation. Total length of females 10.8— Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 13.3. mm. The illustrations were made from specimens from Cali. Diagnosis. Females are distinguished by the wide heart-shaped abdomen with two humps (Fig. 207) and the distinctly shaped dark marks around the slits on the poste- rior of the epigynum (Fig. 210). The atria separate from each other ventrally (Fig. PAUL). Males have a smaller median apophysis (Fig. 213) than that of M. fasciata, in ectal view surrounding a rounded space (Fig. PAS) Natural History. The spiders rest on ex- posed sites: on the barb of a barbed wire, fence posts, and upper surface of leaves. Specimens also were collected on a guay- oba tree in a yard and in sugar cane areas. Moths caught include Spodoptera frugi- perda (a sugar cane pest) and Leucania sp. (Eberhard, 1977, 1981). Distribution. Colombia to northern Peru (Map 3A). Specimens Examined. COLOMBIA Valle: Sof Cali, on plants, 6 June 1948, 12 (E. M. Poulsen, ZMUC); nr. Cali, Jan. 1977, imm. 2 (W. Eberhard, MCZ); Aug. 1977, 12, 4d (W. Eberhard, MCZ); Lago Calima, 1,400 m, 19 Nov. 1977, 12 (W. Eber- hard EG 3-20, MCZ); Rio Tulua, 1,100 m, Aug. 1977, 26 (W. Eberhard, MCZ). PERU Piura: Mal- lagra, Rio Chira, 8 June 1941, 22, imm. (D. L. Friz- zell and H. E. Frizzell, AMNH, CAS). Mastophora pickeli Mello-Leitao Figures 216-222; Map 3D Mastophora pickeli Mello-Leitao, 1931: 73, figs. 6, 18, 24, 25, 2. Female holotype from Tapera, Pernam- buco, Brazil, in MNRJ, 395, examined. The specific name is a noun in apposition after the locality. Roewer, 1942: 901. Platnick, 2001. Glyptocranium pickeli:—Bonnet, 1957: 1998. Note. Vanzolini and Papavero (1968) listed three localities with the name Tap- ero in Pernambuco. I assume this locality is the only one also listed in the Index to Map of Hispanic America (American Geo- graphical Society of New York, 1944), a railroad station, west of Recife. Description. Female holotype. Cara- pace, chelicerae, labium, endites, brown. Sternum patchy orange-brown. Coxae and distal leg articles brown. Abdomen black anteriorly enclosing white streaks, white posteriorly (Fig. 218); ventrally with indis- tinct white square on gray. Carapace with shallow tubercles, glossy, with short setae on sides (Figs. 216, 217). Abdomen with- out setae, with a pair of wide humps bear- ing distinct smaller humps dorsally (Figs. 218, 219) and slight swellings on side (Fig. 219). Total length 9.5 mm. Carapace 4.4 mm long, 3.7 wide in thoracic region, 2.4 wide at lateral eyes. First femur 3.4 mm, patella and tibia 4.8, metatarsus 3.3, tarsus 0.9. Second patella and tibia 3.4 mm, third 2.0, fourth 3.2. Length of first patella and tibia 1.1 times width of carapace. Males are not known. Diagnosis. This species is distinguished by distinct small humps on a larger swell- ing of the abdomen (Figs. 218, 219) and the epigynum with atria approaching each other (Figs. 221, 222). It differs from M. ypiranga, which has a similar epigynum, by having the carapace, viewed from an- terior, wider and more swollen, the ante- rior median eyes facing slightly laterally and ventrally, and the forked tubercles laid back, with their tips facing posteriorly (Fig. 217). Distribution. Known only from the type locality (Map 3D). Specimens Examined. No other specimens have been found. Mastophora cranion Mello-Leitao Figures 223-229; Map 3D Mastophora cranion Mello-Leitao, 1928: 49, pl. 1, @. Female holotype from Tapera, Est. Pernambuco, Brazil, in MNRJ no. 00394, examined. Mello-Lei- tao, 1931: 72, figs. 2, 14, 9. Roewer, 1942: 955. Platnick, 2001. Glyprocranium cranion:—Bonnet, 1957: 1997. Note. For locality information, see note under M. pickeli. Description. Female holotype. Carapace orange-brown, black pigment between an- terior median eyes. Chelicerae, labium, endites brown. Sternum uneven white and brown. Coxae and distal leg articles light brown. Abdomen white with a pair of dor- MASTOPHORA ° Levi 351 sal dark spots, each consisting of stippled black dots (Fig. 218); venter whitish with white square, spinnerets brown. Carapace with long white setae on sides and on clyp- eus and many small tubercles (Figs. 223, 224). Abdomen with wide humps, anterior edge with long setae (Figs. 225, 226). Total length 10.8 mm. Carapace 4.7 mm long, 4.3 wide in thoracic region, 2.8 wide at lat- eral eyes. First femur 4.0 mm, patella and tibia 6.3, metatarsus 4.7, tarsus 1.1. Sec- ond patella and tibia 4.3 mm, third 2.4, fourth 3.7. Length of first patella and tibia 1.5 times width of carapace. Males are not known. Diagnosis. Mastophora cranion is distin- guished from others by black spots on the wide humps of the abdomen (Figs. 225, 226), its long white setae, and its epigyn- um with the atria in line with the outer margin of the seminal receptacles (Fig. 929). Distribution. Known only from the type locality (Map 3D). Specimens Examined. No other specimens have been found. Mastophora longiceps Mello-Leitao Figures 230-236; Map 3D Mastophora longiceps Mello-Leitéo, 1940: 57, fig. 5, ?. Female holotype from Ilha Sao Sebastiao, Est. Sao Paulo, Brazil, in MNRJ, examined. Brignoli, 1983: 273. Platnick, 2001. Glyptocranium longiceps:—Bonnet, 1957: 1998. Description. Female holotype. Carapace beige in center, dark brown on sides. Che- licerae, labium, endites, sternum orange- brown. Coxae and distal leg articles or- ange-brown. Abdomen _ contrastingly marked black and white (Figs. 232, 233); venter gray with a pair of light patches. Carapace shiny with short setae on sides, both black and white; forked horns thick, fingerlike (Figs. 230, 231). Legs with short black and white setae. Eyes indistinct without dark pigment. Abdomen bald with wide humps (Figs. 232, 233). Total length 13.0 mm. Carapace 7.5 mm long, 6.0 wide in thoracic region, 3.8 wide at lateral eyes. First femur 6.3 mm, patella and tibia 9.2, 352 metatarsus 6.4, tarsus 1.6. Second patella and tibia 6.5 mm, third 3.5, fourth 5.6. Length of first patella and tibia 1.5 times width of carapace. Diagnosis. The thick horns (Figs. 230, 231) and wide median plate on the pos- terior of the epigynum (Fig. 235) separate this species from M. melloleitaoi (Figs. 274—280). Distribution. Known only from the type locality (Map 3D). Specimens Examined. No other specimens have been found. Mastophora pesqueiro new species Figures 237-243; Map 3D Holotype. Female holotype from Pesqueiro, Monte- negro, Rio Grande de Sul, Brazil, 14 June 1977 (M. F. “Beurmann), in MCN no. 5730. The specific name is a noun in apposition after the locality in Montenegro. Description. Female holotype. Carapace light brown to dark red-brown, glossy, with two lines of white setae posteriorly on each side and a white edge. Chelicerae, labium, endites, sternum, coxae light brown. Distal leg stawles light brown, darker ventrally, ith white Stace Abdomen marked with two darker framed white patches on each side and a transverse dusky line anteriorly in a median white area (Figs. 239, 240); venter light brown with white square. Car- apace glossy with posterior horns pointed and two small median tubercles (Figs. 237, 238). Abdomen with small, pointed humps (Figs. 239, 240). Total length 10.5 mm. Carapace 4.7 mm long, 4. 6 wide in tho- racic region, 2.3 wide at lateral eyes. First femur Te 3mm, patella and tibia 6.9, meta- tarsus 4.6, tarsus 1.4. Second patella and tibia 5.0 mm, third 2.7, fourth 4.7. Length of first patella and tibia 1.5 times carapace width. Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 Males are not known. Diagnosis. Mastophora pesqueiro is dis- tinguished by the conical horns on the smooth carapace (Figs. 237, 238), the four white marks on the abdomen, and the dis- tinctly shaped humps (Figs. 239, 240). The epigynum has a median bulge on its pos- terior face (Figs. 241-243). Distribution. Known only from the type locality (Map 3D). Specimens Examined. No other specimens have been found. Mastophora piras new species Figures 244-250; Map 3D Holotype. Female yolotype from Emas, Pirassunga, Est. Sao Paulo, Brazil, 2 Nov. 1952 (Pietracatelli, Werner, and Dionisio), in MZSP, 4339. The specific name is an arbitrary combination of letters. Description. Female holotype. Carapace dark brown. Chelicerae light brown. La- bium, endites, sternum light brown. Coxae and distal leg articles began with indistinct darker Bigtches! Abdomen anterior black enclosing some white anterior loops, pos- terior white (Figs. 246, 247); venter with indistinct white square on gray-brown. Carapace heavily sclerotized, shiny, with sides thinner with a punctuate pattern of tiny tubercles. Abdomen triangular heart- shaped with lateral bulges and distinct small humps (Figs. 246, 247). Total length 13.8 mm. Carapace 6.5 mm long, 5.9 wide in thoracic region, 3.3 wide at lateral eyes. First femur 5.3 mm, patella and tibia 7.7, metatarsus 4.8, tarsus 1.6. Second patella and tibia 5:5 mm: third 3/3) fourth 4.8: Length of first patella and tibia 1.2 times width of carapace. Males are not known. Variation. Total length of females 13.0— 13.8 mm. The paratype has some long, white setae on sides of carapace, clypeus, —>, Figures 230-236. Mastophora longiceps Mello-Leitao, female. 230, 231, carapace and chelicerae. 230, frontal. 231, lateral. 232, 233, carapace and abdomen. 232, dorsal. 233, lateral. 234-236, epigynum. 234, ventral. 235, posterior. 236, posterior, cleared. Figures 237-243. MM. pesqueiro new species, female. 237, 238, carapace and chelicerae. 237, frontal. 238, lateral. 239, 240, carapace and abdomen. 239, dorsal. 240, lateral. 241-243, epigynum. 241, ventral. 242, posterior. 243, posterior, cleared. MASTOPHORA ° Levi 353 a0 CORY 2, S longiceps Figures 244-250. OM. piras new species, female. 244, 245, carapace and chelicerae. 244, frontal. 245, lateral. 246, 247, car- apace and abdomen. 246, dorsal. 247, lateral. 248-250, epigynum. 248, ventral. 249, posterior. 250, posterior, cleared. Figures 251-257. M. ypiranga new species, female. 251, 252, carapace and chelicerae. 251, frontal. 252, lateral. 253, 254, carapace and abdomen. 253, dorsal. 254, lateral. 255-257, epigynum. 255, ventral. 256, posterior. 257, posterior, cleared. Scale lines. 1.0 mm; genitalia, 0.1 mm. 354 legs, and anterior of abdomen; the abdo- men is all dark, perhaps because of poor preservation. The illustrations were from the holotype. Diagnosis. Mastophora piras is distin- guished from M. melloleitaoi by the heart- shaped abdomen with small humps (Figs. 246, 247) and by the wider median area of the epigynum in posterior view (Fig. 249). Distribution. Minas Gerais and Sao Pau- lo states of southern Brazil (Map 3D). Paratypes. BRAZIL Minas Gerais: Gavernador Va- ladares, 14/15 Oct. 1981, 1 imm. (N. Sorkin, T. Spitz- man, AMNH). Sdo Paulo: Sio Bernardo, Nov. 1926, 12 (H. Bakkenist, MZSP 8069). Mastophora ypiranga new species Figures 251-257; Map 3E Holotype. Female holotype from Ypiranga, Cap. [city of Sido Paulo], Est. Sao Paulo, Brazil, 1898, in MZSP, 5791. The specific name is a noun in ap- position after the locality. Description. Female holotype. Carapace orange-brown. Chelicerae yellow-brown. Labium, endites, sternum light brown. Coxae and distal leg articles light brown. Abdomen anterior black, posterior white (Figs. 253, 254); venter grayish white with white square. Carapace with low tubercles, with some short white setae on sides (F igs. 251, 252). First femur with S-shaped cur- vature. Abdomen with two humps (Figs. 253, 254). Total length 12.7 mm. Carapace 4.7 mm long, 4.2 wide in thoracic region, 2.7 wide at lateral eyes. First femur 3.7 mim, patella and tibia 5.4, metatarsus 3.3, tarsus 1.1. Second patella and tibia 3.8 mm, third 2.2, fourth 3.6. Length of first patella and tibia 1.2 times width of cara- pace. Males are not known Variation. Total length of females 9.7— Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 12.7 mm. The illustrations were made from the holotype. Diagnosis. Mastophora ypiranga differs from M. pickeli and M. melloleitaoi by the shape of the abdomen (Figs. 253, 254) and by the narrower, almost triangular median plate of the epigynum and the atria ap- proaching each other (Figs. 256, 257). Other differences are that the carapace ap- pears less swollen when viewed from an- terior, the anterior median eyes are di- rected forward, and the horns are erect with the tips pointing dorsally (Fig. 252). The anterior of the abdomen, which over- hangs the carapace, is white. Distribution. Minas Gerais to Santa Ca- tarina states of southern Brazil (Map 3E). Paratypes. BRAZIL Minas Gerais: Vicosa, 1931, 12 (E. J. Hambleton, AMNH). Sdo Paulo: Sao Paulo, July 1928, 12 (R. Cassalo, IBSP 4530). Santa Catar- ina: June 1919, 12 (Luederwaldt, MZSP 8067). Mastophora yacare new species Figures 258-264; Map 3E Holotype. Female holotype from Rincori del Yacaré, Artigas, Uruguay, 20 Jan. 1957, in FCMU. The spe- cific name is a noun in apposition after the locality. Description. Female holotype. Carapace light brown. Chelicerae, labium, endites brown. Sternum orange, underlain by white pigment. Legs brown, femora light- est. Abdomen black over humps, whitish posteriorly (Figs. 260, 261); venter light brown with white square containing two rows of three black spots; spinnerets dark brown. Abdomen with a pair of wide, dor- sal humps (Figs. 260, 261). Total length 11.5 mm. Carapace 4.1 mm long, 4.0 wide in thoracic region, 2.6 wide behind pos- terior lateral eyes. First femur 3.5 mm, pa- tella and tibia 5.1, metatarsus 3.5, tarsus 1.1. Second patella and tibia 3.8 mm, third = Figures 258-264. Mastophora yacare new species, female. 258, 259, carapace and chelicerae. 258, frontal. 259, lateral. 260, 261, carapace and abdomen. 260, dorsal. 261, lateral. 262-264, epigynum. 262, ventral. 263, posterior. 264, posterior, cleared. Figures 265-268. M. abalosi Urtubey and Baez, female, after authors. 265, horns of carapace, frontal. 266, carapace and chelicerae, lateral. 267, carapace and abdomen, dorsal. 268, epigynum, posterior. Figures 269-273. M. comma Baez and Urtubey, female, after authors. 269, horns of carapace. 270, carapace and chelicerae, lateral. 271, 272, carapace and abdomen. 271, dorsal. 272, lateral. 273, epigynum, posterior. MASTOPHORA ° Levi 355 eet fis yi? fae % 258 ae % 259 Ps abalosi ae yacare 260 as n n n a (498 Phe & aoe (ger eo 4 5 \ Ay ie - v 28I 282 oe Neier eth 2s 182 ON extraordinaria 278 283 285 Figures 274-280. M. melloleitaoi Canals, female. 274, 275, carapace and chelicerae. 274, frontal. 275, lateral. 276, 277, carapace and abdomen. 276, dorsal. 277, lateral. 278-280, epigynum. 278, ventral. 279, posterior. 280, posterior, cleared. Figures 281-287. M. extraordinaria Holmberg, female. 281, 282, carapace and chelicerae. 281, frontal. 282, lateral. 283, 284, carapace and abdomen. 283, dorsal. 284, lateral. 285-287, epigynum. 285, ventral. 256, posterior. 287, posterior, cleared. Scale lines. 1.0 mm; genitalia, 0.1 mm. 356 2.0, fourth 3.3. Length of first patella and tibia 1.3 times width of carapace. Males are not known. Diagnosis. Mastophora yacare is distin- guished from M. melloleitaoi by its epigyn- um and the atria, which separate and face sideways (Fig. 264). Distribution. Known only from the type locality (Map 3E). Specimens Examined. No other specimens have been found. Mastophora abalosi Urtubey and Baez Figures 265-268, 458; Map 3E Mastophora abalosi Urtubey and Baez, 1983: 3, figs. 1-11, 2. Female holotype and seven paratypes from the city of Santiago del Estero, Argentina, in the Inst. Animales Venenosos, Santiago del Estero, unavailable. Platnick, 1989: 340. Platnick, 2001. Description. Female (after Urtubey and Badez, 1983). Carapace brown. Chelicerae yellowish. Labium, endites yellow-white. Sternum spotted orange-yellow. Sternum dark orange. Legs yellowish. Abdomen an- terior clear yellow followed by dark band that covers humps (Fig. 267), posteriorly pale yellow; venter with light yellow. Car- apace with numerous tubercles, setae only on sides; tubercles with light tips. Abdo- men with humps (Fig. 267). Total length 12.5 mm. Carapace 5.6 mm long, 5.4 wide in thoracic region. First femur 4.8 mm, pa- tella and tibia 6.8, metatarsus 5.1, tarsus 1.3. Second patella and tibia 5.2 mm, third 2.6, fourth 4.8. Length of first patella and tibia 1.3 times width of carapace. Males are not known. Diagnosis. Mastophora abalosi is distin- guished from M. extraordinaria by some morphological characters (Urtubey and Baez, 1983). The posterior of the epigyn- um was illustrated (Fig. 268) and shows the atria slightly to the lateral of the slits. The egg sac has flaps (Fig. 458). Distribution. Santiago del Estero in northern Argentina (Map 3E). Specimens Examined. No specimens were avail- able. Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 Mastophora comma Baez and Urtubey Figures 269-273, 459; Map 3E Mastophora comma Baez and Urtubey, 1985: 3, figs. 1-9, 2. Female holotype from the city of Santiago del Estero, Argentina, and seven paratypes in the Instituto de Animales Venenosos, Santiago del Es- tero, unavailable. Platnick, 1989: 340. Platnick, 2001. Description. Female (after Baez and Ur- tubey, 1985). Carapace brown. Chelicerae yellow. Sternum orange. Legs orange-yel- low. Abdomen dusky yellow covered with many tiny maroon spots with white setae, spotted with brown spots and a few darker maroon spots, and posteriorly with a few transverse bands (Fig. 271); venter yellow, spinnerets maroon. Intense yellow area between epigastric groove and spinnerets, with a few small maroon spots. Carapace with tubercles (Figs. 269, 270). Abdomen with a pair of dorsal humps tipped by nip- ples (Figs. 271, 272). Total length 12.0 mm. Carapace 3.7 mm long, 4.8 wide in thoracic region. First femur 3.9 mm, pa- tella and tibia 6.4, metatarsus 5.0, tarsus 1.2. Second patella and tibia 4.6 mm, third 2.4, fourth 3.9. Length of first patella and tibia 1.3 times width of carapace. Males are not known. Diagnosis. This species differs from oth- ers by the unusual structure of the abdo- menal humps (Fig. 272) and by the two parallel slits on the posterior face of the epigynum (Fig. 273). The egg sac lacks flaps and has a short stalk (Fig. 459). Distribution. Santiago del Estero in northern Argentina (Map 3E). Specimens Examined. No specimens of this species were available. Mastophora melloleitaoi Canals Figures 274-280; Map 3E Mastophora Mello-Leitdoi Canals, 1931: 20, figs. 1-4, pl. 2, fig. 4, 2. Female holotype, from Rosas, Prov. Buenos Aires, Argentina, in MACN, examined. Mastophora_ mello-leitéo:—Mello-Leitao, 1931, 73: Ine Oy ILIA Se Mastophora mello-leitaoiae:—Roewer, 1942: 900. Glyptocranium melloleitaoi:—Bonnet, 1957: 1998. Mastophora melloleitaoi:—Platnick, 2001. Note. Roewer’s (1942) spelling appears to be an error. Description. Female from Balcarce. Carapace light brown, not shiny. Chelic- erae, endites, labium light brown. Sternum light brown. Legs brown. Abdomen light brown, anterior gray. Near midline ante- riorly a light V-shaped mark, followed by a light upside-down V. Carapace bald, ex- cept for some setae posteriorly. Abdomen humps wide (Figs. 276, 277). Total length 13.0 mm. Carapace 5.5 mm long, 4.8 wide in thoracic region, 3.2 wide at lateral eyes. First femur 4.2 mm, patella and tibia 6.3, metatarsus 4.2, tarsus 1.3. Second patella andatibial4:oyamm) thirds 2. fourth: 4:2) Length of first patella and tibia 1.3 times width of carapace. Males are not known. Variation. Total length of females 9.0— 12.8 mm. The total length of the holotype is 12.8 mm, carapace 4.6 mm wide, patella and tibia 5.8. Humps may have black caps. The illustrations (Figs. 274-278) were made from a female from Balcarce; Fig- ures 279 and 280 are from the holotype. Diagnosis. The female differs from M. extraordinaria in that the carapace is not shiny (Fig. 274), in having the anterior of the abdomen dark (Figs. 276, 277), and in having the slits on the posterior of the epi- gynum almost parallel (Fig. 279). Masto- phora melloleitaoi also lacks the dark me- dian dorsal area on the posterior of the epigynum present in M. extraordinaria (Fig. 286). If cleared, each atrium shows a small median lobe (Fig. 280) not present in other species. Distribution. From Parana State, south- ern Brazil, to Buenos Aires Province, Ar- gentina (Map 3E). Specimens Examined. BRAZIL Rio de Janeiro: Vas- souras, Fazenda de Sao Sebastiao, March 1871, 12 (B. P. Mann, MCZ). Paranda: Curitiba, 21 Jan. 1965, 1 imm. (C. Valle, MZSP 4326). Rio Grande do Sul: Rodeio Bonito, Bagé, 10 Feb. 1967, 12 (C. de Oliv- eira, MCN 473); Santa Maria, 9 May 1973, 12 (D. Link, MCN 01659). ARGENTINA Cordoba: Cruz Alta, 15 Aug. 1948, 12 (J. P. Duret, MACN). Buenos Aires: Ascension, 12 (B. Gerschman, MACN): Bal- carce, 16 Feb. 1950, 12 (Cuccioli, MACN):; Burzaco, MASTOPHORA ® Levi 357 12 (F.C. S., MACN); San Isidro, 19 Apr. 1949, 19 (N. Konmilev, MACN): Zelaya, 2 imm. (J. Pereyra, MACN 523). Mastophora extraordinaria Holmberg Figures 281-287, 460; Map 3F Mastophora extraordinaria Holmberg, 1876: 113. Fe- male from Buenos Aires, Argentina, lost. Bréthres, 1909: 163, figs. 1, 2, 2, egg sacs. Canals, 1931: 17, figs. 1-5, pl. 1, fig. 1. Mello-Leitao, 1931: 70, figs. 7, 19. Roewer, 1942: 900. Platnick, 2001. Glyptocranium extraordinaria:—Bonnet, 1957: 1997. ?Mastophora cinerea Mello-Leitaéo, 1943: 105, fig. 4, imm. Immature holotype from Cordoba, Argenti- na, in MLP, examined. DOUBTFUL NEW SYN- ONYMY. Mastophora intermedia Mello-Leitao, 1945: 240, figs. 14-17, 2. One female holotype from Pindapoy, Mi- siones Prov., Argentina, in MLP, examined. Brig- noli, 1983: 274. NEW SYNONYMY. Note. Holmberg described the size of the prominences of the abdomen and the two brown spots. Mastophora cinerea is a light-colored immature with slightly elongate humps. Its placement is doubtful. (Do immature M. extraordinaria have longer humps?) Mastophora intermedia has humps rounded, as wide as long, and has the same internal genitalia as does M. extraordinar- ia. The differences are a slight median notch of the posterior rim of the epigynum in ventral view, and absence of the median dark area posterior in the epigynum (Fig. 286). Description. Female from Chascomis. Carapace dark brown with narrow white rim. Legs brown, slightly ringed. Abdomen white with a pair of dorsal black patches (Fig. 283). Carapace shiny on tips with low tubercles and three grooves on each side. Abdomen with few setae on sides with one pair of humps; venter with white square. Total length 12 mm. Carapace 4.8 mm long, 4.6 wide in thoracic region, 2.8 wide at posterior lateral eyes. First femur 3.9 mm, patella and tibia 5.5, metatarsus 3.5, tarsus 1.2. Second patella and tibia 4.5 mim, third 2.7, fourth 4.1. Length of first patella and tibia 1.1 times width of cara- pace. Males are not known. 358 Variation. The epigynum is variable in width and the abdominal humps may be wider or higher. Total length of females 95-14 mm. The illustrations were made from the female from Chascomis. Diagnosis. Compared with M. mellolei- taoi, the carapace has low, flat tubercles; the horns of the carapace are relatively smooth and shiny (Figs. 281, 282); and the bulge bearing the median eyes is indis- tinct. The anterior of the abdomen is usu- ally white (Fig. 284). The median plate of the epigynum is flat dorsally, sclerotized, and darker than ventrally (above on Fig. 286); it is raised in the middle in M. mel- loleitaoi. The atria approach each other (Fig. 287), unlike those of M. melloleitaoi (Fig. 280). Natural History. The egg sac is drop- shaped, 11 mm wide, almost the size of the female abdomen (Fig. 460). A female from Gonzales Catan was found on a citrus tree. An immature in Uruguay was col- lected at night “from regular web.” Distribution. From Rio Grande do Sul State, southern Brazil, to Buenos Aires Province, west to Chaco and Cérdoba provinces, Argentina (Map 3F). Specimens Examined. BRAZIL Rio Grande de Sul: Canela, 12 Feb. 1966, 12 (A. Lise, MCN 0119): Gar- ibaldi, 30 Oct. 1974, 12 (O. ame MCN 2381). URUGUAY Montevideo, 1? (J. Canosa, MACN 4182); 12 (E. Cordero, MNRJ 14011); Pun- tas Arroyo Laureles, Tacuaremb6, 1 imm. (FCMU 293): Treinta y Tres, 20 Aug. 1971, 1 2 (FCMU 295). ARGENTINA Chaco: Saenz Peja, Sept. 1933, 19 (B. Ohneiser, MACN 31331). Cordoba: Cérdoba, 22 M. J. Viana, AMNH, MACN 1106); Calamuchita, Dec. 1940, 49 (J. M. Viana, MACN 1005): Dec. 1941, 22 (J. M. Viana, MACN 1106); Agus do Oro, Mar. 1940, 1 imm. (J. A. De Carlo). Buenos Aires: Buenos Aires, 12 (Scly., MNHN 23388); Chascomits, Oct. 1934, 12 (I. Dor, MACN 35983); Cap. Federal, limm. (E. Pizarro, MACN 12782): Florencio Varela, 1 May 1949, 1° (O. de Ferrarini, MLP 13549); Gon- zales Catan, 6 June 1949, 12 (Touson); Hurlingham, Jan. 1954, 12 (Giai, MACN 4359); La Plata, 12 (M. Birabén, MLP 16178); 12 (M. Birabén, BMNH): Moreno, Feb. 1939, 12 (Schiapelli, Gerschman, MACN); 14 Nov. 1943, 12 (S. M. Doello Jurado; MACN 1361); Rosas, 1930, 1° (J. B. Daguerre, MACN 4185). Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 Mastophora leucabulba (Gertsch), new combination Figures 288-295; Map 2F Agatostichus leucabulba Gertsch, 1955: 250: figs. 34, 38, 40, 2. Female holotype from Harlingen, Texas, in AMNH, destroyed. Agathostichus leucabulba:—Brignoli, 1983: 255. Agathostichus leucabulbus: pisincle 2001. Note. Platnick’s change of spelling is not required. Description. Female (after Gertsch, 1955). Carapace reddish brown except for yellow rim and orange median patch en- closing tubercles; tubercles tipped white. Chelicerae Ibvaeseornfislh at base. Labium, en- dites yellowish. Sternum yellowish. Coxae yellowish and distal leg articles yellowish to light brown with fait brown markings on fins leg. Abdomen yellowish to whitish above with dusky flecks, black around the base (Fig. 289). Carapace with woolly se- tae, one large bulb behind median eyes, a large iedian bulb flanked by a pair of sniel bulbs on the side, behind bifid horns, sides with dark warts and cephalic area coarsely roughened, hidden by mat of wooly hairs. betel eyes on connate tu- bercles, median eyes on large elevated tu- bercle at the posterior edge of which is a small tubercle. Abdomen with long humps. Total length 6.7 mm. Carapace 2.8 mim long, 2.9 dee in thoracic region. First femur 3.6 mm, patella and tibia 4.9, meta- tarsus 3.6, tarsus 1.0. Second patella and tibia 3.4 mm, third 1.9, fourth 2.6. Length of first patella and tibia 1.7 times width of carapace. Diagnosis. Mastophora leucabulba is distinguished by the enlarged tubercle be- tween the median eyes (Figs. 288, 290) and the large blunt tubercles and dark col- oration of the carapace (Figs. 289, 290). Doubtful males, considered and labeled M. cornigera by W. Ivie, and listed under that species by Gertsch, may be this spe- cies (Figs. 292-295). Distribution. Southern Texas to Hon- duras (Map 2F). Specimens Examined. TEXAS Duval Co.: San Di- MASTOPHORA ® Levi 359 leucabulba Figures 288-295. Mastophora leucabulba Gertsch. 288-291, female, after Gertsch. 288, carapace and chelicerae, lateral. 289, 290, carapace and abdomen. 289, dorsal. 290, lateral. 291, epigynum, posterior. 292-295, left palpus of presumed male. 292, apical. 293, mesal. 294, ventral. 295, ectal. Figures 296-307. WM. alvareztoroi lbarra and Jiménez new species. 296-303, female. 296, 297, carapace and chelicerae. 296, frontal. 297, lateral. 298-300, carapace and abdomen. 298, dorsal. 299, ventral. 300, lateral. 301-303, epigynum. 301, ventral. 302, posterior. 303, posterior, cleared. 304-307, male. 304, carapace, chelicera and right palpus. 305, dorsal. 306-307, left palpus. 306, mesal. 307, ventral. Scale lines. 1.0 mm; genitalia, 0.1 mm. 360 ego, 29 Apr. 1895, 2d, imm. doubtful determination (USNM). Wilson Co.: Floresville, 29 Apr. 1895, 1 imm., 4d, doubtful determination (AMNH). MEXI- CO Tamaulipas: 64 km S Linares, imm., 1.9 mm long (Gertsch, 1955, AMNH, destroyed, not examined). HONDURAS E of Tela Beach, 6 July 1929, imm. (A. M. Chickering, MCZ). Mastophora alvareztoroi Ibarra and Jiménez new species Plate 1; Figures 296-307; Map 2F Holotype. Female holotype from Rancho Alejandria, Municipio Estaci6n Juarez, Chiapas, Mexico, 25 Sept. 1975 (M. Alvarez del Toro), in MCZ. The species has been named after the collector, the late Miguel Alvarez del Toro, who dedicated his life to the study and protection of the Chiapas fauna and is the author of a book on Chiapas spiders (Alvarez del Toro, 1992). Agathostichus sp. Alvarez del Toro, 1992: 173, fig. ES SR photos Description. Female from Rosaria Iza- pa. Carapace horns white, sides dark brown with eyes, clypeus, and rim yellow- ish (Figs. 296-298). Chelicerae yellowish with a pair of dark patches. Labium, en- dites dark brown, distally light. Sternum dark brown. Coxae and aise leg articles yellowish; first femur with fine black ring and distally with ventral dark patch. Nie domen yellow, dark brown on anterior bor- der, and brown on some hump tips (Fig. 298); venter light brown with white square; a dark brown patch on epigynal area, and no pigment in spinneret area. Carapace hirsute with cone-shaped white tubercles (Figs. 296, 297). Posterior me- dian eyes 0.8 diameter of anterior medi- ans, laterals 0.7. Anterior median eyes 1.7 diameters apart. Ocular trapezoid rectan- gular, slightly wider than long. Chelicerae with one anterior tooth and three posterior teeth. Abdomen heart-shaped, three pairs of dark spots dorsally and sparse black se- tae, hirsute with tufts on white setae on Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 dorsum. A pair of dorsal humps, followed by a series of four median, conical humps and several smaller lateral humps on each side. Setae, also tufts of white setae on small circular and convex white areas, scat- tered on dorsum of abdomen, with scat- tered dark brown setae, especially on an- terior half, also with a few feather-shaped symmetrically arranged dark brown setae (Fig. 298). Total length 8.9 mm. Carapace 3.4 mm long, 3.3 wide in thoracic region, 1.7 wide at lateral eyes. First femur 4.3 mim, patella and tibia 5.5, metatarsus 4.2, tarsus 1.1. Second patella and tibia 3.8 mm, third 2.3, fourth 3.5. Length of first patella and tibia 1.6 times width of cara- pace. Male weakly sclerotized, perhaps just molted. Carapace reddish brown except for white median light patch enclosing white horns. Chelicerae dark yellow. Ster- num dark brown. Coxae, distal leg articles yellowish. Abdomen whitish. Carapace el- evated behind, second midline tubercle slightly longer than first, lateral horns shout half length of median horns, sepa- ration beueen median and lateral horns greater than width of laterals at base; each transverse horn with translucent seta on tip; carapace with woolly white setae. Che- licerae with three posterior median teeth and one posterior tooth. Abdomen with two dorsal paired humps; four anterior, four median, and two posterior feather- shaped setae (Fig. 305). Total length 1.7 mm. Carapace 0. 8 mm long, 0.8 wide in thoracic region. First ne 1.1 mm, pa- tella and aie 1.1, metatarsus 1.1, tarsus 1.03. Second patella and tibia 0.8 mm, third 0.4, fourth 0.6. Length of first patella and tibia 1.4 times width of carapace. Note. Males and females were matched = Figures 308-314. Mastophora soberiana new species, female. 308, 309, carapace and chelicerae. 308, frontal. 309, lateral. 310, 311, carapace and abdomen. 310, dorsal. 311, lateral. 312-314, epigynum. 312, ventral. 313, posterior. 314, posterior, cleared. Figures 315-321. Mastophora leucacantha (Simon), female. 315, 316, carapace and chelicerae. 315, frontal. 316, lateral. 317, 318, carapace and abdomen. 317, dorsal. 318, lateral. 319-321, epigynum. 319, ventral. 320, posterior. 321, posterior, cleared. MASTOPHORA ° Levi 361 Figures 322-328. M. brescoviti new species, female. 322, 323, carapace and chelicerae. 322, frontal. 323, lateral. 324, 325, carapace and abdomen. 324, dorsal. 325, lateral. 326-328, epigynum. 326, ventral. 327, posterior. 328, posterior, cleared. Figures 329-335. WM. conifera (Holmberg), female. 329, 330, carapace and chelicerae. 329, frontal. 330, lateral. 331, 332, carapace and abdomen. 331, dorsal. 332 lateral. 333-335, epigynum. 333, ventral. 334, posterior. 335, posterior, cleared. Scale lines. 1.0 mm; genitalia, 0.1 mm. 362 because they were collected in the same area in Chiapas, have similar cone-shaped carapace tubercles and horns, and both sexes have feather-shaped setae. Variation. Living females look all woolly. Total length of females 8.6—8.9 mm, cara- pace 3.3—4.0 mm wide. An immature, 4.3 mm total length, is similar to the adult fe- male in relative proportions, horns, and pointed abdomen, but has fewer abdomi- nal humps (only the pair and the four in midline). As in males of the genus, a row of long setae is present on the anterior of the first and second metatarsi and tarsi. The illustrations (Figs. 298-300) were made from the paratype collected with the holotype. Diagnosis. Mastophora alvareztoroi is distinguished from M. leucabulba by the shape and humps of abdomen (Figs. 298— 300), by the cone-shaped tubercles on the carapace (Figs. 296, 297), by having feath- er-shaped setae (Figs. 298, 305), Rind by the epigynum having atria bent toward each other (Figs. 302, 303). Natural History. Some specimens came from coffee groves in Finca Irlanda and Rosario Izapa. The specimens were be- tween | and 2 m above the ground. Mark Stowe (in correspondence) wrote that this fuzzy spider bears a strong resemblance to congregations of fuzzy scale insects in the same habitat in Texas. Distribution. Southern Texas and Chia- pas, Mexico (Map 2F). Paratypes. MEXICO Chiapas: Rancho Alejandria, Municipio Estacién Juarez, 25 Sept. 1975, 12 (M. Alvarez del Toro, MLJ); Finca Inlanda, 870 m, 15°10'N, 92°21'W, 65 km NNW of Tapachula, 12 Aug. 1987, 1d (G. Ibarra, A. Garcia, M. Moreno, ECOTAR): Rosario Izapa, Municipio Tuxtla Chico, 430 m, 14°59'N, 92°09'W, 18 km ENE of Tapachula, 25 Aug. 1995, 12 (A. Ventura, ECOTAR). Specimens Examined. TEXAS Hidalgo Co.: Santa Ana Natl. Wildlife Refuge, 18 Dec. 1983, 12 (M. K. Stowe 112, MKS). MEXICO Chiapas: Finca Ham- burgo, ca. 15°10’N, 92°19’W, 950 m, Municipio de Tapachula, 16 Nov. 1994, 1 imm. (G. Ibarra, ECO- TAR). Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 Mastophora soberiana new species Figures 308-314; Map 2F Holotype. Female holotype from Pipeline Road, Ca- nal Zone [in Soberiana National Park, Panama], 26 July 1976 (Y. Lubin, G. G. Montgomery), in MCZ. The specific name is a noun in apposition after the locality. Description. Female holotype (in poor physical condition). Carapace yellowish, sides of thorax dark brown, a wide, light rim and tubercles with white pigment (Figs. 308, 309). Chelicerae, labium, en- dites yellowish. Sternum brown. Coxae, distal leg articles yellowish. Abdomen white dorsally (Fig. 310), white ventrally, except for yellowish genital area and spin- nerets. Carapace with spine-shaped tuber- cles and white setae (Figs. 308, 309). Ab- domen with three median humps and nu- merous small lateral humps, with tufts of white setae (Figs. 310, 311). Total length 8.3 mm. Carapace 3.7 mm long, 3.4 wide in thoracic region, 1.8 wide behind pos- terior lateral eyes. First femur 4.7 mm, pa- tella and tibia 5.8, tarsi lost. Second patella and tibia) 422 imi) third)! 3a tomntamors Length of first patella and ie 1.7 times width of carapace. Males are not known. Diagnosis. This species is distinguished from M. alvareztoroi by the longer cara- pace tubercles and by having Ths atria of the epigynum located dorsally (Fig. 314). Distribution. Panama (Map 2F). Specimens Examined. No other specimens have been found. Mastophora leucacantha (Simon), new combination Figures 315-321; Map 3G Agatostichus leucacantha Simon, 1895: 885, fig. 947, carapace. Immature holotype from Rio Salobro, Bahia, Brazil, in MNHN, 8486, examined. Mello- Leitao, 1931: 67. Gertsch, 1955: 250. Agathostichus leuwcacantha:—Simon, 1897: 473. sRoewer 1942: 900. Agathostichus leucacanthus:—Bonnet, 1955: 182. Platnick, 2001. Note. In his publications, Simon (1895, 1897) did not include the female symbol as he did for other species, indicating that the specimen was immature. Roewer (1942) and Gertsch (1955) cited it as fe- male. Platnick (2001) considered the name of Simon (1895) to be a nomen nudum be- cause the description was shared by the genus and species. But this is valid for 19th century descriptions (International Code of Zoological Nomenclature, art. 12.2.6 [International Commission on Zoo- logical Nomenclature, 1999]). Description. Female from the Organ Mountains. Carapace yellowish white with white marks and median white tubercles, brown triangle on posterior slope; anterior point of triangle dark between forked tu- bercles (Fig. "315). Chelicerae yellowish. Labium dark brown, endites yellowish. Sternum dark brown. Coxae and distal leg articles yellowish white. Abdomen yellow- ish white with a small black mark on each side (Fig. 317); venter yellowish white with white square surrounded by a gray line; dorsum and sides with indistinct gray marks. Carapace with woolly setae and long tubercles (Figs. 315-317). Median eyes on a swelling, wench lateral pair on a swelling. Median Benue trapezoid almost square. Chelicerae with three anterior teeth, one posterior tooth. Legs with white setae. Abdomen with a pair of dorsal humps and tufts of white setae (Figs. 317, 318). Total length 8.3 mm. Carapace 3.7 mm long, 3.6 wide in thoracic region, 2.2 wide at lateral eyes. First femur 3.6 mm, patella and tibia 4.6, metatarsus 3.3, tarsus 1.0. Second patella and tibia 3.6 mm, third 2.1, fourth 3.2. Length of first patella and tibia 1.3 times width of carapace. Males are not known. Variation. The immature holotype, total length 4.0 m, has carapace tubercles short- er and lacks some lateral ones; venter of abdomen with black square. The illustra- tions were made from the adult female from Organ Mountains. The tufts of setae on the abdomen were prominent on the female from Organ Mountains when first MASTOPHORA ® Levi 363 examined (by the author in 1969), but have mostly been lost as result of handling. Diagnosis. Mastophora leucacantha is distinguished by the long median horn that is almost as long as the carapace, and dif- fers from M. Dinner ror roi by having the ab- domen rounded behind, sphereas M. al- vareztoroi has the abdomen lobed behind, and differs from M. leucabulba by having all tubercles behind the eyes and from M. soberiana by the shape of the epigynum (Fig. 320). Distribution. Bahia to Rio de Janeiro states, Brazil (Map 3G). Specimens Examined. Rio de Janeiro: Cachoeirin- ha, Montaigne Orgues [Serra Orgaios, Organ Moun- tains |, 1902. 19 (iss R. Wagner, MNHN 26035). Mastophora brescoviti new species Figures 322-328; Map 3G Holotype. Female holotype from Jardim Botanico, Porto Alegre, Rio Grande do Sul, Brazil (A Brescovit), in MCN no. 26135. The species has been named after the collector and arachnologist A. D. Brescovit. Description. Female holotype. Carapace with symmetrical white lines on head re- gion, sides of cephalic area dark brown, sides of thorax light brown with dark streaks and speckles, and many downy white setae (Figs. 322, 323). Chelicerae patchy brown. Abyann: endites, sternum dark brown. Coxae light brown and distal leg articles yellowish iin temo brown rings. Abdomen light brown, darker ante- riorly between humps and pedicel, darker patches on each side, humps darkest, with bunches and individual white setae (Fig. 324); venter light brown. Lateral eyes on bulges. Carapace with median tubercle longest. Abdomen dorsally with a pair of humps and a median swelling bearing white setae (Figs. 324, 325). Total length 9.2 mm. Carapace 3.7 mm long, 3.4 ede in thoracic region, 1.8 wide Benne pos- terior lateral eyes. First femur 4.0 mm, pa- tella and tibia 5.1, metatarsus 3.7, tarsus 1.2. Second patella and tibia 3.8 mm, third Dy rowiada Sex Males are not known. 364 Diagnosis. Mastophora brescoviti is dis- tinguished by the long posterior, median tubercle, and unusual shape of the abdo- men with a median swelling behind the humps (Figs. 324, 325). The posterior of the epigynum has a pair of diagonal swell- ings (Fig. 327) not present in other spe- cies. Distribution. Only known from Porto Alegre, Brazil (Map 3G). Specimens Examined. No other specimens have been found. Mastophora conifera (Holmberg) Figures 329-335; Map 3G Heterocephala conifera Holmberg, 1876: 143. Female from Boradero [Prov. Buenos Aires], Argentina, lost. Mastophora conifera:—Canals, 1931: 18, figs. 1—5, pl. 1, fig. 2. Mello-Leitao, 1931: 71, figs. 4, 16. Roewer, 1942: 900. Platnick, 2001. Glyptocranium coniferum:—Bonnet, 1957: 1996. Description. Female from Tigre [in poor condition]. Carapace, sternum, legs or- ange-brown. Abdomen anteriorly black, posteriorly lighter gray, with some black streaks (Figs. 331, 332); venter black with a pair of white spots. Carapace with tips of horns thin, setae on sides of thoracic area (Figs. 329, 330). Anterior median eyes largest, laterals smallest. Abdomen with numerous dorsal tubercles (Figs. 331, 332). Total length 12.0 mm. Carapace 5.3 mm long, 4.6 wide in thoracic region, 2.8 wide at lateral eyes. First femur 4.3 mm, patella and tibia 5.3, metatarsus 3.8, tarsus broken. Second patella and tibia 4.3 mm, third 2.4, fourth 3.7. Length of first patella and tibia equals 1.2 times width of cara- pace. Males are not known. Diagnosis. Mastophora conifera is dis- tinguished by the tubercular abdomen Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 (Figs. 331, 332) and by the epigynum, which in posterior view has a pair of de- pressions containing short, ventrally con- verging slits (Fig. 334). Distribution. Santa Fé and Buenos Ai- res provinces, Argentina (Map 3G). Specimens Examined. ARGENTINA Santa Fé: Co- lonia Macias, Nov. 1942, imm. shriveled (J. M. Viana, MACN). Buenos Aires: Tigre, 1902, 12, once dried up (J. Brethes, MACN 5896). Mastophora corpulenta (Banks) Figures 336-342, 461; Maps 2F, 4A Ordgarius corpulentus Banks, 1898: 251, pl. 15, fig. 8. Female holotype from San José del Cabom Baja California, Mexico, in CAS, destroyed. Neotype here designated the holotype of M. lenca. Mastophora corpulenta:—Roewer, 1942: 900. Plat- nick, 2001. Mastophora lenca Gertsch, 1955: 247, figs. 28-30, 32, 33, 2. Female holotype from Zamorano [?Zambra- no], Honduras, in AMNH, examined. Brignoli, 1983: 274. Platnick, 2001. NEW SYNONYMY. Glyprocranium corpulentum:—Bonnet, 1957: 1997. Note. Banks (1898) described and pic- tured (fig. 8) elongated tubercles on the abdomen and lateral tubercles on the side of the carapace. The only North American or Central American species known to have both these characters is M. lenca. The type locality of Ordgarius corpulentus is uncertain, because, as Banks himself pointed out, the collection was handled by G. Marx before being turned over to Banks after Marx’s death, and Marx locality labels are confused. Description. Female holotype. Carapace dark brown, with short white setae not covering tubercles; tubercles with light tips (Figs. 336, 337). Sternum dark orange. Legs dark brown. Median eyes on bulge, lateral eyes on bulges. Abdomen gray, with long humps (Fig. 339); venter with white square. First tarsus with S-shaped curva- => Figures 336-342. Mastophora corpulenta (Banks), female. 336, 337, carapace and chelicerae. 336, frontal. 337, lateral. 338, 339, carapace and abdomen. 338, dorsal. 339, lateral. 340-342, epigynum. 340, ventral. 341, posterior. 342, posterior, cleared. Figures 343-350. MM. rabida new species, female. 343, 344, carapace and chelicerae. 343, frontal. 344, lateral. 345, 346, carapace and abdomen. 345, dorsal. 346, lateral. 347-350, epigynum. 347, ventral. 348, posterior. 349, posterior, cleared. 350, seminal receptacle, median. MASTOPHORA ° Levi 365 Figures 351-361. M. escomeli new species. 351-357, female. 351, 352, carapace and chelicerae. 351, frontal. 352, lateral. 353, 354, carapace and abdomen. 353, dorsal. 354, lateral. 355-357, epigynum. 355, ventral. 356, posterior. 357, posterior, cleared. 358-360, male left palpus, stained. 358, apical. 359, mesal. 360, ventral. 361, ectal. Scale lines. 1.0 mm; genitalia, 0.1 mm. 366 ture. Total length 11.0 mm. Carapace 5.4 mm long, 5.2 wide in thoracic region, 2.8 wide behind posterior lateral eyes. First femur 4.7 mm, patella and tibia 7.6, meta- tarsus 5.4, tarsus 1.4. Second patella and tibia 4.8 mm, third 2.4, fourth 4.1. Males are not known. Diagnosis. Mastophora corpulenta is distinguished from M. diablo by the small dorsal knobs in the depression of the pos- terior of the epigynum (Fig. 341). The egg sac is fig-shaped, lacks lateral flaps, and has a thick stalk (Fig. 461). Distribution. Central America (Map AA). Specimens Examined. NICARAGUA Leé6n, Aban- gasca, 13 Dec. 1994, 1 subadult 2 (J. M. Maes, MCZ). Mastophora rabida new species Figures 343-350; Map 4A Holotype. Female holotype and immature female paratype from Rabida Island, Galapagos Islands, Ecuador, 12 May 1981 (Y. D. Lubin, 319), in MCZ. The specific name is a noun in apposition after the locality Note. The female holotype is a penulti- mate instar, ready to molt. The exuvium is loose above the epigynum. The epigynum is mature but not sclerotized. Description. Female holotype. Carapace dark orange. Chelicerae, labium, endites orange. Secu orange with white pig- ment. Coxae and distal leg articles dark dusky orange. Abdomen white with some faint gray marks (Fig. 345); venter with white square. Carapace appearing downy, covered with tubercles; tubercles with light tips (Figs. 343, 344). Median eyes on bulge, lateral eyes on bulges. Abdomen slightly wider than long wath small humps (Figs. 345, 346). Total length 7.7 mm. Car- Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 apace 3.1 mm long, 2.9 wide in thoracic region, 1.8 wide at lateral eyes. First femur 2.8 mm, patella and tibia 4.2, metatarsus 3.2, tarsus 1.0. Second patella and tibia 3.0 mm, third 1.7, fourth 2.8. Length of first patella and tibia 1.4 times width of cara- pace. Males are not known. Diagnosis. Mastophora rabida is distin- guished by the ventral loops of the slits on the posterior of the epigynum (Figs. 348, 349). Natural History. From Y. Lubin (per- sonal correspondence): “#319. nocturnal araneid on orb web. During day sits on twigs. 1 female, 1 juvenile. [NB: maybe it was the juvenile on an orb web? I didn't specify in the notes. YL]. #510. Tagus Cove, Isabella [This is a mangrove area].” Notes from field book 19: 30: “on Croton bush, hanging from thread with legs 3,4. Legs 1,2 Rell outwards, flexed. No Taelew: Spider Smerchcs legs forward when I hum, then moves to edge of leaf and adopts same posture.” Distribution. 4A), Galapagos Islands (Map Paratypes. Galapagos Islands: Isabella Island, Ta- gus Cove, on Croton scouleri at night, 13 May 1983, 1 imm. (Y. Lubin 510, MCZ). Mastophora escomeli new species Figures 351-361; Map 4A Escomel, 1918, Glyptocranium gasteracanthoides: 136 (misidentification). Holotype. Female holotype from Valle de Majes, nr. Arequipa, Depto. Arequipa, Peru, 1920 (E. Esco- mel), in MNHN. The species has been named after the collector and author of a paper on the venoms of Mastophora. Note. A similar specimen examined by —> Figures 362-365. Mastophora obtusa Mello-Leitao, immature female. 362, 363, carapace and chelicerae. 362, frontal. 363, lateral. 364, 365, carapace and abdomen. 364, dorsal. 365, lateral. Figures 366-372. WM. felis Piza, female. 366, 367, carapace and chelicerae. 366, frontal. 367, lateral. 368, 369, carapace and abdomen. 368, dorsal. 369, lateral. 370-372, epigynum. 370, ventral. 371, posterior. 372, posterior, cleared. Figures 373-379. MM. holmbergi Canals, female. 373, 374, carapace and chelicerae. 373, frontal. 374, lateral. 375, 376, cara- pace and abdomen. 375, dorsal. 376, lateral. 377-379, epigynum. 377, ventral. 378, posterior. 379, posterior, cleared. MASTOPHORA ° Levi 367 Figures 380-386. MM. reimoseri new species, female. 380, 381, carapace and chelicerae. 380, frontal. 381, lateral. 382, 383, carapace and abdomen. 382, dorsal. 383, lateral. 384-386, epigynum. 384, ventral. 385, posterior. 386, posterior, cleared. Scale lines. 1.0 mm; genitalia, 0.1 mm. 368 Canals (MACN) was labeled M. satan, presumably because of its long first legs. Description. Female holotype. Carapace orange-brown. Chelicerae brown. Labium, endites, sternum orange. Coxae orange, distal leg articles brown. Abdomen brown- ish, underlain by some white patches (Figs. 353, 354); venter with white square. Carapace with many large tubercles, tu- bercles on sides; tubercles with light tips, with short white setae between, but not covering tubercles, and longer white setae on sides (Figs. 351, 352). Median eyes on a bulge; lateral eyes on bulges. First tarsus with S-shaped curvature. Abdomen with long humps (Figs. 353, 354). Total length 13.5 mm. Carapace 5.0 mm long, 5.0 wide in thoracic region, 3.0 wide at lateral eyes. First femur 4. a mm, patella and tibia Wat metatarsus 6.3, tarsus 1.6. Second patella ands tibia 4.8 mm: third: 227. fourth 4.3. Length of first patella and tibia 1.5 times width of carapace. Male of uncertain affiliation. Color and shape as in other males. Total length 1.7 mm. Carapace 0.83 mm long, 0.81 wide in thoracic region, 0.54 wide a lateral eyes. First femur 0.72 mm, patella and tibia 0.81, metatarsus 0.49, tarsus 0.13. Second patella and tibia 0.71 mm, third 0.39, fourth 0.54. Length of first patella and tib- ia 1.0 times width of carapace. Note. Affiliation of males with females is uncertain. Variation. Total length of females 12.0— 13.5 mm. Length of free patella and tibia 15> e/atimmes Sadeh of carapace. The fe- male from Ica is more setose, with long setae on the legs and many shorter setae on the Aileen: The depressions of the female epigynum are larger and the bulge between the slits is less distinct. The illus- trations were made from the female ho- lotype. Diagnosis. Mastophora escomeli is dis- tinguished from M. gasteracanthoides by having a swelling betwecn the slits on the posterior of the “epigynum (Fig. 356) and having the atrium ventral to the seminal receptacles (Fig. 357). In M. gasteracan- Bulletin Museum of Comparative Zoology , Vols ISG, INO. 5 thoides, the slits are in a shared depression and the atria are dorsal to the seminal re- ceptacles (Figs. 415, 416). Natural History. Specimens were found on grapevines near Arequipa, and were known to readily bite grape workers as they pruned the plants, causing necrotic lesions. Distribution. Dry coastal region of Peru (Map 4A). Paratypes. PERU Lima: Lomas de Lachay, 26 May 1996, 12 (N. Llerana Martinez, MUSM). Arequipa: Arequipa, 1912, 22 (E. Escobal, MNHN); 12 (E. Escobal, MACN 4198). Specimens Examined. PERU Ica: Ica, 1992, 1° (Cascavilea-Rubio, MACN). La Libertad: Cerro Campana, N Trujillo, 23 May 1989, many 6, imm. (A. Salas, MUSM),. Mastophora obtusa Mello-Leitao Figures 362-365; Map 4C Mastophora obtusa Mello-Leitao, 1936: 134, fig. 2 imm. Immature holotype from Pesqueira, Pernam- buco, Brazil, in MNRJ, 41845, examined. Roewer, 1942: 955. Platnick, 2001. Glyptocranium obtusum:—Bonnet, 1957: 1998. Note. Pesqueira is located in the note with the description of M. pickeli. Description. Female holotype. Carapace reddish brown with white rim. Chelicerae, labium, endites brown. Sternum light brown. Coxae and distal leg articles brown. Abdomen very light brown; venter with white square. Carapace with short white setae on sides (Figs. 363, 364). Abdomen high with a pair of humps. Total length 4.8 mm. Carapace 1.8 mm long, 1.7 wide in thoracic region, 1.2 wide at lateral eyes. First femur 1.6 mm, patella and tibia 2.3, metatarsus 1.3, tarsus 0.5. Second patella and tibia 1.7 mm, third 0.8, fourth 1.8. Length of first patella and tibia 1.4 times width of carapace. Diagnosis. Although the type is imma- ture, M. obtusa is distinguished from many other species by the high abdomen and the humps on a joined swelling (Fig. 364). The high sides of the carapace, the shape of the tubercles, and the lack of pigment pattern on the abdomen suggest that the species belongs to the M. gasteracanthoi- des group of species. Perhaps this is an im- mature of M. satan. Specimens Examined. No other specimens have been found. Mastophora felis Piza Figures 366-372; Map 4C Mastophora felis Piza, 1976: 83, fig. 1. Female holo- type from Piracicaba, Sao Paulo, Broa, in MZAQ no. AO105, examined. Brignoli, 1983: 273. Platnick, 2001. Note. The holotype was embedded in difficult-to-remove fungal mycelium. Description. Female holotype. Carapace dark brown with tips of tubercles light and a thin white rim, each posterior median eye in a light patch. Chelicerae brown. La- bium, whites brown. Sternum brown. Coxae and distal leg articles orange-brown. Abdomen brownish gray (Figs. 368. 369): venter with a median white square. Cara- pace with long tubercles, the median of the horn’s base with multiple tubercles (Figs. 366, 367), covered with short white setae between tubercles. Median and lat- eral eyes on bulges. Legs with white setae. Abdomen with a pair of long dorsal humps (Fig. 369). Total length 13.0 mm. Cara- pace 6.3 mm long, 6.4 wide in thoracic re- gion, 3.8 wide at lateral eyes. First femur 5.8 mm, patella and tibia 10.6, metatarsus 8.8, tarsus 2.3. Second patella and tibia 6.7 mm, third 3.6. fourth 5.5. Length of first patella and tibia 1.6 times width of cara- pace. Males are not known. Variation. Total length of females 11.3— 13.0 mm. Length of first patella and tibia 1.4—1.6 times width of carapace. The illus- trations were made from the female ho- lotype. Diagnosis. Mastophora felis is distin- guished from all others having long wide humps and carapace with fabercles on sides and by having the atria of the epi- gynum ventral to fie seminal receptacles (at 11 and 1 h in Fig. 372) and the slits with a lateral lip (Fig. 371). The carapace tubercles are longer than those of M. satan and the posterior eyes are on light patches. MASTOPHORA °® Levi 369 Distribution. Rio de Janeiro and Sao Paulo states, Brazil (Map 4C). Paratypes. BRAZIL Rio de Janeiro: Santo Anténio, Rio Bonito [22°42'S, 42°37'W], 1933, 12 (S. Reme- tente, IBSP 418). Sdo Paulo: ?Campinas, July 1982, 12 (C. Froelich, IBSP 4968). Mastophora holmbergi Canals Figures 373-379; Map 4C Mastophora Holmbergi Canals, 1931: 22, figs. 1-5, pl. 3, fig. 5; pl. 4, figs. 7, 8, 2. Female from km 701, Santiago del Estero, in MACN, 24133 [7140], ex- amined. Mastophora holmbergi:—Mello-Leitao, 1931, 73, figs. 10, 21, 2. Roewer, 1942, 900. Platnick, 2001. Glyptocranium holmbergi:—Bonnet, 1957, 1997. Description. Female holotype. Carapace dark red-brown to black, light transverse band in front of posterior median eyes. Sternum orange. Legs orange-brown. Ab- domen with Pant pattern (Figs. 375); ven- ter with white square. Carapace with many tubercles with light tips especially on sides, and a few downy setae; lateral eyes on bulges (Figs. 373, 374). First tarsus slightly S-shaped. Abdomen with narrow humps. Total length 11.0 mm. Carapace 5.2 mm long, 5.3 made in thoracic region, 3.0 wide at later al eyes. First femur 5.7 mm, patella and tibia 10.0, metatarsus 9.2, tarsus 2.0. Second patella and tibia 6.2 mm, third 3.2, fourth 5.0. Length of first patella and tibia 1.9 times width of carapace. Males are not known. Variation. Total length of females 11.0— Se) mana, ene ‘ilmecaciens were made from the female holotype. Diagnosis. Unlike Mastophora reimo- seri, ih holmbergi has long first legs. In the epigynum fhe atria are sable in ven- tral view (Fig. 377) and the slits are par- allel, but at their ventral ends the slits bend toward each other (Fig. 378). Distribution. Paraguay, to Santiago del Estero, Argentina (Map 4C). Specimens Examined. PARAGUAY Rca. del .. . [il- legible], Nov. 1940, 12 (Cranwell-Giai, MACN 1630). 370 Mastophora reimoseri new species Figures 380-386; Map 4C Holotype. Female holotype from Asuncion, Paraguay, 1908 (E. Reimoser), in NHMW. The species has been named after the collector and Austrian arach- nologist, E. Reimoser. Description. Female holotype. Speci- men faded. Carapace yellow-brown. Che- licerae, labium, endites yellow-brown. Sternum, legs golden brown. Carapace with many tubercles and short setae, clyp- eus with longer setae (Figs. 380, 381). Me- dian eyes and lateral eyes on bulges. First femora bent at their distal ends with long setae at point of greatest curvature. Ab- domen with erennict humps and short se- tae, base of humps with longer setae (Figs. 382, 383). Total length 8.5 mm. Carapace 4.0 mm long, 4.0 wide in thoracic region, 2.3 wide at lateral eyes. First femur 3.7 mm, patella and tibia 6.2, metatarsus 4.5, tarsus 1.2. Second patella and tibia 4.0 mim, third 2.1, fourth 3.5. First patella and tibia 1.6 times width of carapace. Males are not known. Diagnosis. The species is distinguished by ies musical note—shaped Pree of the slits in the epigynum (Fig. 385), and by having shorter legs than M. holmbergi. Distr ibution. Known only from Roun: cion, Paraguay (Map 4C). Specimens Examined. No other specimens have been found. Mastophora satan Canals Figures 387-398, 462, 463; Map 4B Mastophora satan Canals, 1931: 25, figs. 1-5, pl. 3, fig. 6, 2. Female holotype from La Rioja, Argen- tina, in MACN, 5260, examined. Mello-Leitio, 1931: 73, figs. 11, 22, 2. Roewer, 1942: 901. Plat- nick, 2001. Glyptocranium satan:—Bonnet, 1957: 1998. Description. Female from Cérdoba. Carapace dark brown with many short light setae and narrow white rim. Sternum red-brown. Legs red-brown with long white setae. Abdomen whitish with dusk- iness on humps and sides (Fig. 389); ven- ter with white square. Carapace with many tubercles, lateral eyes on bulges, median Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 eyes on bulge (Figs. 387, 388). Abdomen humps very long (Figs. 389, 390). Total length 14.0 mm. Carapace 6.4 mm long, 6.6 wide in thoracic region, 3.5 wide at lat- eral eyes. First femur 7.0 mm, patella and tibia 11.5, metatarsus 10.0, tarsus 2.2. Sec- ond patella and tibia 7.0 mm, third 3.3, fourth 5.7. Length of first patella and tibia 1.7 times width of carapace. Male from Rio Grande do Sul, Brazil. Carapace beige with a triangular white patch. Seas legs beige. Abdomen whitish. Carapace with four tubercles: ab- domen with two humps. Total length ed mm. Carapace 0.92 mm long, 0.81 wide in thoracic region, 0.58 wide a lateral eyes. First fonatin 0.87 mm, patella and tibia 0.92, metatarsus 0.49, tarsus 0.31. Second patella and tibia 0.74 mm, third 0.44, fourth 0.55. Note. The association of male and fe- male is uncertain. The male is placed with the most common species in the area; also, a female of the species has been collected at the collecting site. Variation. Total length of females 9.7— 17.5 mm. The holotype is 11.2 mm total length, carapace 5.7 mm wide and long, the first patella and tibia 9.7. Length of first patella and tibia 1.4—-1.6 times cara- pace width in females from Brazil, 1.7 in specimens from Uruguay, 1.5 from Men- doza, 1.6 from La Pampa, 1.5 from Entre Rios. The illustrations were made of a fe- male from Cérdoba Province, Argentina, a female from La Rioja Province (Fig. 394), a female from Tucumaén Province (Fig. 395), and of the male from Rio Grande do Sul, Brazil. Diagnosis. Mastophora satan is separat- ed fon M. gasteracanthoides and M. dia- blo by the long first tibia and metatarsus, each 9.7 mm or longer. The epigynum dif- fers from that of M. diablo by showing the atria as a dark patch in the dorsal slope of a depression (Figs. 392, 394, 395), whereas in M. satan, atria are outside and lateral to the depression (Figs. 404, 406). The epi- gynum is similar to that of M. gasteracan- fhoiles but differs in ventral view, where MASTOPHORA ° Levi 371 ae M10) ds Figures 387-398. Mastophora satan Canals. 387-395, female. 387, 388, carapace and chelicerae. 387, frontal. 388, lateral. 389, 390, carapace and abdomen. 389, dorsal, with male. 390, lateral. 391-395, epigynum. 391, ventral. 392, 394, 395, posterior. 393, posterior, cleared. 392, 393, (Cordoba). 394, (holotype from La Rioja). 395, (Tucuman). 396-398, male left palpus. 396, mesal. 397, ventral. 398, ectal. Figures 399-407. M. diablo new species, female. 399, 400, carapace and chelicerae. 399, frontal. 400, lateral. 401, 402, carapace and abdomen. 401, dorsal. 402, lateral. 403-407, epigynum. 403, ventral. 404, 406, posterior. 405, 407, posterior, cleared. 404, (Chaco). 406, 407, (Santiago del Estero). Scale lines. 1.0 mm; genitalia, 0.1 mm. 372 M. gasteracanthoides has a distinct sclerite a short distance anterior to the posterior edge (Fig. 414), whereas M. satan has the whole median area slightly sclerotized (Fig. 391). The median apophysis of the palpus of the male (Fig. 398) is more curved than that of M. gasteracathoides (Fig. 420). The egg sac lacks flaps and has only a short stalk (Figs. 462, 463). Distribution. From Pernambuco State, eastern Brazil, to La Pampa Province, Ar- gentina (Map 4B). Specimens Examined. BRAZIL Pernambuco: Tap- era, 12 (B. Pickel, MNRJ 391). Bahia: Feiora de San- tana, July 1994, 12 (S. D. Cunha, IBSP 16246). Sao Paulo: Santo André, 16 June 1965, 12 (L. Daga, IBSP 1931); Seminario Santa Terezinha, Tieté, 5 May 1953, 12 (IBSP 887); Sao José do Rio Preto, 9 Feb. 1964, 12 (Vizotto, MZSP 3471): 1 June 1964, 1 egg sac, 12 (Vizotto, MZSP 3470). Santa Catarina: Bla menau, 12 (NHMW). Rio Grande do Sul: 12 (P. Buck, MNRJ 41644); Rodeio Bonito, Bogé, 5 June 1980, 12 (E. W. Aguiar, MCN 9103); Canela, 10 Feb. 1966, 29 (A. A. Lise, MCN 0752); Sao Leopoldo, 5 Mar. 1965, 12 (C. Valle, MZSP 4797); Porto Alegre, 12 Apr. 1926, 12 (R. Gliesch, MNRJ 392); Parque Zoologico, Sapucaia do Sul, 9 Dec. 1985, 22 (A. E. Tovares, MZSP 14079); Porto Alegre, 17 Mar. 1955, 1 imm. (T. de Lema, MCN 01628); 16 June 1963, 1° (A. Lise, MCN 01820): 6 Oct. 1988, 12 (R. Villanova, MCP 105); Belem Velho, Porto Alegre, 17 July 1979, 12 (V. Mott, MCN 2608): Morro Santana, Porto Ale- gre, 5 May 1984, 1d (S. M. Silva, MCN 12204); 13 Sept. 1984, 12 (A. A. Lise, MCN 29426); Santa Ma- ria, Aug. 1986, 12 (MCP 10340); Santo Anténio da Patulha, 30 Oct. 1980, 12 (T. K. Moreira, MCN 9456). URUGUAY Villasboas, 1953, 12 (L. Lecour, FCMU); Paso del Cerro (Artigas), May 1956, 192 (C. Fuques, FCMU)); Artigas, Sept. 1959, 12 (C. Fuques, FCMU): Mar. 1965, 12 (C. Fuques, FCMU 1965); Ruta 3, Salto, 3 Aug. 2001, 1 (V. Vazquez, Williams, FCMU 562); Cuareim, Espinillares, Artigas, 12 Mar. 1956, 12 (C. Fuques, FCMU 296). ARGENTINA Misiones: 1940, 12 (Exp. Zotla. Armanini, MACN 2050). Catamarca: Sijan, Nov. 1964, 12 (Ahumada, MACN). La Rioja: 22 (Sr. Giacomelli, MACN 4186). Tucumdn: Tucuman, 12 Dec. 1984, 12 (FMLT 2159). Santiago del Estero: Santiago del Estero, 20 Apr. 1958, 19 ( (J. W. Abalos, MACN); La Banda, 1958, 12 (J. Abalos, MACN); July 1958, 12 (D. Bravo, MACN): Tabla Redonda Depto. Banda, 23 Dec. 1959, 12, egg sacs, imm. (J. W. Abalos, MACN). Mendoza: Mendoza, 1907, 12 (E. Reimoser, NHMW); San Rafael, Feb. 1940, 12 (D. Pereyra, MACN 1799). Cordoba: Mina Clavero, April 1973, 12 (Stirbel, MACN). Entre Rios: Concordia, 1931, Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 12 (MNBJ 57953). Santa Fé: Santa Fé, 1931, 12 (M Birabén, MNRJ 522). Buenos Aires: Ireneo Portela, 30 April 1922, 12 (Scheimer, MACN). La Pampa: General Pico, Feb. 1952, 12 (Williamson, MACN); 30 Mar. 1958, 22, doubtful determination (William- son, MACN). Mastophora diablo new species Plate 1; Figures 399-407, 464; Map 4D Holotype. Chaco, Argentina, MACN, no. 5432. Mastophora gasteracanthoides:—Mello-Leitao, 1931: 69, in part. Canals, 1931: 19 (misidentified gaster- acanthoides Nicolet). : Female holotype from Colonia Benitez, Sept. 1959 (Bachmann) in Note. Most specimens of this species had been misidentified as M. gasteracan- thoides in collections. Description. Female holotype. Carapace brown, with many downy short setae and narrow white rim. Sternum brown. Legs brown with long white setae. Abdomen brownish white; venter with white square. Abdomen humps long (Figs. 401, 402). Total length 13.0 mm. Carapace 5.3 mm long, 5.8 Sade in thoracic region, 3.2 wide at Teter al eyes. Abdomen 15 mm _ high. First femur 5.5 mm, patella and tibia 8.6, metatarsus 6.3, tarsus 1.3. Second patella and tibia 5.7 mm, third 3.0, fourth 4.8. Length of first patella and Abie 1.5 times wath of carapace. Males are not known. Variation. Total length of females 10.3— 16.7 mm. Length of feet patella and tibia LSS etimes sate of carapace. Figures 399-405 were made from the fannie ho- lotype; Figures 406 and 407 were from specimens from Santiago del Estero. Diagnosis. Differs from M. satan in hav- ing first tibia less than 9 mm total length and from both M. satan and M. gastera- canthoides by having the atria outside and lateral to the depressions in posterior view of the epigynum (Figs. 404407). The egg sac lacks flaps and has a heavier stalk (Fig. 464) than that of M. satan. Natural History. A large syrphid fly (14 mm total length) was collected in Moreno, Buenos Aires, with one adult. MASTOPHORA ° Levi 373 Figures 408-421. Mastophora gasteracanthoides (Nicolet). 408-416, 421, female. 408, 409, carapace and chelicerae. 408, frontal. 409, lateral. 410-413, carapace and abdomen. 410, 412, dorsal, with male. 411, 413, lateral. 410, 411, (Santiago). 412, 413, (Chillan). 414-416, epigynum. 414, ventral. 415, posterior. 416, posterior, cleared. 417-420, male left palpus. 417, apical. 418, mesal. 419, ventral. 420, ectal. 421, female, frontal view with right legs. Scale lines. 1.0 mm; genitalia, 0.1 mm. (ey) ~ Distribution. Northern and central Ar- gentina to La Pampa Province (Map 4D). Paratypes. ARGENTINA Formosa: Las Lomitas, Oct. 1966, 12 (A. Vogt, MACN). La Rioja: 29 (Prof. Gémez, MACN 4187). Tucumdn: El Timbé, 27 May 1952, 22 (J. Campos, FMLT 994); Los Bosques, 12 BMNH); Dept. Burruyaco, E] Haranjo, 13 June 1964, 12 (M. Inés Cortez, FML 1693). Santiago del Estero: El Zanj6n, 5 June 1960, 12 (J. W. Abalos, MACN); Fernandez, 4 Apr. 1960, 12 (J. W. Abalos, MACN); Frias, 2 Oct. 1970, 12 (J. W. Abalos, MACN); La Banda, 24 May 1958, 1¢ (J. Areas, MACN); Los Juries, March 1959, 12 (L. Remedi, MACN); Santiago del Estero, 18 Sept. 1958, 1 (D. Luna, MACN); 1959, 1° (J. W. Abalos, MACN). Cor- rientes: Bella Vista, Nov. 1944, 12 (Silberman, MACN). Entre Rios: Basavilbaso, 12 (U. Podesta, MACN 4189). Santa Fé: Roldan, June 1943, 12 (Es- cuela 230, MACN). Cordoba: Argiiello, Feb. 1946, 12 (J. A. De Carlo, MACN 1657); Bajo Grande, 12 MLP 15563). Buenos Aires: Castela, 27 July 1958, 12 (Ing. Favret); Moreno, Jan. 1946, 192 (R. D. Schiapelli, MACN 1658). La Pampa: Mira Pampa, April 1949, 12 (C. Vigliorcho, MLP 16638); General Pico, March 1951, 12 (C. Ballani, MLP 13642). Mastophora gasteracanthoides (Nicolet) Figures 7, 408-421, 464; Map 4B Epeira gasteracanthoides Nicolet, 1849: 485, pl. 5, fig. 7a, b, 2. Specimens from gardens and fields of ental provinces, Santiago, Chile, in MNHN, lost. Glyptocranium gasteracanthoides:—Simon, 1895: 882, fig. 946, 2. Bonnet, 1957: 1997. Mastophora_ gasteracanthoides:—Porter, 1918: 139. Roewer, 1942: 901. Archer, 1963: 19. Platnick, 2001. Mastophora gasteracanthoides oxalidis Archer, 1963: 16. Female holotype, males, and imm. from Loma de Pefiuelas, 6 km al sur de La Serena, Coquimbo, in AMNH, examined. Platnick, 2001. NEW SYN- ONYMY. Note. The short humps of recently col- lected specimens from Santiago (Figs. 410, 411) differ from those of ie specimens illustrated by Nicolet (1849) and Simon (1895), which have higher humps. Older specimens kept in MNHN from Chillan (Figs. 412, 413) had longer humps but 4 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 genitalia and carapace similar to those of recently collected specimens. The subspecies named by Archer (1963) had no diagnosis. Description. Female from Santiago. Carapace brown, tubercles lighter. Chelic- erae lighter. Sternum brown. Legs lighter brown. Abdomen brownish white with some asymmetrical darker patches and humps (Figs. 410, 411); venter with white square. Total length 12 mm. Carapace 4.8 mm long, 4.8 wide in thoracic region, 2.5 wide at lateral eyes. First femur 4.0 mm, patella and tibia 6.3, metatarsus 4.8, tarsus 1.3. Second patella and tibia 4.3 mm, third 2.6, fourth 4.0. Length of first patella and tibia 1.3 times eden of carapace. Male. Carapace brown, with white me- dian band covering the two median tuber- cles and the pair of horns. Sternum, legs brown. Abdomen dusky white anterior, venter with indistinct white spots. Abdo- men without humps (Fig. 7). The palpus has one weak patellar seta. Total length 2.3 mm. Carapace 0.9 mm long, 0.9 wide in thoracic region, 0.7 wide at lateral eyes. First femur 0.9 mm, patella and tibia 1.0, metatarsus 0.6, tarsus 0.3. Second patella and tibia 0.9 mm, third 0.5, fourth 0.7. Length of first patella and tibia 1.1 times width of carapace. Note. Males came out of an egg sac col- lected with females. Variation. Total length of females 9.6— 13.5 mm. The illustrations of the female are made from a female from Santiago, ex- cept Figures 412 and 413, which are of a female from Chillan. Diagnosis. Mastophora gasteracanthoi- des differs from M. diablo by having the atria show as dark spots in the posterior slope of the depression in posterior view of the epigynum (Fig. 415), and from M. > Figures 422-433. Ordgarius magnificus (Rainbow). 422-428, female. 422, 423, carapace and chelicerae. 422, frontal. 423, lateral. 424, 425, carapace and abdomen. 424, dorsal, with male. 425, immature, lateral. 426-428, epigynum. 426, ventral. 427, posterior. 428, posterior, cleared. 429-433, male. 429, carapace and chelicerae, lateral. 430, dorsal. 431-433, left palpus. 431, mesal. 432, ventral. 433, ectal. MASTOPHORA ® Levi 375 5 422 ie « S Wrnagaltiels Cladomelea akermani Figures 434-444. Cladomelea akermani Hewitt. 434-440, female. 434, 435, carapace and chelicerae. 434, frontal. 435, lateral. 436, 437, carapace and abdomen. 436, dorsal, with male. 437, lateral. 438-440, epigynum. 438, ventral. 439, posterior. 440, posterior, cleared. 441-444, male. 441, dorsal. 442-444, left palpus. 442, mesal. 443, ventral. 444, ectal. Scale lines. 1.0 mm; genitalia, 0.1 mm. 376 satan by having shorter first legs and a dis- tinct sclerotized plate on the epigynum, anterior of the posterior margin, in ventral view (Fig. 414). The palp of the male has a median apophysis straighter (Fig. 420) than that of M. satan (Fig. 398). The egg sac (Fig. 465) lacks flaps and oo has a thin stalk. Distribution. Central Chile (Map 4B). Specimens Examined. CHILE 12 (MNHN, 114). Coquimbo: 6 km S of La Serena, 23-30 Nov. 1961, 12, egg sac with d and imm. 2 (A. F. Archer, AMNH#). Valparaiso: Quilput [Quilpué], 1904, 12 (C Porter, MNHN 23457). Metropolitana: Santiago, 4 June 1947, 1 imm. (R. Donoso, AMNH); Feb. 1955, 1? (I. Pedag., AMNH); Quilicura, Oct. 1979, 12 (L. Pena, AMNH), Aug.—Sept., 2 imm. (L. E. Pefia, AMNH); Renca, W of Santiago, 2 Oct. 1984, 1? (L. E. Pea, AMNH); Los Espejo, 6 Nov. 1973, 12 (C. L. Cartagena, MCZ). Nuble: Chillan, 12 (Delfin, MNHN, 23520). Ordgarius Keyserling Ordgarius Keyserling, 1886: 114. Type species by monotypy, M. monstrosus Keyserling, 1886, from Queensland, Australia. Neave, 1940: 453. Roewer, 1942: 902. Bonnet, 1958: 3200. Davies, 1988: 318, mea, BO, OU 2, Cre Euglyptila Simon, 1908: 151. Type species designated by Bonnet, 1956, E. acanthonotata from Tonkin [northern Vietnam]. Neave, 1939b: 325. Roewer, 1942: 903. Bonnet, 1956: 1810. NEW SYNONY- MY. Note. Two species of Euglyptila were described by Simon, both from immatures. Euglyptila acanthonotata is in MNHN, lost; the other, E. nigrithorax, 2.5 mm total length was found, described, and illustrat- ed by Emerit (1980). It is an immature of Ordgarius sexspinosus Thorell, 1894, total length 14 mm, and is illustrated by Yin, 1997: 384. NEW SYNONYMY. Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 Ordgarius magnificus (Rainbow) Figures 422-433, 466 Dicrostichus magnificus Rainbow, 1897: 523, pl. 17, fig. 8, 2. Holotype from Mount Kembla, New South Wales, Australia, not examined. Roewer, 1942: 900. Ordgarius magnificus:—Davies, 1988: 316. Description. Female. Carapace yellow- ish, dark brown in groove between ce- phalic and thoracic areas, a dark band above eyes, tubercles white. Sternum light orange. Coxae and distal leg articles light orange. Abdomen yellowish white with ventral white square. Carapace with me- dian eyes on stalk, with horns quite small (Figs. 422, 423). Eyes subequal, median eye area almost square. Chelicerae fang groove with three anterior teeth, one small posterior tooth. Abdomen narrowing to posterior end, with tubercles (Figs. 424, 425). Total length 13.5 mm. Carapace 6.3 mm long, 6.7 wide in thoracic region, 3.3 wide at lateral eyes. First femur 6.0 mm, patella and tibia 7.3, metatarsus 4.7, tarsus 1.4. Second patella and tibia 7.0 mm, third 3.9, fourth 5.8. Length of first patella and tibia 1.1 times width of carapace. Male. Carapace beige, white pigment spots at base of spines. Sternum dark brown. Legs light beige. Dorsum of ab- domen white, enter black. Carapace (Fig. 429), abdomen as in Figure 430. Row of setae on tarsi as in male “Mastophora. Pal- pal patella with no setae. No endite tooth, no coxal hook. Total length 1.7 mm. Car- apace 0.88 mm long, 0. 81 wide in thoracic region, 0.52 wide at lateral eyes. First fe- mur 0.78 mm, patella and tibia 0.89, meta- tarsus 0.57, tarsus 0.30. Second patella and tibia 0.78 mm, third 0.39, fourth 0.59. Length of first patella and tibia 1.1 times sae of carapace. = Figures 445-468. Egg sacs of Mastophora species, including species of Ordgarius and Cladomelea. 445, M. timuqua. 446, M. bisaccata. 447, M. stowei. 448, M. yeargani. 449, 450, M. phrynosoma. 449, (Kentucky). 450, (Florida). 451, M. carpogastra. 452, M. vaquera. 453, 454, M. hutchinsoni. 453, (after Kaston, 1981). 454, (New Jersey). 455, M. cornigera. 456, M. archeri. 457, M. fasciata. 458, M. abalosi (after Urtubey and Baez, 1983). 459, M. comma (after Baez and Urtubey, 1985). 460, M. extraordinaria. 461, M. corpulenta. 462, 463, M. satan. 462, (Santiago del Estero). 463, (Rio Grande do Sul). 464, M. diablo. MASTOPHORA ® Levi SU yeargani stowei timuqua 453 hutchinsoni archeri corpulenta abalosi Ordgarius \ 466 magnificus | ' 467 Cladomelea akermani gasteracanthoides saian 465, M. gasteracanthoides. 466, Ordgarius magnificus (after Davies, 1988). 467, O. monstrosus (after Davies, 1988). 468, Cladomelea akermani. Scale lines. 1.0 mm; all except 466 and 467 are approximately the same magnification. 378 Genitalia. The epigynum like Mastopho- ra in having only a posterior lobe ventrally (Fig. 426) and having two slits and diag- nostic sculpturing on the posterior face (Fig. 427) and having indistinct atria (Fig. 428). The palpus of the male has no con- ductor but has the tip of the tegulum sclerotized (at 11 h in Fig. 433) and the median apophysis more sclerotized than that of Mastophora (Figs. 431-433). Variation. The female examined came from Olderley, Brisbane, Queensland, the male from Mulgowie, SE Queensland (QMB). Natural History. Unlike most Masto- phora, this species ties leaves together and may have a diurnal retreat. The female uses the second leg to swing a bolas. The egg sac of O. magnificus is spindle-shaped (Fig. 466), that of O. monstrosus resem- bles those of Mastophora and has minute flaps (Fig. 467). Distribution. Australia. Cladomelea Simon Cladomelea Simon, 1895: 886, figs. 949, 950. Type species by original designation Cyrtarachne longi- pes O. P.-Cambridge, 1877: 559, from West Africa. Neave, 1939a: 750. Roewer, 1942: 900. Bonnet, 1956: 1097. Note. Cladomelea longipes is very simi- lar to C. akermani. Cladomelea akermani Hewitt Figures 434-444, 468 Cladomelea akermani Hewitt, 1923: 63, figs. 4,5, 2. Female holotype from Pietermaritzburg, Natal, not examined. Roewer, 1942: 500. Leroy, Jocqué, and Leroy, ICES I, S. S Description. Female. Carapace light or- ange-brown, distal ends of projections black. Chelicerae, labium, endites orange. Sternum light orange-brown. Coxae or- ange-brown; legs brown. Dorsum of ab- domen whitish with a pair of brown tu- bercles (Fig. 436); venter whitish with a median, transverse, white rectangle. Car- apace with median eyes on a bulge, three projections and long white setae, no horns (Figs. 434, 435). Height of clypeus equals Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 about five diameters of anterior median eye. Abdomen widest in middle, dorsum with numerous rounded tubercles, not completely symmetrical (Figs. 436, 437). Total length 15.5 mm. Carapace 5.4 mm long, 5.2 wide, 2.3 wide at lateral eyes. First femur 6.7 mm, patella and tibia 10.8, metatarsus 8.4, tarsus 1.4. Second patella and tibia’ 7.3 mm: third 3.3) fourth) 4976 Length of first patella and tibia 1.9 times width of carapace. Male. Carapace, labium, endites, ster- num dark brown. Coxae, distal leg articles light. Dorsum of abdomen maculated black, gray, and white (Fig. 441); venter dark brown. Carapace rugose, without tu- bercles, posterior area swollen. Height of clypeus equals 1.8 diameters of anterior median eye. Endite without tooth. Palpal patella without macroseta. First coxa with- out hook. Row of setae on tarsi, as in male Mastophora. Abdomen widest in middle, dorsum sclerotized with three humps and two pairs of sclerotized discs (Fig. 441). Total length 1.6 mm. Carapace 0.94 mm long, 0.72 wide, 0.51 wide at lateral eyes. First femur 0.80 mm, patella and tibia 0.91, metatarsus 0.48, tarsus 0.34. Second patella and tibia 0.70 mm, third 0.41, fourth 0.57. Length of first patella and tib- ia 1.3 times width of carapace. Genitalia. The epigynum is as in Mas- tophora, having only a posterior lip ven-_ trally (Fig. 438) and having two slits and diagnostic sculpturing on the posterior face (Fig. 439) and tiny atria (Fig. 440). | The palpus of the male has a distinct con-— ductor supporting the embolus (at 1 h in Fig. 443) and the median apophysis more sclerotized than that of Mastophora (Figs. 449-444), | Variation. A second female from Pieter- | maritzburg, Natal (AMNH), examined had | the height of the clypeus only four diam- | eters of the anterior median eye. Female | examined and illustrated from Umgeni | Valley Reservation, Kwa—Zulu—Natal, | South Africa, the male came from Umgeni Valley project near Howick, South Africa | (NMP). | | | | I i Natural History. The spider is found in grasslands on grass of the Kwa—Zulu—Natal area around Pietermaritzburg, South Af- rica (Leroy et al., 1998). The egg sac is drop-shaped and attached to a grass blade (Fig. 468). The female Handles the bolas with a second leg and swings it in a hori- zontal plane. Distribution. South Africa. LITERATURE CITED ALVAREZ DEL Toro, M. 1992. Arafias de Chiapas. Chiapas, México: Universidad Aut6noma de Chiapas, Tuxtla Gutiérrez. 297 pp. AMERICAN GEOGRAPHICAL SOCIETY. 1944. Index to Map of Hispanic America. Vol. 5. Geographical Names in Brazil. Washington, DC: U.S. Govern- ment Printing Office. ARCHER, A. F. 1963. Catalogo de las arafias Chilenas de las familias de la division Metarachnae. 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Nouvelles espéces d’Argiopidae, Memorias del Instuto de Butantan, Sao Paulo, 3: 327-331. YEARGAN, K. V. 1988. Ecology of a bolas spider, Mas- tophora hutchinsoni: phenology hunting tactics, and evidence for aggressive chemical mimicry. Oecologia, 74: 524-530. . 1994. Biology of bolas spiders. Annual Re- view of Entomology, 39: 81-99. YEARGAN, K. V., AND L. W. QUATE. 1996. Juvenile bolas attract psychodid flies. Oecologia, 106: 966-271. . 1997. Adult male bolas spiders retain juve- nile hunting tactics. Oecologia, 112: 272-276. YIN, CHANGMIN. 1997. Arachnida, Araneae: Aranei- dae. Beijing, China: Fauna Sinica, Academia Sin- ica, Science Press. 460 pp. 382 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5 INDEX Valid names are printed in italics. Page numbers refer to main references, starred page numbers to illus- trations. abalosi, 355*, 356, 377* Acacesia, 315 Acantharachne, 314 Acantharanea, 314 Agathostichus, 315 Agatostichus, 315 akermani, 375*, 377*, 378 alachua, 329*, 330 Alpaida, 315 altiventer, 325 alvareztoroi, 359*, 360 apalachicola, 326, 327* Araneus, 315 archeri, 346, 347*, 377* bicurvata, 345 bisaccata, 327*, 330, 331*, 377° bisaccatum, 330 brescoviti, 361*, 363 carpogaster, 340 carpogastera, 340 carpogastra, 327*, 340, 377* carpogastrum, 340 catarina, 335*, 339 Celaenia, 315-317 Cladomelea, 314, 316-18, 378 Coelossia, 314 comma, 355*, 356, 377* conifera, 361*, 364 coniferus, 364 cornigera, 343*, 344, 377* cornigera, 342 cornigerum, 345 cornigerus, 345 corpulenta, 364, 365*, 377* corpulentum, 364 corpulentus, 364 corumbatai, 335*, 339 cranion, 349*, 351 Cyrtarachne, 315-317 Cyrtophora, 315 diablo, 32:7) Siles, SOS ont Dicrostichus, 314 dizzydeani, 349*, 350 Eriophora, 315 escomeli, 365*, 366 Euglyptila, 376 Exechocentrus, 314 extraordinaria, 355*, 357, 377* fagoides, 340 fasciata, 347*, 348, 377* fasciolata, 318, 325 felda, 326, 327* felis, 367*, 369 gasteracanthoides, 327*, 373*, 374, 377* gasteracanthoides, 366, 372 gasteracanthoides oxalidis, 374 Glyptocranium, 315 haywardi, 335*, 339 Heterocephala, 315 holmbergi, 367*, 369 hutchinsoni, 342, 343*, 377* intermedia, 72 Kaira, 315-317, 325 lara, 335*, 340 lenca, 364 leucabulba, 358, 359%* leucabulbus, 358 leucacantha, 361*, 362 leucacanthus, 363 longiceps, 351, 353* magnificus, 375*, 376, Mastophora, 315-317 melloleitaoi, 355*, 356 Metepeira, 315 monstrosus, 377* multilineata, 330 obesus, 330 obtusa, 367*, 368 | obtusum, 368 | occidentalis, 348 | Ocrepeira, 315 Ordgarius, 314, 316, 317, 376 | Pasilobus, 315-318 pesqueiro, 352, 353* phrynosoma, 333*, 336, 377* pickeli, 349*, 350 pickeli occidentalis, 348 piras, 352, 353* Poecilopachys, 315-317 rabida, 365*, 366 reimoseri, 367*, 370 satan, 370, 371*, 377* satsuma, 325, 327* seminole, 337*, 341 soberiana, 361*, 362 stowei, 331*, 334, 377* Taczanowskia, 315-317 timuqua, 328, 329*, 377* vaquera, 337*, 342, 377* yacare, 354, 355* yeargani, 333*, 336, 377* ypiranga, 353*, 354 > ne ri A MONOGRAPHIC REVISION OF THE ANT GENUS PRISTOMYRMEX a . (HYMENOPTERA: FORMICIDAE) MINSHENG WANG 97 2003 VOLUME 157, NUMBER 6 10 April 2003 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY BreviorA 1952— BULLETIN 1863— Memoirs 1865-1938 JouNsonia, Department of Mollusks, 1941-1974 OCCASIONAL PAPERS ON Mo.uusks, 1945-— SPECIAL 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, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4, Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology 4 and Evolutionary Biology: Essays in Honor of Emest E. Williams. 729 pp. 6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. ee ey ee RUPEE AONE AERTS a Ea Hi or Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the 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 of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter's Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1947. (Complete sets only.) Proceedings of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2003. A MONOGRAPHIC REVISION OF THE ANT GENUS PRISTOMYRMEX (HYMENOPTERA: FORMICIDAE) MINSHENG WANG' CONTENTS No Strac tate ek eee au eee as oe Lae RE RS 383 Tiniirahinetivor 383 (GOW ESCH OTS) meee ie ene ee A 385 Measurements and Indices _...- 385 A Brief History of the Genus Pristomyrmex _. 386 Genus eristomunmnes, Navi een ere 387 List of Pristomyrmex Names with Synonymies 392 Key to the World Species of Pristomyrmex (Workers) eae eet et ee ee 393 Mherpunctatuss Groupee 404 Pristomyrmex OIIBU SUIS Sho hg ere 404 Pristomyrmex fossulatus (Forel)... 406 ristOmummer PUlCheTe Spos M.)) sees teres 408 Pristomyrmex punctatus (F. Smith) 410 BRiSCOMUTMEX MUA USS. (IN esate tee teseen 415 Whererbranius Group oe ee 417 Pristomyrmex cribrarius Arnold 418 itherquadridens Group) == ee 420 Pristomyrmex africanus Karavaiev 423 Pristomyrmex bicolor Emery stat. n. 425 Pristomyrmex brevispinosus Emery 428 Pristomyrmex collinus sp. n. 432 Pristomyrmex costatus sp. 0 434 Pristomyrmex curvulus SPDEAli ghosts seelel en aa 437 nristomunmex edad Morel) mars: sen 440 Pristomyrmex erythropygus Taylor 44] PiPSHOUMUIT UD PE JUGS Os WN ee 443 Pristomyrmex foveolatus Taylor 446 Pristomyrmex UT SUTUS ES |) 1) ae eae 449 Pristomyrmex lometsmimitsssp sails eens 450 Hei StOMY TEL IMO CES CUS) S|) IN. yeeen eae eee 452 Pristomyrmex nitidissimus Donisthorpe —. 453 Pristomyrmex OCCULEUSTC Sera as sere eneaae 455 Pristomyrmex orbiceps (Santschi) 456 Pristomyrmex quadridens Emery — 459 Pristomyrmex quadridentatus (André) 463 Pristomyrmex quindentatus sp. n. 467 Pristomyrmex sulcatus Emery stat. n. _......... 469 Pristomyrmex thoracicus Taylor. 473 Pristomyrmex trachylissus (F. Smith) 474 istomu mime, (hOZOTm BOOM = 2a ieee ee 476 ‘Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts 02138. Bull. Mus. Comp. Zool., Pristomyrmex wheeleri Taylor 478 Pristomyrmex wilsoni Taylor 481 Mhextnispinosuss Group pe ere ee 483 Pristomyrmex bispinosus (Donisthorpe) este Pristomyrmex browni sp. n Pristomyrmex trispinosus (Donisthorpe) RACE 488 Mhevepisatusn Group) semis enema ene 489 Rristomueynnnen (ACCT OSUS)S)) 410.) eee ee 491 Rristomuy nme: DOLLOTL ES) \sgesee seems 492 Pristomyrmex coggii Emery —--------- 493 Pristomyrmex inermis sp. Mi. 2. 496 RristomumimexciaT SUS espe ah) ae eee 497 Pristomyrmex levigatus Emery 499 RiistomunmexiOMGtssy Sail. eeeet sesame cee 502 Pristomurenne xn ts Cid us Ist © iy aeseee amen oee eee 503 Pristomyrmex mandibularis Mann — 505 Pristomyrmex minusculus sp. M. —------ 507 Pristomyrmex obesus, Mann 2 = 509 Pristomyrmex simplex CO theae eee eer 512 DVS) 7D iRROy NAGHVIS! (Gako\ONS) 2 ee 514 Pristomyrmex profundus sp. n. -.......------- 515 ihenvmbripenniss Group yess = seen 516 Pristomyrmex fuscipennis (F. Smith) 517 PristOmmuymimen fViClett iI Iiy eee eens ee 518 Pristomyrmex pollux Domisthorjem=s =e 521 Pristomyrmex reticulatus Donisthorpe 524 Pristomyrmex umbripennis (F. Smith) 525 INiomnem INfuvdlummn 528 Pristomyrmex parvispina Emery 528 Ackaitowvled came mts eee seen eee 539 IRE LETETN CE Sie ee ts inna clean ESE ON eh ONS Rese 539 ABSTRACT. The ant genus Pristomyrmex is revised as a whole for the first time. The genus is redefined, and seven species groups are erected and discussed. Illustrations are present for all 52 species. A key to the worker caste is provided. Twenty-one new species are described: 20 from the Oriental region and one from Mauritius. Thirteen names are newly synony- mized, and two former infraspecific taxa are elevated to species rank. INTRODUCTION Pristom yrmex, an ant genus of moderate size, contains 52 living species, but fossils 157(6): 383-542, April, 2003 383 384 have not been discovered. Pristomyrmex occurs primarily in the Oriental region, but six endemic species are present in the eastern rainforest of Australia and five en- demic species in Africa. In addition, in Mauritius there are three native species, one of which also occurs on Reunion I[s- land. Lastly, one species, Pristomyrmex punctatus, has invaded temperate China, Korea, and Japan. This species has also been detected at two entry ports in the United States and thus shows potential for spread via human commercial actions. Pristomyrmex belongs to the subfamily Myrmicinae. It possesses a raised trans- verse ridge or a few toothlike prominences on the dorsal labrum in all female castes, including workers, ergatoid queens, and queens. This character is also shared by the myrmicine genera Acanthomyrmex, Myrmecina, and Perissomyrmex. As a re- sult, these four living genera are grouped together in the tribe Myrmecinini (Bolton, 1994, 1995, personal communication; Brown, 1971). Pristomyrmex is unique in the tribe because it is the only genus pos- sessing 11 antennal segments in all three female castes and 12 segments in the male. Most Pristomyrmex species dwell in the rainforest, foraging as predators or scav- engers. An Asian species, P. punctatus, however, occurs in open and disturbed habitats (e.g., bare hills, agricultural areas, and beaches). These ants prefer to nest in soil, litter, or rotten wood; in rotten parts of living trees; in dead standing trees; or around plant roots. Pristomyrmex is of great interest be- cause it exhibits several unusual biological and evolutionary phenomena. The absence of morphologically normal queens and re- production primarily by unmated workers in P. punctatus (=P. pungens) is a highly unusual life history in the Formicidae. It has attracted much attention from those who hope to obtain insight into the nature of reproductive conflict within colonies since, in this species, reproductive division of labor occurs among morphologically identical workers (Itow et al., 1984; Mu- Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 zutani, 1980, 1982; Peeters, 1993; Tsuji, 1988a,b,c, 1990a,b, 1994, 1995; Tsuji and It6, 1986). Ergatoid queens, a_ special wingless female caste morphologically in- termediate between the queen and the worker, are present in at least four species: P. punctatus, P. africanus, P. wheeleri, and P. mandibularis; two of them (P. africanus and P. wheeleri) possess both queen and ergatoid queen castes. Character displace- ment, showing that two species possess a greater difference in sympatric than allo- patric populations, has also been reported in this genus by Taylor (1965). In addition, simulating death, slowness of movement, and nocturnal foragers are also recorded in Pristomyrmex (Donisthorpe, 1946; Tay- lor, 1965; Weber, 1941). Colony size varies greatly among species, ranging from about a dozen to several thousand workers (Don- isthorpe, 1946; Itow et al., 1984; Mann, 1919; Taylor, 1965, 1968). Although Pristomyrmex is biologically promising, the taxonomic foundation of — the genus is poor. Much of the literature on Pristomyrmex is more than 50 years old and consists of isolated descriptions of spe- cies or infraspecific forms. Only a handful of papers present more comprehensive studies of the Australian and African su- bfaunas, respectively (Bolton, 1981; Taylor, 1965, 1968). The tropical Asian region, however, containing the bulk of the de- scribed taxa, has been in taxonomic chaos, for many years obscuring a better under- | standing of the evolution and radiation of | this interesting group. This survey takes the whole Pristomyr- mex into consideration. I believe that only after that the entire genus covering all zoo- geographical regions is comprehensively | investigated can a full set of characters to | define the genus be summarized, the spe- _ cies groups correctly erected, and the re- | lationships between species properly ana- lyzed and then the possible origin and the | evolution of the genus hypothesized. I present a detailed description of the | taxonomic characters for the worker caste © of each species. These characters not only REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang are useful for the species identity but also provide important information for a fur- ther study on the phylogeny within the ge- nus. I also include illustrations and de- scriptions of males for many species as possible. This was done for three reasons. First, two species (P. pollux and P. reticu- latus) were described, each from a single male specimen, many years ago. Without examining other available males, I would not be able to assign these two species to their appropriate species group, and the discovery of other new species would then be impeded. Second, the males of most ant genera are very poorly characterized and thus cannot be curated properly in museum collections. Finally, I feel that these males contain some clues for the study of the phylogenetic relationships of the genus. COLLECTIONS AMNH American Museum of Natural History, New York, N.Y., U.S.A. Australian National Insect Col- lection, Canberra City, Australia Bishop Museum, The State Mu- seum of Natural and Cultural History, Honolulu, Hawaii, U.S.A. Natural History Museum, Lon- don, U.K. California Academy of Sciences, San Francisco, California, U.S.A. Institute of Zoology, Academy of Sinica, Beijing, China Institute of Zoology, Ukrainian National Academy of Sciences, Kiev, Ukraine Natural History Museum of Los Angeles County, Los Angeles, California, U.S.A. Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy Museum of Comparative Zool- ogy, Harvard University, Cam- bridge, Mass., U.S.A. Muséum d Histoire Naturelle, Geneva, Switzerland ANIC BMHH BMNH CASC IZAS IZUA LACM MCSN MCZC MHNG 385 Museum of Nature and Human Activities, Sanda, Hyogo, Japan Muséum National d’ Histoire Naturelle, Paris, France Museum fiir Naturkunde der Humboldt- Universitit zu Berlin, Berlin, Germany National Arthropod Collection, Mount Albert Research Center, Auckland, New Zealand Naturhistorisches Museum, Ba- sel, Switzerland Naturhistorisches Museum, Vi- enna, Austria Naturhistoriska Stockholm, Sweden Oxford University Museum, Ox- ford, U.K. South African Museum, Cape Town, South Africa National Museum of Natural History, Washington, D.C., U.S.A. MEASUREMENTS AND INDICES Head Width (HW). Maximum width of head, in full-face view, excluding the eyes (Fig. 1). Head Width Including the Eyes (HWE). Maximum width of head across the eyes, in full-face view. This measurement is used only in the male. Head Length (HL). Length of the head in full-face view, excluding the mandibles (Fig. 1), measured from the midpoint of a straight line across the occipital margin to either the apex of the median tooth (if it is present) of the anterior clypeal margin or the midpoint of a line connecting the apexes of the two lateral teeth (if the me- dian tooth is absent) of the anterior clypeal margin or the midpoint of the anterior clypeal margin (if the anterior margin lacks any teeth). Cephalic Index (CI). HW/HL xX 100. Scape Length (SL). Length of the an- tennal scape, including the lamella encir- cling the base of the scape but excluding the basal condyle (Fig. 1). Scape Index (SI). SL/HW X 100. MNHA MNHN MNHU NACA NHMB NHMV NHPS Piksmuseet, OXUM SAMC USNM 386 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 S) Figures 1-3. Measurements illustrated for this study (worker of Pristomyrmex longispinus sp. n.). 1: Head, full-face view; 2: Entire body, lateral view; 3: Dorsal view of alitrunk, petiole, and postpetiole. Pronotal Width (PW). Maximum width of the pronotum in dorsal view (Fig. 3). Alitrunk Length (AL). Diagonal length of the alitrunk in lateral view, from the an- teriormost point of the pronotum to the apex of the metapleural lobe (Fig. 2). Eye Length (EL). Maximum length of the eye. Total Length (TL). Til + TL2 + TL3 (see Fig. 2). (Note: The measurements of TL do not deal with those individuals whose gasters are abnormally contracted or prolonged or whose petioles are raised too high or too low.) TL1: A line mea- sured from the apex of the closed man- dibles to the midpoint of a straight line across the occipital margin, in full-face view. TL2: A straight line from the anter- iormost point of the pronotum to the point at which the posterior margin of postpetiole meets the uppermost point of an articulation. TL3: A line from the an- terior-uppermost point of the articulation to the apex of gaster. Pronotal Spine Length (PSL1). A straight distance from the base to the apex of pronotal spine (see Fig. 2). Propodeal Spine Length (PSL2). A straight distance from the base to the apex — of propodeal spine (see Fig. 2). Postpetiole Index (PPI). PPW/PPL xX } 100 (PPW: maximum width of the post- — petiole in dorsal vies; PPL: length of the | postpetiole in dorsal view). All measurements are taken in milli-— meters. Note: For the Australian species, my measurements differ slightly from Taylor's (1965, 1968). For the maximum measur- able width of head, I have excluded the eyes, which were included by Taylor. A BRIEF HISTORY OF THE GENUS PRISTOMYRMEX The genus Pristomyrmex, when erected | by Mayr (1866), contained one species, P. | pungens Mayr (=a junior synonym oe Je REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang punctatus (F. Smith)), a member of the present punctatus group. At that time, Mayr did not realize that five species de- scribed by F. Smith (1858, 1860, 1861, 1863, 1865), that is, Myrmica fuscipennis, Myrmica punctata, Myrmica_ trachylissa, Myrmica umbripennis, and Solenopsis lae- vis, also belonged in the new genus. Mayr (1866) provided a description of the genus, which actually was simply derived from some characters of the species P. puncta- tus (F. Smith). A second member of the punctatus group was introduced by Sants- chi (1916) when he transferred Tetramo- rium (Xiphomyrmex) fossulatum Forel to Pristomyrmex. The content of the genus Pristomyrmex expanded for the fist time when Mayr (1886) transferred Myrmica trachylissa F. Smith to Pristomyrmex. Pristomyrmex tra- chylissus is now a member of the quadri- dens group. After that, many species of the quadridens group were discovered (Bol- ton, 1981; Donisthorpe, 1949c; Emery, 1887, 1895, 1897, 1900; Forel, 1914; Ka- ravaiev, 1931, 1933; Taylor, 1965, 1968), and several more names were added to the group when Odontomyrmex André and Hylidris Weber were designated by Forel (1915) and Brown (1953) as a subgenus and a synonym of Pristomyrmex, respec- tively. But, Mann (1919) found no evi- dence supporting the subgenus Odonto- myrmex. The members of the third species group (i.e., umbripennis group) of Pristomyrmex were recognized first by Emery. He de- scribed a new species (Pristomyrmex pic- teti) in 1893 and transferred a species (Myrmica fuscipennis F. Smith) to Pristo- myrmex in 1901. Donisthorpe expanded the umbripennis group: He _ transferred Myrmica umbripennis F. Smith and Sole- nopsis laevis F. Smith to Pristomyrmex and described Pristomyrmex pollux and Pris- tomyrmex reticulatus (1932, 1946, 1949a). When Emery (1897) described Pristo- myrmex coggii, Pristomyrmex levigatus, and Bristom yrmex lucidus, representatives of the present levigatus group were added 387 to the genus for the first time. Pristomyr- mex cribrarius, the sole member of the cribrarius group, was described by Arnold (1926). Lastly, Brown (1971) synonymized the genus Dodous Donisthorpe, adding species belonging to the present trispino- sus group to the genus. Thus, the genus Pristomyrmex became clearly delimited and assumed its modern form. GENUS PRISTOMYRMEX MAYR Pristomyrmex Mayr, 1866: 903. Type species: Pristo- myrmex pungens Mayr, op. cit.: 904 [=Myrmica punctata F. Smith, 1860: 108; = Pristomyrmex punctatus (F. Smith) ]; by monotypy. Odontomyrmex André, 1905: 207. Type species: Odontomyrmex quadridentatus André, op. cit.: 208; by monotypy. [As a subgenus, thus synonym, of Pristomyrmex by Forel, 1915: 53.] Hylidris Weber, 1941: 190. Type species: Hylidris myersi Weber, loc. cit. (=Pristomyrmex africanus Karavaiev); by original designation. [Synonymy by Brown, 1953: 9.] Dodous Donisthorpe, 1946: 145. Type species: Do- dous trispinosus Donisthorpe, loc. cit.; by original designation. [Synonymy by Brown, 1971: 3.] Diagnosis of worker, queen, and erga- toid queen. Combination of the following asterisked four characters (i.e., characters 2, 7, 11, and 29 in the worker caste) sep- arating Pristomyrmex from other myrmi- cine genera. Definition: Worker. Possessing the fol- lowing combination of characters: 1. Small (TL 1.74, HL 0.46, HW 0.46) to large-sized (TL 7.06, HL 1.68, HW 1.74) monomorphic myrmicine ants. *2. Mandible somewhat subtriangular; masticatory margin of mandible with he ee to five teeth, Shieh have one or the other of the following six basic arrangements: (1) the strongest apical + the second strongest preapical + the smallest third + the acute basal tooth, dia- stema lacking, as in levigatus group and in profundus group, or (2) the strongest apical + the second strongest preapical + two smaller teeth of similar size, diastema indis- tinct or lacking, as in umbripennis group, or (3) the strongest apical + the second 388 strongest preapical + a shorter (first) diastema (sometimes the first diastema is not distinct) + a small denticle + a longer (second) diaste- ma + a small basal denticle, as in both P. bispinosus and P. trispino- Sus, OF the apical + the preapical + a lon- ger diastema + a small denticle + a shorter diastema (sometimes the second diastema is indistinct) + a small basal denticle, as in PR. browni, or the strongest apical + the second strongest preapical + a distinct di- astema + a basal tooth (which is sometimes formed by the fusion of the two small teeth) or two (or three) small teeth of similar size, as in punctatus group, cribrarius group, and most members of the quardridens group, or the strongest apical + the second strongest preapical + an intercalary tooth + a very short diastema (or this diastema indistinct) + two small teeth of similar size, as shown in P. trachylissus. 3. Basal margin of mandible with a broad-based triangular or an acute and prominent tooth, or only curved, not form- ing tooth, or almost straight. 4. Median part of clypeus shieldlike, projecting posteriorly between the bases of the antennae; lateral parts of clypeus in front of antennal insertions usually re- duced to ridges but rarely (in the two Ori- ental species P. divisus and P. pulcher) de- veloped so that the antennal fossae do not reach the lateral anterior margins of clyp- eus. 5. Anterior clypeal margin usually with a median tooth and one to three pairs of lateral denticles (or crenulate shapes) but sometimes the median tooth rudimentary (as in some species of the levigatus group) and sometimes anterior clypeal margin lacking any distinct denticles (as in P. pro- fundus, P. divisus, and P. pulcher). 6. Ventral surface of clypeus with a me- (5) (6) Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 dian tooth or two lateral teeth, or with a transverse ridge, or without any ridge or tooth. *7. Dorsal labrum with a raised trans- verse ridge or a few toothlike prominenc- es, present on the anterior portion of la- brum in most species. 8. Palp formula l,2, 1372/20 253)4)3 0% 53. 9. Frontal lobes absent in punctatus and trispinosus groups or weak, as in lev- igatus, profundus, and quadridens groups, or somewhat expanded, as in umbripennis group; as a result, the articulations of the antennae are mostly or entirely exposed in full-face view. 10. Frontal carinae usually developed, extending to the level of the posterior mar- gins of eyes, but sometimes frontal carinae absent or very short, as in the trispinosus group, in P. trogor, and in P. longispinus. *11. Antennae with 11 segments; apical three segments forming a distinct club. 12. Base of each antennal scape encir- cled by a narrow lamella, except in P. pro- fundus; this lamella usually with a broad and deep notch on the center of dorsal surface in the umbripennis group but en- tire in the other species groups. 13. Antennal scrobes usually absent or weakly developed, but in P. profundus, the scrobes are deep and well developed. 14. Eyes present in all known species, situated approximately at the midlength of the sides of the head; usually moderate- sized, but small in the several species (P. boltoni, P. coggii, P. longus, P. eduardi, P. picteti, and P. pollux). 15. Alitrunk usually lacking dorsal su- tures, but in the three species of the trispi- nosus group, a promesonotal suture or im- pression present. 16. Pronotum unarmed, or armed with a pair of tubercles, teeth, or spines of vary- ing sizes. 17. Mesonotum usually unarmed, but with a pair of thick, blunt, and digitlike short prominences in P. trispinosus, and sometimes weakly tuberculated in P. bispi- nosus and P. browni. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 18. Propodeum armed with a pair of teeth or spines, except in P. inermis. 19. Metapleural lobes usually subtrian- gular, or each with a blunt-rounded to semicircular apex, but indistinct in P. pro- fundus. 20. Fore tibial spurs pectinate. Middle and hind tibiae sometimes without any spur, sometimes with either simple or hair- like spurs. 21. Propodeal spiracles circular and high-positioned on the lateral surfaces of the propodeum. 22. Metapleural gland bullae large, sep- arated from the propodeal spiracles, and positioned above the posterior lower cor- ners of propodeum. 23. Petiole in profile nodiform or wedge-shaped, pedunculate, usually with a long anterior peduncle. 24. Subpetiole sometimes without a ventral process, sometimes bearing a nar- row semitranslucent lamella. In P. acero- sus, a pinlike process is present. 25. Postpetiole in profile nodiform, usually rounded dorsally. 26. Petiole spiracle, postpetiole spira- cle, and first gastral spiracle visible. 27. Dorsal surfaces of head and alitrunk smooth, or possessing either scattered fo- veolate punctures, or foveolate-reticulate sculpture, or developed rugoreticulum, or regular striate sculpture. Gaster unsculp- tured. 28. Dorsal surfaces of head and alitrunk usually with numerous hairs, but only a few hairs present on the dorsal alitrunk in P. fossulatus, P. orbiceps, and P. trogor. Petiole and postpetiole each usually with one to three pairs of hairs, but sometimes more pairs of hairs present; sometimes petiole and postpetiole lacking hairs. First gastral tergite usually without hairs or with a few sparse hairs, but sometimes first gas- tral tergite covered with numerous, evenly distributed, erect or suberect hairs. *29. Anterior clypeal margin lacking a median seta at the midpoint of the margin, instead usually having two to three pairs of 389 long, forward-projecting hairs flanking the midpoint of margin. 30. Sting slender and long. Female. Usually alate, but in some spe- cies (P. punctatus, P. mandibularis), only ergatoid queens have been found. In some species (P. wheeleri, P. africanus), both alate and ergatoid queens exist. Alate Queen. Characters similar to those of worker in the structure and shape of mandible, palp formula, clypeus, frontal lobes, frontal carinae, antennae, meta- pleural lobes, tibial spurs, petiole node, postpetiole, and sting as well as in the sculpture of body. But larger, with slightly or much larger eyes, than in the conspe- cific worker; three ocelli present. The ali- trunk with wings and flight sclerites; well- marked dorsal sutures present. Pronotal spines usually absent, but in some species, the pronotum is armed with a pair of teeth that are much shorter than in conspecific worker; propodeal teeth or spines usually shorter than those of conspecific worker. Wing venation as shown in Figures 4—5. On the forewings, the marginal cell (see Holldobler and Wilson, 1990: 9) is always open; R + Sc thick (for the explanation of symbols used, see Brown and Nutting, 1950); A short, far from the anal angle; A, Cu-A, Mf2+3 usually reduced to vestigial lines distally; cross-vein m-cu and r-m ab- sent; cross-vein cu-a usually present but sometimes broken in larger species (such as P. picteti, P. umbripennis ) and some- times rudimentary or very weak in some samples of a few smaller species (e.g., P. orbiceps, P. lucidus); 1r absent; anal lobe usually indistinct in smaller species but present in larger species. Hind wings with- out anal lobe. (Note: The venation of the both fore and hind wings of alates, in Pris- tomyrmex, is rather stable, with only slight variations within the different species. For example, on the forewings, Mf2+3, some- times becomes an almost entirely vestigial line, but sometimes it is distinct and rather long; Rsf4 + Rsf5 is rather thick and long in some larger species but thin and short in some smaller species). 390 5 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 7 Figures 4-7. General forewing and hindwing venation of alate queens and males of Pristomyrmex. 4: Forewing of alate queens; 5: Hindwing of alate queens; 6: Forewing of males; 7: Hindwing of males. [The forewing and hindwing venation of the alate queens of the following 13 Pristomyrmex species was examined: P. brevispi- nosus, P. collinus, P. orbiceps, P. quadridens, P. quindentatus, P. sulcatus, P. levigatus, P. lucidus, P. obesus, P. fuscipennis, P. picteti, P. pollux, and P. umbripennis. The males of 16 Pristomyrmex species were examined: P. brevispinosus, P. flatus, P. trogor, P. longispinus, P. orbiceps, P. quadridens, P. quadridentatus, P. sulcatus, P. browni, P. trispinosus, P. obesus, P. levigatus, P. picteti, P. pollux, P. umbripennis, and P. punctatus.] Ergatoid Queen. General characters, in- cluding the pronotal prominences and size of body, similar to those of the conspecific worker. Ocellus present (one ocellus in P. mandibularis but three ocelli in P. punc- tatus, P. wheeleri, and P. africanus); apter- ous, but mesonotum more convex than in conspecific worker; pro-mesonotal suture present in P. mandibularis but represented by an impression in P. punctatus, P. whee- leri, and P. africanus. Male. Possessing the following combi- nation of characters (summarized accord- ing to 54 specimens falling into at least 17 species): 1. Small to moderate size (TL 2.40— 6.04, HL 0.48—0.94, HW 0.51—0.98, HWE 0.62-1.10), usually smaller than the con- specific queen. 2. Head, in full-face view, across and in- cluding the eyes, usually broader than long (Figs. 261-269). 3. Mandibles vestigial, very small, rounded or toothlike, far from meeting, as indicated by an arrow in Figure 262. 4. Anterior margin of labia broadly concave at center; Mors of labrum with- out any transverse ridge or toothlike prom- inences (see Figs. 262, 264, 269). 5. Eyes very large, well developed, and convex, situated at the sides of head. 6. Antennae filiform, 12 segments, lack- ing a lamella encircling the base. Scapes short, usually distinct shorter than the maximum length of eye; of the other 11 funicular segments, the first segment shortest, the apical segment longest, the remaining nine segments much longer than their broad. Three ocelli conspicuous and well developed, situated on the vertex of the head. 8. Antennal sockets set back from the posterior margin of the clypeus. 9. Antennal scrobes absent. 10. Frontal carinae absent or very short and weak. 11. Frontal lobes absent so that the ar- ticulations of the antennae are completely exposed in full-face view. 12. Palp formula as in the conspecific worker in seven species examined (i.e., P. punctatus, P. quadridens, P. curvulus, P. brevispinosus, P. sulcatus, P. picteti, and P. pollux). 13. Clypeus convex in the middle, not projecting posteriorly upward between the bases of antennae; its shape transverse, Or REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang somewhat semicircular; its anterior margin entire, without any denticles, usually rath- er straight but sometimes arched. 14. Cheeks very short. 15. Alitrunk robust, with wings, well- developed flight sclerites, and well-marked sutures. 16. Pronotum narrow in middle, over- hung by mesoscutum in lateral view, lack- ing any armaments. 17. Mesonotum well developed, con- sisting of a large mesoscutum, a rather large mesoscutellum, and two small axillae. Notauli usually distinct, forming a Y shape, but sometimes they show a V shape, and sometimes they are absent or very weak. Parapsidal furrows usually absent, but sometimes they are superficially im- pressed. 18. Metanotum transverse, overhung by mesoscutellum. 19. Propodeum showing a sloping dor- sal surface; propodeal armaments absent or present; if present, they are usually shorter than in the conspecific worker. 20. Metapleural lobes present, sub- triangular, or toothlike, or blunt-rounded to semicircular. 21. Venation (Figs. 6-7) as in alate queen. 22.. Legs slender; fore tibial spurs pec- tinate; middle and hind tibiae usually lack- ing any spurs but sometimes simple spurs are present. 23. Petiole with a long or a rather long anterior peduncle. In dorsal view, sides of petiole subparallel. Petiole node low, lower than in the conspecific worker and queen; subpetiole lacking any lamella or toothlike projection. 24. Postpetiole node rather low, lower than in the conspecific worker and queen. In profile, subpostpetiole usually lacking any projections, but sometimes bearing a small tooth. 25. Positions of spiracles on propo- deum, petiole, postpetiole, and first gastral segment similar to those in the conspecific worker and queen. narrow, 391 26. Usually much less sculptured than conspecific worker and queen. 27. Numerous hairs present on the en- tire dorsal surfaces of body. (Note: The genitalia of males is not dis- sected. ) The male of Pristomyrmex can be dis- tinguished within the tribe Myrmecinini by the following characters: Pristomyrmex Antennae: 12 segments Mandibles: Very small, toothlike, not meeting Petiole: With a long anterior peduncle Forewing: Without m-cu cross-vein; marginal cell open Acanthomyrmex Antennae: 13 segments Mandibles: Subtriangular, with six to eight teeth, meeting when they are closed Petiole: Similar to that of Pristomyrmex Forewing: PWith m-cu cross-vein; mar- ginal cell closed Myrmecina Antennae: 13 segments Mandibles: Similar to those of Pristo- myrmex Petiole: Without an anterior peduncle Forewing: Without m-cu cross-vein: marginal cell closed Perissomyrmex Antennae: ?10 segments Mandibles: Unknown Petiole: Unknown Forewing: Unknown Larva. According to Wheeler and Wheeler’s (1954, 1960, 1973, 1976) stud- ies, the larva of Pristomyrmex has the fol- lowing combination of characters: 1. Stout and rather short. 2. Head extremely long and narrow. 3. Thorax more slender than abdomen and forming a neck, which is curved ven- trally. Diameter greatest near middle of abdomen, decreasing gradually toward head; posterior end rounded. 4. Body without tubercles. 392 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 5. Mandibles subtriangular, without medial blade; apical tooth curved medially and usually acute; subapical medial tooth small. 6. Body hairs numerous, with five or six types, including anchor-tipped hairs. Head hair few, short to moderately long. 7. Gula spinulose. 8. Anterior surface of labium densely spinulose. 9. Palps lateral. Pupa. Not enclosed in cocoons (Wheel- er and Wheeler, 1976). LIST OF PRISTROMYRMEX NAMES WITH SYNONYMIES (Currently valid names are in boldface) acerosus: sp. n. africanus: Pristomyrmex africanus Karavaiev =beni =mbomu =myersi =primus aruensis: Pristomyrmex quadridens var. aruensis Karavaiev =quadridens beni: Hylidris myersi subsp. beni Weber =africanus bicolor: stat. n.: Pristomyrmex trachy- lissa var. bicolor Emery =taurus syn. n. bispinosus: Dodous bispinosus Donis- thorpe boltoni: sp. n. brevispinosus: Pristomyrmex brevi- spinosus Emery =yaeyamensis syn. n. browni: sp. n. castaneicolor: Pristomyrmex castaneico- lor Donisthorpe =umbripennis castor: Pristomyrmex castor Donis- thorpe =umbripennis coggit: Pristomyrmex coggti Emery collinus: sp. n. costatus: sp. n. cribrarius: Pristomyrmex cribrarius Arnold curvulus: sp. n. divisus: sp. n. eduardi: Pristomyrmex eduardi Forel erythropygus: Pristomyrmex ery- thropygus Taylor flatus: sp. n. formosae: Pristomyrmex brevispinosus r. sulcatus var. formosae Forel, 1912: 54. unavailable name fossulatus: Tetramorium (Xiphomyr- mex) fossulatum Forel foveolatus: Pristomyrmex foveolatus Taylor fuscipennis: Myrmica fuscipennis F. Smith hirsutus: sp. n. inermis: sp. n. japonicus: Pristomyrmex japonicus Forel =punctatus laevigatus: Hylidris laevigatus Weber =orbiceps laevis: Solenopsis laevis F. Smith =umbripennis largus: sp. n. levigatus: Pristomyrmex levigatus Emery =mendanai syn. n. longispinus: sp. n. longus: sp. n. lucidus: Pristomyrmex lucidus Emery mandibularis: Pristomyrmex mandi- bularis Mann mbomu: Hylidris myersi subsp. mbomu Weber =africanus melanoticus: Pristomyrmex subsp. melanoticus Mann =obesus mendanai: Pristomyrmex mendanai Mann =levigatus minusculus: sp. n. modestus: sp. n. myersi: Hylidris myersi Weber =africanus nitidissimus: Pristomyrmex nitidissi- mus Donisthorpe obesus: Pristomyrmex obesus Mann =melanoticus syn. n. obesus REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 393 =pegasus syn. n. occultus: sp. n. orbiceps: Xiphomyrmex orbiceps Sants- chi =laevigatus orbiculatus: Pristomyrmex orbiculatus Donisthorpe =quadridens parumpunctatus: Pristomyrmex parum- punctatus Emery =umbripennis parvispina: Pristomyrmex parvispina Emery, 1900: 678. Nomen nu- dum pegasus: Pristomyrmex pegasus Mann =obesus picteti: Pristomyrmex picteti Emery =tingiana syn. n. pollux: Pristomyrmex pollux Donis- thorpe primus: Hylidris myersi subsp. primus Weber =africanus profundus: sp. n. pulcher: sp. n. punctatus: Myrmica punctata F. Smith =japonicus =pungens syn. n. pungens: Pristomyrmex pungens Mayr =punctatus quadridens: Pristomyrmex quadri- dens Emery =aruensis syn. n. =orbiculatus syn. n. quadridentatus: Odontomyrmex quad- ridentatus André =queenslandensis queenslandensis: Pristomyrmex (Odon- tomyrmex) quadridentatus var. queenslandensis Forel =quadridentatus quindentatus: sp. n. reticulatus: Pristomyrmex reticula- tus Donisthorpe rigidus: sp. n. simplex: sp. n. sulcatus: stat. n.: Pristomyrmex brevi- spinosus subsp. sulcatus Emery taurus: Pristomyrmex taurus Stitz =bicolor thoracicus: Pristomyrmex thoracicus Taylor tingiana: Pristomyrmex picteti var. tin- giana Stitz =picteti trachylissus: Myrmica trachylissa_ F. Smith trispinosus: Dodous trispinosus Donis- thorpe trogor: Pristomyrmex trogor Bolton umbripennis: Myrmica umbripennis F. Smith =castaneicolor syn. n. =castor syn. n. =laevis syn. n. =parumpunctatus syn. n. wheeleri: Pristomyrmex wheeleri Taylor wilsoni: Pristomyrmex wilsoni Taylor yaeyamensis: Pristomyrmex yaeyamen- sis Yamane and Terayama =brevispinosus KEY TO THE WORLD SPECIES OF PRISTOMYRMEX (WORKERS) Note: P. fuscipennis and P. reticulatus, whose worker castes are presently unknown, are not includ- ed in the key. Tle Dorsum of alitrunk in profile not arched, with mesonotum much higher than propodeal dorsum, that is, a vertical cliff present between mesonotum and propodeal dorsum (Fig. 8). An- tennal scrobes well developed and deep. Basal margin of mandible with a strong tooth adjacent to the basal tooth of masticatory margin so that five teeth are set close together (Fig. 27). Base of antennal scape without a circling lamella (Fig. 80) (profundus group; Asia: Sabah) —----- Siena ees profundus (p. 515) Dorsum of alitrunk in profile, excluding armaments, more or less arched- shaped, never showing a _ vertical cliff between mesonotum and pro- podeal dorsum (Figs. 9-25). Anten- nal scrobes absent or shallow. Tooth on basal margin of mandible either absent or present; if present, it is on about midway, not adjacent to the basal tooth of masticatory margin (Figs. 28, 29, 35). Base of antennal scape with a circling lamella (Figs. SHS 2)) ie aE ae ae ce es 2; 394 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 TABLE 1. A LIST OF PRISTOMYRMEX SPECIES AND THEIR BIOGEOGRAPHIC DISTRIBUTION. Species Group Species Name ORI PAL AUS AFR IOI punctatus group P. divisus sp. n. oF P. fossulatus (Forel) + P. pulcher sp. n. + *P. punctatus (F. Smith) + + rigidus sp. n. * cribrarius group cribrarius Arnold + quadridens group africanus Karavaiev + bicolor Emery +f brevispinosus Emery ate collinus sp. n. + costatus sp. 0. aie curvulus sp. n. is eduardi Forel +f erythropygus Taylor 4p flatus sp. n. +f foveolatus Taylor 2s hirsutus sp. n. a longispinus sp. n. ati modestus sp. n. a5 + + nitidissimus Donisthorpe occultus sp. n. orbiceps (Santschi) oo quadridens Emery + quadridentatus (André) + quindentatus sp. n. sulcatus Emery thoracicus Taylor + trachylissus (F. Smith) + trogor Bolton + wheeleri Taylor a wilsoni Taylor 4s bispinosus (Donisthorpe) browni sp. n. trispinosus (Donisthorpe) acerosus sp. n. boltoni sp. n. coggit Emery inermis sp. Nn. largus sp. n. levigatus Emery longus sp. n. lucidus Emery > mandibularis Mann *P. minusculus sp. n. P. obesus Mann P. simplex sp. n. profundus group P. profundus sp. n. umbripennis group P. fuscipennis (F. Smith) P. picteti Emery P. pollux Donisthorpe P. reticulatus Donisthorpe P. umbripennis (F. Smith) neh as) ash awlas) mshi as) as) eh melas) ae) ae) is) as)! Se) fas) das) As) taehlac) tas) as) las) be) N AS ae) AS} aS) gs) as) Bs) as) else) As) As) ashe trispinosus group +++ levigatus group S+ttt+t+++++¢¢¢¢¢4¢4¢4¢4+ + Total number of species 1 vl 5 3 Total number of the endemic species 36 6 5 3 Notes: ORI, PAL, AUS, AFR, and IOI are abbreviated, respectively, from the Oriental region, the Palaearctic region, Australia, Africa, and Indian Ocean Islands. AFR refers to the African continent only. Pristomyrmex has not been recorded from Madagascar. “*” symbol indicates that P. punctatus and P. minusculus occur in the two regions, respectively. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 395 25 Figures 8-25. Alitrunks of Pristomyrmex workers, lateral view. 8: P. profundus sp. n.; 9: P. levigatus Emery; 10: P. inermis sp. n.; 11: P. minusculus sp. n.; 12: P. picteti Emery; 13: P. pollux Donisthorpe; 14: P. brevispinosus Emery (non-type); 15: P. brevispinosus Emery (syntype); 16: P. foveolatus Taylor; 17: P. sulcatus Emery (syntype); 18: P. sulcatus Emery (non-type); 19: P. quadridentatus (André); 20: P. wheeleri Taylor; 21: P. longispinus sp. n.; 22: P. trispinosus (Donisthorpe); 23: P. browni sp. n.; 24: P. bicolor Emery; 25: P. wilsoni Taylor. Figure 26. Propodeal spines of the worker of Pristomyrmex browni sp. n., dorsal view. 396 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 —~ — eee es ——_—>> See eee Ae aes 34 35 36 —— a | ——_ 37 : ; 2 : SP 38 39 Figures 27-39. Mandibles of Pristomyrmex workers. 27: P. profundus sp. n.; 28: P. levigatus Emery; 29: P. mandibularis Mann; 30: P. picteti Emery; 31: P. quadridens Emery; 32: P. quindentatus sp. n.; 33: P. quadridentatus (André); 34: P. trachylissus (F. Smith); 35: P. rigidus sp. n.; 36: P. punctatus (F. Smith); 37: P. browni sp. n.; 38: P. trispinosus (Donisthorpe); 39: P. bispinosus (Donisthorpe). QD. Masticatory margin of mandible with New Guinea; Indonesia; Pohnpei four teeth; the third tooth, counting Is.; Palau Is.; Yap I.; Tonga Is.; Wal- from the apex, smallest, distinctly lis Is.; found rarely in N. Queens- smaller than the basal one; diastema land, Australia) _. minusculus (p. 507) absent between the preapical and Pronotummunanned (Hig). 5 the third tooth (Figs. 28-29) (levi- 5, Postpetiole in profile with an arched an- gatus group; Asia, Australia) 3 terior face and a steeply sloping pos- Masticatory margin of mandible with terior face and the apex of postpe- three to five teeth; if four teeth pre- sent, then the third tooth, counting from the apex, similar in size to the basal one; diastema either present or indistinct between the preapical and the third tooth (Figs. 30-39) _ 14 oy Propodeum unarmed (Fig. 10). Petiole node in profile wedge-shaped (Fig. 45) (Asia: Papua New Guinea) -------- a ee re Lr et oe inermis (p. 496) Propodeum armed with a pair of teeth or spines (Figs. 9, 11). Petiole node in profile nodiform, not wedge- shaped (Figs. 40-44) 4 4. Pronotum armed with a pair of teeth (Fig. 11) (Asia and Pacific Is.: Papua tiole pointing posterior-upwardly (Fig. 43); in dorsal view, postpetiole usually longer than broad, very rare- ly about as long as broad. Petiole node with a single evenly blunt- rounded apex (Fig. 43). Head broader; HW mostly >1.00 (Asia: Papua New Guinea) -.... lucidus (p. 503) Postpetiole in profile with a somewhat evenly convex dorsum, lacking an abruptly steep posterior face (Figs. 40—42, 44) and in dorsal view broad- er than long. Petiole node in profile with a distinct anterodorsal angle (Figs. 40-42, 44). Head narrower; Dea IN Varco lel 010 Veena ae nk SO ee REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 397 Figures 40-58. Petiole nodes and postpetioles of Pristomyrmex workers. 40A: Dorsal surface of the petiole node of P. obesus Mann, dorsal view; 41A: Dorsal surface of the petiole node of P. /Jongus sp. n., dorsal view; 40B, 41B, 42-58: Petiole nodes and postpetioles, lateral view: 40B: P. obesus Mann; 41B: P. longus sp. n.; 42: P. acerosus sp. n.; 43: P. lucidus Emery; 44: P. mandibularis Mann; 45: P. inermis sp. n.; 46: P. punctatus (F. Smith); 47: P. rigidus sp. n.; 48: P. cribrarius Arnold; 49: P. quadridens Emery; 50: P. africanus Karavaiev; 51: P. nitidissimus Donisthorpe; 52: P. collinus sp. n.; 53: P. flatus sp. n.; 54: P. curvulus sp. n.; 55: P. longispinus sp. n.; 56: P. hirsutus sp. n.; 57: P. sulcatus Emery; 58: P. modestus sp. n. 398 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 8. Eyes smaller, with two to three omma- tidia in the longest row (Asia: New (@uin'ea)) eee boltoni (p. 492) Eyes larger, usually with five to seven ommatidia in the longest row ___. 9 9. Larger species, with HW 0.90-0.96 and cy ZEN HL 0.90-0.90 (Pacific Is.: Pohnpei 59 Zany Pa LT he iia eRe eee largus (p. 497) Smaller species; HW < 0.80, HL < 0.80 Figures 59-60. Gasters of Pristomyrmex workers, lateral 10 Cea ene Sone hol ae )e “th al Sr rt view. 59: Gaster of P. simplex sp. n.; 60: Gaster of P. obesus ; PEVOle witht ay Ong Mann. pinlike process (Fig. 42) (Pacific Is.: New Hebrides) acerosus (p. 491) Ventral surface of petiole without a long 6. Dorsum of petiole node laterally com- pinlike process (Figs. 40, 41, 44) — 11 pressed and in dorsal view distinctly Le Each side of petiole with a longitudinal longer than broad (Fig. 41A) (Asia: carina that separates the tergite * New Fat oe ranges ‘ 502) from the sternite (Fig. 40). Basal orsum of petiole node not laterally Ries maya eee compe eee and in dorsal view Lara cecny on He ube about as broad as or broader than long, not longer than broad (Fig. A TAN) fap sateen aie eas 220 tS, aE Dorsum of alitrunk unsculptured, smooth and shining. Dorsum of head between frontal carinae usu- ally smooth and shining (Fig. 61), except for a few punctures border- cific Is.: Papua New Guinea, Solo- mon Is., Nama Is., New Britain Is.) 5 LOE fo, Se) Neel levigatus (p. 499) Sides of petiole unsculptured and smooth, lacking a longitudinal carina (Fig. 44). Basal margin of mandible with a prominent tooth (Fig. 29) (Raxerinv® 105.9 VEMD) een tet nate s “ ~“ imperromtale camin ae) ees 8 : : Dorsum’ of alitrumk with,some scattered’ et ee mandibularis (p. 505) foveolate punctures. Dorsum of 12(7). Entire first gastral tergite evenly clothed head between frontal carinae with with numerous erect or suberect scattered foveolate punctures or fo- hairs (Fig. 60) (Pacific Is.: Solomon veolate-reticulate sculpture (Figs. LSS) Pea ee ooh eee obesus (p. 509) (OGY i a ae ge en SAN ee 12 Only a few hairs present usually near the o> CS ay pas: SS L) S V2 \e I\N ) 65 66 67 Figures 61-67. Characters on the sculpture of the dorsal heads of Pristomyrmex workers, full-face view, excluding a portion of the mandibles. 61: Smooth head; 62-63: Scattered foveolate punctures; 64: Foveolate-reticulate sculpture; 65: Rugoreticulum; 66: Regular striations around the antennal fossae and on the genae; 67: Regular striations on the entire dorsum of the head. 13. 14(5 15. 16. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang base of the first gastral tergite (Fig. BIO) Pe eS ROE PS ae ae Eee Dorsum of head, except for the scrobes, with foveolate-reticulate sculpture; punctures often aligned so that it seems that several longitudinal ru- gae appear between frontal carinae. Eyes smaller, EL = 0.06—0.08, with three to four ommatidia in the lon- gest row (Asia: Papua New Guinea) 13 a ea ees owe Ne coggii (p. 493) Dorsum of head between frontal carinae with some scattered foveolate punc- tures; space between foveolae smooth. Eyes larger, EL = 0.09- 0.12, usually containing five (rarely four) ommatidia in the longest row (Asia: Papua New Guinea) ------------ suerte aceite ikabie ioe Ane Ae simplex (p. 512) Masticatory margin of mandible with four teeth, lacking a distinct diaste- ma (Fig. 30). Lamella, circling the base of antennal scape, with a broad and deep notch on the center of the dorsal surface (Fig. 82). Petiole node in profile longer than high (umbripennis group; ein) Poa Poranl a Masticatory margin of mandible with three to five teeth; usually with a distinct diastema (Figs. 31-33, 35- 39); if (very rarely) diastema indis- tinct, masticatory margin with five teeth (Fig. 34). Lamella, circling the base of antennal scape, entire, with- out a notch on the center of the dor- sal surface (Fig. 81). Petiole node in profile usually higher than long __. Eyes larger, usually consisting of 20 or more ommatidia, containing six to seven ommatidia in the longest row. Propodeum with a pair of Roothilike armaments that are shorter than the distance between their bases. About one-third of antennal scape usually laterally compressed near the base (Asia: Papua New Guinea, Indone- 15 7, S12) pea aa emus umbripennis (p. 525) Eyes smaller, generally consisting of less than 10 ommatidia, with three to four ommatidia in the longest row. Propodeum with a pair of spines that are longer than the distance be- tween their bases. Antennal scape not laterally compressed near the [SING ceo sine eda te en oo ee Propodeal spines longer and strongly up- curved at their apices (Fig. 13). Larger species with HL 1.42—1.54, HW 1.42-1.58 (Asia: W. Malaysia, N. Borneo) Propodeal spines shorter and not strong- 16 pollux (p. 521) 18. 19. ly upcurved at their apices (Fig. 12). Smaller species with HL 1.04-1.36, HW 1.02-1.40 (Asia: Papua New Guinea, Indonesia, Singapore, Ma- laya, Sabah, Brunei, Philippines) -—- 399 wb: 2ca Jui dane Ne ee te picteti (p. 518) 17(14). Pronotum unarmed. Eyes larger, usually containing seven or more (very rare- ly six) ommatidia in the longest row. Dorsal surfaces of head and alitrunk with well developed rugoreticulum or many foveolate punctures (punc- tatus group; Asia, Africa) Pronotum usually with a pair of teeth or spines; if (very rarely) pronotal teeth or spines absent, then either eyes smaller, with two to five ommatidia in the longest row, or dorsal surfaces of ARE and head between frontal carinae unsculptured and smooth __ Dorsum of alitrunk with well developed coarse reticulum. |Propodeum armed with a pair of long spines. Antennal scapes longer, usuall >0.78; one-sixth to one-fifth of the length of the scapes projecting be- yond the occipital margin. Palp for- TOGNDU Kee Dy Gis aubeee see ae ak a ned nl oe Dorsum of alitrunk with scattered fove- olate punctures. Propodeum armed with a pair of short spines. Antennal scapes shorter, with the length 0.54— 0.60, only close to the occipital mar- gin. Palp formula 4,3 (South Africa) 18 bo bo 19 Sa arse to Oe See fossulatus (p. 406) Lateral portions of clypeus, in front of antennal fossae, developed and not reduced to narrow margins (Figs. 73, 74). Anterior clypeal margin lacking distinct denticles (Figs. 65, 68). Median portion of clypeus not fey eee re OTe AD RE Ns Nr se once Lateral portions of clypeus reduced to a narrow margin in front of the anten- nal fossae (Fig. 75). Anterior clypeal margin with five to seven denticles. Median portion of clypeus more or TSS euhheit totes sae ees Ree ee RP Rien Dorsum of head with scattered foveolate punctures; spaces between foveolae smooth (Fig. 63). Frontal carinae short, not extending to the level of the posterior margins of eyes in full- face view. Alitrunk in dorsal view with a deep longitudinal furrow at middle (Asia: Philippines) —------- 20 21 RRs LIN te Pals ea ACE. divisus (p. 404) Dorsum of head entirely sculptured with coarse reticulum (Fig. 65). Frontal carinae long, extending to the level of the posterior margins of eyes in 400 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 71 Figures 68-72. Characters on the anterior clypeal margins of Pristomyrmex workers. 68: Anterior clypeal margin entire; 69: Anterior clypeal margin with two teeth; 70: Anterior clypeal margin with three teeth; 71: Anterior clypeal margin with seven teeth; 72: Anterior clypeal margin with three prominences. full-face view. Alitrunk in dorsal with two minute toothlike promi- view lacking a deep longitudinal fur- nences. Dorsal alitrunk convex, not row at the midline (Asia: Malaya) - depressed. Sculpture of the sides of Pe SOPRE PRE ites. ON uno 2 pulcher (p. 408) pedicel segments more coarse (Asia: 21(19). Two or more pairs of erect hairs present Thailand, Malaya, Sarawak, Sabah, on the dorsum of petiole node (Fig. Brunei, Sumatra) rigidus (p. 415) 46). Basal margin of mandible al- 22(17). Promesonotal suture present. Propodeal most straight, without a distinct spines developed and long, in dorsal tooth (Fig. 31). Ventral surface of view joining together at base and clypeus lacking toothlike promi- forming a “fork” (Fig. 26). Alitrunk nences. Dorsal alitrunk more or less in profile with a convex promeson- depressed, with marginate sides. otum and a deeply concave propo- Sculpture of the sides of pedicel deal dorsum (Figs. 22, 23). Dorsum segments lighter and finer (wide- of head, at least on the genae and spread in the east and south of Asia; around the antennal sockets, with occasionally intercepted at entry regular striate sculpture, lacking fo- ports in North America) ———----- veolate punctures or rugoreticulum eae punctatus (p. 410) (Figs. 66, 67) (trispinosus group; In- A pair of hairs present on the dorsum of dian Ocean Islands) _.... 23 petiole node (Fig. 47). Basal margin Promesonotal suture absent. Propodeal of mandible with an acute or broad- armaments in dorsal view usually based triangular tooth (Fig. 35). well separated at the base and not Ventral surface of clypeus usually resembling a fork. In rare case, where the propodeal spines are set close together at the base, the dor- sum of alitrunk, in profile, lacks a deeply concave propodeum (Fig. 25). Dorsum of head smooth or sculptured with foveolate punctures or with rugoreticulum, but never showing regular striate sculpture . 25 23. Propodeal spines in dorsal view diver- gent, in profile almost straight. Figures 73-75. Lateral portions of the clypei of Pristomyrmex Larger species with HW > 1.00, HL workers in front of antennal fossae. 73: P. divisus sp. n.; 74: eNO ESI, 2 SO ans Ane 24 P. pulcher sp. n.; 75: P. punctatus (F. Smith). Propodeal spines in dorsal view subpar- 24. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang allel (Fig. 26), in profile view bent at about a right angle near the base (Fig. 23). Smaller species with HW 0.82—0.90, HL 0.88—1.01, SL 0.80— 0.97 (Indian Ocean Is.: Mauritius, Reunion [.) Dorsum of head and alitrunk entirely covered with regular long coarse striations (Fig. 67). Mesonotum with a pair of strong, blunt digitlike prominences (Fig. 22) (Indian Ocean Is.: Mauritius) Dorsum of head smooth and _ shining, with rugae only present around the antennal fossae, on the genae and sometimes around the centrical disc of dorsal head (Fig. 66). Dorsum of alitrunk smooth and shining. Me- sonotum lacking well-developed digitlike | prominences (Indian Ocean Is.: Mauritius) --------------------- 25(22). Sides of postpetiole with several coarse 26. longitudinal rugae (Fig. 48). In pro- file view, the posterodorsal and pos- teroventral corners of the petiole node right-angled. Palp formula 4,3 (cribrarius group; Africa: Mozam- bique, South Africa) browni (p. 485) trispinosus (p. 488) bispinosus (p. 484) ae cele eat Maree ate cribrarius (p. 418) Sides of postpetiole unsculptured or at most with a single longitudinal ruga (Figs. 49-58). In profile view, the posterodorsal and _posteroventral corners of the petiole node not right-angled. Palp formula 1,3 or 2,2 or 2,3 (quadridens group; Asia, Aus- tralia, Africa) Mandibular dentition arranged as an api- cal tooth + a preapical + a diastema + three small denticles of similar size (Fig. 32). Sometimes, the three small denticles are fused together so that they are not clearly visible, but the length of the masticatory margin covered by the three small denticles is slightly longer than that of diaste- ma. Pronotum either unarmed or armed with a pair of short triangular spines that are shorter than the pro- jexolenll sormnes eo Mandibular dentition arranged as an api- cal + a preapical + a diastema + one or two denticles, and the length of the masticatory margin covered by the one or two denticles is dis- tinctly shorter than that of diastema (Figs. 31, 33, 35). If (very rarely) mandibular dentition not as de- scribed previously but arranged as 2) bo “I 6 Nw ~“ 30. 31. an apical + a preapical + a small denticle + a very short diastema (or diastema indistinct) + two small denticles (Fig. 34), then the prono- tum is armed with a pair of long ro- bust spines that are much longer than propodeal spines (Fig. 24) Pronotum unarmed. Eyes smaller, with three ommatidia in the longest row (Asia: Sumatra) Pronotum armed with a pair of short tri- angular spines. Eyes larger, usually with five to six (rarely with four) om- matidia in the longest row Dorsal surfaces of head and alitrunk only with scattered shallow foveolate punctures; dorsum of alitrunk with a smooth and unsculptured median longitudinal strip (Asia: Indonesia) Dorsal surfaces of head, except for scro- bal areas, and alitrunk entirely cov- ered with well developed coarse ru- goreticum (Asia: Sarawak, Sabah) -- head between frontal carinae either smooth or with some scattered fo- veolate punctures but lacking fove- olate-reticulate sculpture or rugore- tL GUL (ha aes eee ee eae etree Dorsal surfaces of both alitrunk and head with foveolate-reticulate sculp- tunejor mugoreticulum a ee Area of dorsal head between frontal ca- rinae unsculptured and smooth. Dorsum of alitrunk without foveo- late spun ctuie Sue ee ee Dorsal surfaces of both alitrunk and head between frontal carinae with some scattered foveolate punctures Alitrunk, in dorsal view, with a transverse ridge at the approximate position of metanotal groove (Fig. 19). Anterior clypeal margin with three strong teeth (Fig. 70) Alitrunk, in dorsal view, unsculptured, lacking a transverse ridge at the ap- proximate position of metanotal groove. Anterior clypeal margin usu- ally with five to seven small denti- cles (Fig. 71) First gastral tergite with numerous, evenly distributed, suberect hairs. Alitrunk in dorsal view with several short rugae present at the juncture between the pronotum and the me- sonotum (Australia: New South Wales) First gastral tergite lacking any suberect 401 eduardi (p. 440) 28 quindentatus (p. 467) occultus (p. 455) . Dorsal surfaces of both alitrunk and 30 31 40 34 erythropygus (p. 441) 402 eee 76 77 aE pe) Oe ys wes Se 78 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 oe Figures 76-79. Characters on the ventral surfaces of the clypei of Pristomyrmex workers. 76: A tooth; 77: Two toothlike prominences; 78: A short ruga; 79: A long transverse ridge. hairs. Alitrunk in dorsal view without rugae at the juncture between the pronotum and the mesonotum _._.. 33 33. Propodeal spines longer, subequal to or longer than pronotal spines (Fig. 20). Ventral center of clypeus with a weak and short ruga (Fig. 78). Head broader, with HW 0.97-1.34 and CI 103-116 (Australia: New South Wales, SE Queensland) BY esi NIVEA OE wheeleri (p. 478) Propodeum with a pair of teeth or short spines, much shorter than pronotal spines (Fig. 19). Ventral surface of clypeus with a long, well-developed transverse ridge (Fig. 79). Head nar- rower, with HW 0.80-1.08 and CI 93-101 (Australia: New South Wales, Queensland) pect eine ih el quadridentatus (p. 463) 34(31). Pronotum tuberculate, lacking teeth or spines (Africa: Ivory Coast, Ghana, Nigeria, Cameroon, Gabon, Congo and Angola) orbiceps (p. 456) Pronotum armed with a pair of teeth or ES] ON DONS a tS ee 35 35. Petiole and postpetiole without erect hairs (Fig. 50). Frontal carinae ab- sent. Pronotum with a pair of tri- angular short spines. Ventral surface of clypeus with two toothlike prom- inences (Fig. 77) (Africa: Zaire) ------ trogor (p. 476) Petiole and postpetiole with at least one to two pairs of hairs (Figs. 49, 52). Frontal carinae present and usually extending to the level of the poste- rior margins of eyes. In rare cases 80 81 82 Figures 80-82. Lamella, circling the base of the antennal scape of the Pristomyrmex worker, absent (80), entire (81), or with a broad and deep notch (82). 36. 3M. 38(36). 39. where the frontal carinae are very short or absent, the pronotum is armed with a pair of well-devel- oped, long spines that are longer than the distance between their ba- ses (Fig. 21). Ventral surface of clyp- eus either with a transverse ruga, or with a tooth at center, or without any ruga or tooth, but never show- ing two toothlike prominences —_- 36 Pronotum with a pair of triangular short spines, much shorter than the dis- tance between their bases 37 Pronotum with a pair of long spines, lon- ger than distance between their ba- ses (Figs. 21, 25) Petiole node in profile lacking distinct anterior face distinguishable from the upper surface of peduncle (Fig. 53). Larger species, with HW 0.98— 1.04, HL 0.94-1.02, EL 0.22-0.24 (Asia: Philippines) flatus (p. 443) In profile, anterior face of petiole node distinct from the dorsal surface of peduncle (Fig. 52). Smaller species, with HW 0.77—0.94, HL 0.82-0.94, EL 0.14-0.18 (Asia: Philippines) -— collinus (p. 432) Propodeal spines short or moderately long, much shorter and slender than pronotal spines (Fig. 21) Propodeal spines exceptionally long, subequal in length to or slightly lon- ger than pronotal spines (Fig. 25) (Australia: Queensland) ------------------ eS he abe seh ea A Ee wilsoni (p. 481) Petiole node in profile lacking distinct anterior face distinguishable from the upper surface of peduncle (Fig. 55). Clypeus unsculptured, lacking a median longitudinal carina. Frontal carinae short, not extending to the level of the posterior margins of eyes (Asia: Philippines) longispinus (p. 450) In profile, anterior face of petiole node distinct from the dorsal surface of peduncle (Fig. 54). Clypeus with a median longitudinal carina. Frontal carinae long, extending to the level of the posterior margins of eyes (Asia: Philippines) _ curvulus (p. 437) 39 REVISION OF THE ANT GENUS PRISTOMYRMEX ° 40(30). Petiole and postpetiole lacking erect Ventral surface of hairs (Fig. 50). clypeus with two toothlike promi- nences (Figs. 77) (Africa: Ghana, Cameroon, Capen! Angola, Kenya, Zaire, Sudan) Petiole and postpetiole, respectively, with one to five pairs of erect hairs. Ventral surface of clypeus either with a transverse ruga (Figs. 78, 79) or with a toothlike prominence (Fig. TASH ry oy Seen en UR noe ate a Smaller species (HW 0.82-1.02, HL 0.82-1.02, EL 0.14—0.20). Ventral center of clypeus with a toothlike prominence. Usually one to two pairs of hairs present, respectively, on the dorsal surfaces of petiole node and postpetiole (Asia and Pa- cific Is.: Papua New Guinea; Indo- nesia; Pohnpei Is.) africanus (p. 423) 41 fabs 02a pee quadridens (p. 459) Larger species (HW 1.22-1.24, HL 1.10-1.16, EL 0.24—0.25). Ventral surface of clypeus with a transverse ruga, lacking a toothlike prominence at center. Four to five pairs of short hairs present, respectively, on the dorsal surfaces of petiole node and postpetiole (Fig. 51) (Asia: New Guinea) evenly distributed, erect or suberect hairs. Petiole node with a single evenly blunt-rounded apex (Fig. 56) (Asia: Philippines) _ First gastral tergite lacking erect hairs. Peuole node not showing a single evenly blunt-rounded apex but with a higher anterodorsal angle than the posterodorsall(Hige oi) pee Masticatory margin of mandibles with five teeth; diastema very short or in- distinct between the preapical and the third tooth (Fig. 34). Basal mar- gin of mandible with a central, broadly curved lobe. Anterior clyp- eal margin with three prominences, that is, a median tooth and a broad, low convex lobe on each side (Fig. 72) (Asia: Sarawak, Sabah, Borneo) Masticatory margin of mandibles with three to four teeth; diastema distinct and long, present between the preapical and the third tooth (Figs. 31, 33). Basal margin of mandible almost straight, without a distinctly curved lobe. Anterior clypeal mar- gin usually with five to seven denti- cles (Fig. 71) nitidissimus (p. 453) 42(29). First gastral tergite with numerous, hirsutus (p. 449) 43 trachylissus (p. 474) 44, 45. A6. 47(45). 49. Wang 403 Pronotal spines exceptionally long, usu- ally exceeding 0.40 (very rarely 0.37), usually ‘longer than the dis- tance between the bases of two pro- notal spines (Fig. 24) (Asia: Suma- tra, Java, Malaya, Sarawak, Sabah, Borneo, Philippines) bicolor (p. 425) Pronotum armed with a pair of teeth or spines ($0.32; Figs. 14, 15, 17, 18) that are always shorter than the dis- tances that separate their bases ___ 45 Propodeal spines long (0.19-0.30), much longer than the pronotal armaments (Bigs 16) Spalpstormulal 213 sae 46 Propodeal spines short (0.04—-0.13), usu- ally shorter than but sometimes slightly longer than the pronotal ar- maments (Figs. 14, 15), palp for- mula 1,3 Postpetiole in dorsal view much broader than long, with PPI 133-150. An- tennal scapes shorter (SL 0.70-0.82, SI 81-93) (Australia: Queensland) Jn. Wee ew ae eae foveolatus (p. 446) Postpetiole in dorsal view slightly broad- er than long, with PPI 109-121. An- tennal scapes longer (SL 0.86—-0.98, SI 97-103) (aera: Queensland) ey oe en EN thoracicus (p. 473) Petiole node in profile slightly longer than high, with the anterodorsal an- gle on approximately the same level as or weakly higher than the poster- odorsal; dorsum and sides of petiole node with seven to eight foveolate punctures (Fig. 58) Nees Sarawak) Dimi Fra es ah as modestus (p. 452) Petiole node in profile higher than long, with the anterodorsall angle disanee ly elevated above the posterodorsal; dorsum and sides of petiole node without foveolate punctures (Fig. DID) ence ek Ree eS Rael eee 48 Ventral surface of clypeus lacking a cen- tral prominent tooth but usually bearing a transverse ruga (Asia: Ma- laya, Singapore, Sar Ale Borneo, Sabah, Philippines) _. costatus (p. 434) Ventral surface of clypeus with a central prominent tooth (Fig. 76) __- 49 Pronotum with a pair of moderately long spines (0.14—0.20), usually much longer (sometimes slightly longer) than the propodeal armaments (0.07—0.13) (Figs. 18) (Asia: Malaya, Thailand, Nepal, Burma, China) -—--- Soe ese Re 27 sulcatus (p. 469) Pronotum with a pair of teeth or short spines (0.06—0.10), usually similar in length to or slightly shorter than the propodeal armaments (0.04—0.12) 404 (Figs. 14, 15) (Asia: Sumatra, Sula- wesi, Malaya, Sarawak, Sabah, Thai- land, Philippines, Taiwan, Japan) —- brevispinosus (p. 428) THE PUNCTATUS GROUP Worker. Small to medium sized (HL 0.62—0.98, HW 0.64—0.94, TL 2.62-3.44), with the following combination of charac- ters: (1) Masticatory margin of mandibles with three to four teeth arranged as the strongest apical + the second strongest preapical + a long diastema + two small basal denticles of similar size (or a broad basal tooth). z (2) Palp formula 5,3 in four Oriental species and 4,3 in the single South African species. (3) Frontal carinae present. (4) Lateral portions of clypeus reduced to a narrow margin in front of the antennal fossae in three species (P. punctatus, P. rig- idus, and P. fossulatus) but developed in the other two species (P. divisus and P. pulcher). (5) Frontal lobes indistinct or absent. (6) Lamella that encircles the base of antennal scape, entire. (7) Alitrunk in profile showing a contin- uous convex dorsum and in dorsal view lacking sutures. (8) Pronotum unarmed. (9) Propodeum with a pair of spines. (10) Petiole node in profile more or less wedge-shaped, lacking distinct anterior face distinguishable from the upper sur- face of peduncle. This group has five species. Four occur in the Oriental region, one of which (P. punctatus ) has spread to warm-temperate areas in the eastern Palaearctic. The re- maining species (P. fossulatus) is confined to South Africa. Pristomyrmex punctatus is a unique species within the genus. It is the only Pristomyrmex extending its range to the temperate zone, and it is further char- acterized by the possession of a unique life history that may preadapt it for dispersal by natural and human transport. Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 The punctatus group is closely related to the cribrarius and quadridens groups because all female castes of these species groups possess a distinct diastema after the preapical tooth on the masticatory margin of mandibles (except in P. trachylissus, which has five teeth on the masticatory margin). Though sharing the previously described mandibular character, the trispi- nosus group is relatively distant from the punctatus group because it possesses so many autapomorphic characters, for ex- ample, frontal carinae absent, dorsal ali- trunk with a promesonotal suture or im- pression, propodeal spines in dorsal view showing a “fork”, some regular striation present on the dorsal surfaces of head and alitrunk. The punctatus group differs from the cribrarius group by lacking a pair of pro- notal spines in the workers and by showing the anterior face of the petiole node not distinct from the upper surface of pedun- cle in the female castes. The punctatus group differs from the quadridens group by possessing palp formulae of 5,3 and 4,3 (the quadridens group possesses palp for- mulae of 2,2, 1,3 and 2;3). Pristomyrmex divisus sp. n. Figures 83-84 Diagnosis (Worker). Lateral portions of clypeus in front of antennal fossae, devel- oped, not reduced to margins, so that the antennal fossae are placed well behind the anterior clypeal margin; dorsal head only — with scattered foveolate punctures. Holotype Worker (MCZC). TL 3.06, HL 0.76, HW 0.80, CI 105, SL 0.82, SI 103, EL 0.22, PW 0.56, AL 0.74. Paratypes, 35 workers MHNG). (MCZC, BMNH, ANIC, | Worker. TL 3.06-3.40, HL 0.72-0.82, | HW 0.74-0.86, CI 98-111, SL 0.78—0.90, | SI 98-110, EL 0.21-0.24, PW 0.53-0.64, | AL 0.72-0.80, PPW 0.26-0.30, PPL 0.18— | 0.22, PPI 123-156 (n = 20). Mandibles with a few longitudinal rugae — but smooth near the masticatory margin. | Dentition of the masticatory margin of REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 405 Figures 83-84. Pristomyrmex divisus sp. n. 83: Worker head, full-face view; 84: Worker, lateral view. mandible: the strongest apical tooth + the second strongest preapical + a long dia- stema + a broad, truncated basal tooth (or two minute denticles). A weak minute prominence present about midway on the basal margin of mandible. Clypeus with a strong median longitudinal carina extend- ing through the frontal area; on each side of the median clypeal carina, a few addi- tional rugae are usually present. Anterior clypeal margin lacking denticles. Median portion of clypeus higher than frontal area; lateral portions of clypeus developed, not reduced to margins. Ventral surface of clypeus lacking any toothlike prominences but usually with a few rugae. Palp formula 5,3. Frontal carinae short, not extending to the level of the posterior margins of eyes. Antennal scrobes absent. Frontal lobes ab- sent; thus, the antennal articulations are completely exposed. Antennal scapes, when lying on the dorsal head, surpassing the occipital margin of head by one-sixth to one-fifth of their length. Eyes large and prominent, containing § to 10 ommatidia in the longest row. Dorsum of alitrunk in dorsal view marginated, more or less de- pressed, and usually with a deep longitu- dinal furrow at middle. Pronotum un- armed. Propodeal spines well developed, acute and long, much longer than the dis- tance between their bases. Metapleural lobes small, dentiform, and acute. Petiole in profile with a long peduncle; dorsum of peduncle, together with the anterior face of petiole node, forming a long declivity that reaches the top of petiole node. Ven- tral surface of petiole lacking any process. Postpetiole in profile with a convex dor- sum, in dorsal view somewhat transverse- rectangular and much broader than long. Dorsum of head with numerous large and shallow foveolate punctures; space be- tween foveolae smooth; ventral head with denser foveolate punctures. Dorsal surface of alitrunk with reticulate rugae. Petiole al- ways, and postpetiole usually, with a coarse longitudinal ridge on each side. In dorsal view, petiole node and postpetiole each usually bounded by a rim; dorsums of both petiole and postpetiole, except for rims, very smooth and polished. Gaster unsculp- tured. Dorsal surfaces of head and alitrunk with numerous erect to suberect short hairs. A pair of hairs present, respectively, near the top of both petiole node and post- petiole. First gastral tergite without hairs. Two or three pairs of long, forward-pro- jecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect to suberect short hairs. Color uniform reddish-brown; appendages sometimes slightly lighter. Queen and Male. Unknown. Comments. This species is so far known only from the Philippines, and its closest relative without doubt is P. pulcher, from Malaysia. The workers of two species share the following three characters that are not seen in the other three members of the 406 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 85-86. Pristomyrmex fossulatus (Forel). 85A: Worker head, full-face view; 85B: Showing a short transverse ruga on the ventral clypeus; 86: Worker, lateral view, hairs omitted from the petiole node and postpetiole. punctatus group (P. punctatus, P. rigidus, and P. fossulatus): (1) lateral portions of clypeus, in front of the antennal fossae, de- veloped, making the antennal fossae well behind the anterior clypear margins; (2) anterior clypeal margin lacking distinct denticles; and (3) the median portion of clypeus not flat but somewhat concave. In the workers of P. punctatus, P. rigidus, and P. fossulatus, the anterior clypeal margin is equipped with five to seven denticles, and the lateral portions of clypeus in front of the antennal fossae are reduced to margins (in other words, the antennal fossae reach the lateral anterior margins of clypeus), and the median portion of clypeus is more or less flat. The workers of P. divisus are easily sep- arated from those of P. pulcher. The ce- phalic dorsum shows rugoreticulum in P. pulcher but scattered foveolate punctures in P. divisus; the frontal carinae do extend to the level of the posterior margins of eyes in P. pulcher but not so in P. divisus; a pronounced median longitudinal furrow is present on the dorsal surface of alitrunk in P. divisus but absent in P. pulcher. Holotype Worker. Philippines: Duma- guete, 1949, J. W. Chapman. Paratypes. 18 workers with same data as holotype; 14 workers, Philippines: Duma- guete (J. W. Chapman); three workers, Philippines: Dumaguete, Silliman Univer- sity, 9.v.1949 (Domingo Empeso). Ecological Information. Unknown. Pristomyrmex fossulatus (Forel) Figures 85—86 Tetramorium (Xiphomyrmex) fossulatum Forel, 1910: 428. Syntype workers, South Africa: Natal, Will Broak (Wroughton) (MHNG) lexamined]. Pristomyrmex fossulatus (Forel) Santschi, 1916: 51. Diagnosis (Worker). Masticatory margin of mandible with a long diastema after the preapical tooth; palp formula 4,3; eyes with § to 10 ommatidia in the longest row; pronotum lacking teeth or spines; dorsal surfaces of head and alitrunk with scat-— tered foveolate punctures. Worker. TL 2.63-2.92, HL 0.62-0.71, HW 0.64—0.75, CI 98—106, SL 0.56—0.61, SI 81-88, EL 0.17—0.18, PW 0.44—0.50, AL 0.64—0.74, PPW 0.26—0.28, PPL 0.16— ONSRE RISA Noss (na—s >) Mandibles smooth and shining. Denti- tion of the masticatory margin of mandi- ble: an apical tooth + a preapical tooth + a long diastema + a truncated basal tooth. Basal margin of mandible lacking a tooth- like prominence or curved lobe. Clypeus with a strong median longitudinal carina. Anterior clypeal margin with a median denticle and two to three others on each side, but sometimes two lateral denticles | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang are fused together. Lateral portions of clypeus reduced to margins, and antennal fossae reaching the lateral anterior mar- gins of clypeus. Ventral center of clypeus with a short transverse ruga. Palp formula 4,3. Frontal carinae extending to the level of the posterior margins of eyes. Antennal scrobe short, shallow, but distinct, mar- gined by the frontal carina and a longitu- Final ruga. Frontal lobes absent; thus, the anal articulations are completely ex- posed. Antennal scapes, when lying on the dorsal head, close to the occipital margin of head. Eyes large, containing 8 to 10 om- matidia in the longest row. Pronotum with a pair of blunt tubercles, lacking teeth or spines. Propodeum armed with a pair of spines, about as long as the distance be- tween their bases. Metapleural lobes sub- triangular. Petiole node in profile wedge- shaped, with a triangular apex. Subpetiole with a narrow flange. Postpetiole in profile higher than long, aa a rounded dorsum, in Moral view transverse-rectangular and much broader than long. Dorsum of head, except for the scrobal areas, with numer- ous scattered foveolate punctures. Similar but sparser punctures present on the dor- sal surface of alitrunk. Petiole and post- petiole each usually with a longitudinal ruga on each side. Gaster unsculptured. Several pairs of hairs present on the dor- sum of head beyond the level of the an- tennal insertions. A row of forward-pro- jecting hairs present on the anterior clyp- eal margin. Hairs on the rest of the body as follows—mesonotum (one pair), petiole (zero to one pair), and postpetiole (one to two pairs dorsally)—frequently lost by abrasion (Bolton, 1981: p. 286). First gas- tral tergite lacking erect or suberect hairs. Scapes and tibiae with short hairs. Color reddish-brown; appendages yellow-brown. Queen. I have not seen the queen of this species, but Menozzi (1942: 172) gave a description of this caste. Male. Unknown. Comments and Discussion. The position of P. fossulatus within Pristomyrmex is somewhat complicated. Pristomyrmex fos- 407 sulatus occurs only in South Africa and shares certain character states with P. or- biceps, an African member of the quadri- dens group. The workers of P. fossulatus are similar to those of P. orbiceps in (1) the masticatory margin of mandible with a diastema, (2) the pronotum with a pair of broad and blunt tubercles, (3) the size of eyes, (4) the length of propodeal spines, and (5) hairs on the head and body (e.g., two to three pairs of erect hairs along fie frontal carinae behind the level of anten- nal insertions, a pair on the occipital cor- ners, a pair on the mesonotum). But it is very hard for me to place P. fossulatus into the quadridens group because this species has four segments of the maxillary palpi in the workers. In the 25 members of the quadridens group, the maxillary palp of 18 species examined, including P. orbiceps, is one or two segments. Pristomyrmex fossulatus, however, shares important similarities with four Asian species of the punctatus group @P. divisus, P. pulcher, P. punctatus, and P. rig- idus). These critical characters, as shown in the workers, include (1) a high palp for- mula, (2) the masticatory margin of man- dible with a diastema after the preapical tooth, (3) pronotum without teeth or spines, (4) anterior face of the petiole node indistinguishable from the upper surface of its anterior peduncle, and (5) postpeti- ole in dorsal view much broader than long. Thus, P. fossulatus is assigned to the punc- tatus group. P. fossulatus « can be separated from the four species as follows: In the workers of P. fossulatus, the propodeal spines are moderately long; the antennal scapes, falling into the range 0.56 to 0.61 (SI 81-88), are only close © the occipital margin of head, when laid on the dorsal head; the dorsum of the alitrunk has only scattered foveolate punctures. In P. divi- sus, P. pulcher, P. punctatus, and P. rigidus, the propodeal spines are well developed and long, much longer than the distance henwecnt their asoe. the antennal scapes, with the length 0.70 to 0.94 (usually above 0.78) and SI 91-118 (usually above 98), 408 obviously surpass the occipital margin of head; the dorsum of the alitrunk is covered fully with a developed rugoreticulum. It is possible that P. fossulatus evolved from the P. divisus-P. pulcher-P. punctatus-P. rigidus ancestor but later was divergent from the clade consisting of the four Asian species. It must be pointed out that if P. fossu- latus is designated as a member of the quadridens group, the cribrarius group, also possessing 4,3 palp formula, would become insignificant. As a result, P. cri- brarius (another African species) would fall into the quadridens group. Pristomyrmex fossulatus differs from two African species of the quadridens group (P. orbiceps and P. africanus) in the workers as follows: P. fossulatus Palp formula 4,3 Dorsum of head between frontal cari- nae with numerous foveolate punc- tures Dorsum of alitrunk with some foveolate punctures Petiole node in profile wedge-shaped, with a triangular apex P. orbiceps Palp formula 1,3 Dorsum of head between frontal cari- nae smooth and shining, except for few punctures bordering frontal cari- nae Dorsum of alitrunk smooth and shining Petiole node in profile relatively mas- sive, and broad-rounded dorsally P. fossulatus Palp formula 4,3 Ventral surface of clypeus with a weak transverse ruga Eyes larger, with 8 to 10 ommatidia in the longest row Petiole node in profile wedge-shaped, with a triangular apex, lacking a dis- tinct concave between the anterior face of the node and the dorsal sur- face of the peduncle Pronotum with a pair of tubercles Dorsal head behind the level of anten- Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 nal insertions with three to four pairs of hairs; dorsal alitrunk at most with a pair of hairs P. africanus Palp formula 1,3 Ventral surface of clypeus with two strong toothlike prominences Eyes smaller, with four to five ommatid- ia in the longest row Petiole node in profile not wedge- shaped; a distinct concave present be- tween the anterior face of the node and the dorsal surface of the pedun- cle Pronotum usually with a pair of teeth or spines but sometimes with a pair of tubercles Dorsal head behind the level of anten- nal insertions with numerous (at least 10 pairs of) hairs; dorsal alitrunk with several pairs of hairs Distribution. This species was previous- ly known only from the type series col- lected from South Africa (Bolton, 1981). Two more specimens (ANIC, MCZC) ex- amined here add the following records: South Africa: Cape Province, Grahams- town, Old P. E. Road (L. H. Weatherill). Ecological Information. Unknown. Pristomyrmex pulcher sp. n. Figures 87-88 Diagnosis (Worker). Lateral portions of — clypeus in front of antennal fossae, devel- | oped, not reduced to margins, so that the antennal fossae are placed well behind the — anterior clypeal margin; dorsal surfaces of — head and alitrunk, as well as the sides of pronotum, with well-developed coarse ru- goreticulum. Holotype Worker (BMNH). TL 2.96, | HL 0.76, HW 0.72, CI 95, SL 0.82, ST 114, | EL 0.20, PW 0.53, AL 0.76. Paratypes, 11 | workers (BMNH, MCZC, ANIC). Worker. TL 2.70-3.04, HL 0.70-0.77, | HW 0.69-0.77, CI 95-100, SL 0.76-0.84, | ST 103-114, EL 0.19-0.21, PW 0.50-0.54, | AL 0.70-0.78, PPW 0.25—-0.29, PPL 0.17- | 0.20, PPI 135-156 (n = 11). REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 409 Figures 87-88. Pristomyrmex pulcher sp. n. 87A: Worker head, full-face view; 87B: Showing a curved ruga on the ventral clypeus; 88: Worker, lateral view. Mandibles usually with a few longitudi- nal rugae but smooth near the masticatory margin. Dentition of the masticatory mar- gin of mandible: the strongest apical tooth + the second strongest preapical + a long diastema + a broad basal tooth usually fused by two small denticles. A broad- based triangular short tooth present about midway on the basal margin of the man- dible. Clypeus with sculpture consisting of a strong median longitudinal carina, a transverse carina (sometimes curved or weak or broken), and a few additional short carinae. Anterior clypeal margin lacking distinct denticles. Lateral portions of clypeus, in front of antennal fossae, de- veloped, not reduced to narrow margins. Ventral surface of clypeus usually with a somewhat “M” ruga that is sometimes in- terrupted at the middle. Palp formula 5,3. Frontal carinae strong, extending to the level of the posterior margins of eyes. Slightly impressed scrobal areas present lateral to the frontal carinae. Frontal lobes nearly absent; thus, the antennal articula- tions are completely exposed. Antennal scapes, when lying on the dorsal head, sur- passing the occipital margin of head by one-sixth to one-fifth of their length. Eyes large and prominent, containing seven to nine ommatidia in the longest row. Occip- ital margin straight or feebly concave. In dorsal view, dorsum of the alitrunk mar- ginated and rather depressed. Pronotum unarmed. Propodeal spines well devel- oped, acute, and long, much longer than the distance between their bases. Meta- pleural lobes triangular. Pedicel segments as in Figure 88. Dorsum of petiole pedun- cle, together with the anterior face of the node, forming a long declivity that reaches the apex of the node. Ventral surface of petiole without any appendages or with only a very narrow rim. Postpetiole in pro- file convex dorsally, in dorsal view trans- verse-rectangular and much broader than long. Dorsal surfaces of head and alitrunk, as well as the sides of pronotum, with de- veloped coarse rugoreticulum. Sides of the rest of the alitrunk irregularly rugulose. Antennal scapes with a few longitudinal rugae. Sides of both petiole and postpeti- ole usually with a few coarse longitudinal rugae. In dorsal view, petiole and postpe- tiole each bounded by a rim; dorsal sur- faces of petiole and postpetiole, except for rims, very smooth and polished. Gaster unsculptured. Dorsal surfaces of head and alitrunk with numerous erect or suberect hairs. A pair of similar hairs present bilat- erally near the top of petiole and two pairs usually on the dorsal postpetiole as in Fig- ure 88. Two or three pairs of long, for- ward-projecting hairs present near the an- terior clypeal margin. Scapes and tibiae with some scattered erect or suberect 410 hairs. Gaster lacking erect or suberect hairs. Femurs, tibiae light-yellow, in con- trast with head antennae, alitrunk, tarsi, pedicel segments, and gaster, which are reddish-brown. Queen and Male. Unknown. Comments. Although P. pulcher, an Ori- ental species, is very similar, in appearance of the workers, to P. punctatus, the two species are in fact not closely related with- in the punctatus group. Pristomyrmex pulcher, together with P. divisus, consti- tutes a clade that is the sister group of the clade formed by the species P. rigidus and P. punctatus. The separation of the two clades is summarized under P. divisus. Ad- ditional characters separating the workers of P. pulcher from those of P. punctatus are as follows: P. pulcher A broad, short tooth present on the bas- al margin of mandible Dorsum a petiole node with only a pair of erect hairs Leg bicolored, with femur and _ tibia light-yellow and tarsus reddish-brown Sculpture on the dorsal surfaces of head and alitrunk, and the sides of petiole and postpetiole relatively coarse Propodeal spines shorter, more robust, and slightly bent basally P. punctatus Tooth of the basal margin absent or in- conspicuous Dorsum of petiole node with two or more pairs of hairs Leg uniformly colored, light-red or red- teie brown Sculpture on the dorsal surfaces of head and alitrunk, and the sides of petiole and postpetiole relatively fine Propodeal spines relatively slender, and straight The separation of P. pulcher from P. div- isus is discussed under P. divisus. Holotype Worker. Malaysia: Negri Sem- bilan, Pasoh For. Res., primary forest, lit- ter sample, 3.iv.1994 (L. Ficken). Paratypes. Five workers with same data Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 as holotype; three workers, Malaysia: Ne- gri Sembilan, Pasoh For. Res., litter sam- ple, 3.iv.1994 (M. Brendell, K. Jackson and L. Ficken); three workers, Malaysia: Neg. Sembilan, Pasoh For. Res., litter sample, xi.1994 (M. Brendell, K. Jackson, and S. Lewis). Ecological Information. This species has been collected from a little sample taken in primary forest. Pristomyrmex punctatus (F. Smith) Figures 89-93 Myrmica punctata F. Smith, workers, Indonesia: Bachian I. (OXUM) [examined]. Pristomyrmex punctatus (F. Smith) Mayr, 1886: 361. Pristomyrmex pungens Mayr, 1866: 904. Holotype worker, Malaysia: Malacca (?) (NHPS) [examined]. Syn. n. Pristomyrmex japonicus Forel, 1900: 268. Syntype workers, Japan: Osaka (K. Yamada) (MHNG) [ex- amined]. [Synonymy by Viehmeyer, 1922: 207]. 1860: 108. Syntype (A. R. Wallace) Diagnosis (Worker). Eyes with eight or more ommatidia in the longest row; pron- otum unarmed; propodeal spines straight, and long, much longer than the distance between _ their bees: dorsal surfaces of both head and alitrunk covered fully with rugoreticulum; dorsum of petiole node with two or more pairs of hairs. Worker wily 2462-32225 Miley 0270201845 HIW 0168-0584 C942 1055S L1057/8=0:S61 SI 102-118, EL 0.15—-0.18, PW 0.48-0.56, AL 0.70—-0.84, PPW 0.24—0.27, PPL 0.17— OVA, JAP MAO) Gi —= 70), Mandibles usually with a few fine lon- gitudinal rugae but smooth near the mas- ticatory margin. Dentition of the mastica- tory margin of mandible: the strongest api- cal tooth + the second strongest preapical + a long diastema + a broad basal tooth | ' (or two small basal denticles). Basal mar- — gin of mandible rather straight, lacking a Gustine tooth. Clypeus eine leliiee’ more or less depressed, with a median longitudinal — carina extending posteriorly through the | | frontal area. In some specimens, a few | short weak rugae present on each side of | the median carina of the clypeus. Anterior — clypeal margin equipped with a row of REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang sale [Z 41] Figures 89-92. Pristomyrmex punctatus (F. Smith). 89A: Worker head, full-face view; 89B: Showing a curved ruga on the ventral clypeus; 90: Worker, lateral view; 91: Ergatoid queen head, full-face view; 92: Ergatoid queen, lateral view. denticles, but sometimes median denticle indistinct or absent or becoming a broad- truncated lobe. Lateral portions of clypeus reduced to margins, so that the antennal fossae reach the anterior clypeal margin. Ventral surface of clypeus usually with a curved ruga as in Figure 89B. Palp for- mula 5,3. Frontal carinae distinct, extend- ing to the level of the posterior margins of eyes. Antennal scrobes weak. Frontal lobes almost completely absent; thus, the anten- nal articulations are entirely exposed. An- tennal scapes, when lying on the dorsal head, surpassing the occipital margin of head by one-sixth to one-fifth of their length. Eyes large, with eight or more om- matidia in the longest row. Occipital mar- gin feebly concave. Dorsum of the alitrunk in dorsal view marginated and more or less depressed (Fig. 93). Pronotum unarmed. Propodeal spines long, acute, but slender. Metapleural lobes dentiform and acute. Petiole node in profile wedge-shaped, with a triangular apex. Postpetiole in profile convex dorsally; in dorsal view transverse- rectangular, much broader than long, and also broader than the petiole. Dorsal sur- faces of head and alitrunk as well as sides of pronotum covered fully with rugoreti- culum, but scrobal areas with only several transverse rugae. Sides of the rest of the alitrunk with numerous irregular rugae. Sides of petiole and postpetiole usually with a few fine longitudinal rugae; some- times rugae very weak and broken. Gaster unsculptured. Dorsal surfaces of head and alitrunk with numerous erect to suberect long hairs. Two (or more) pairs of hairs 412 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figure 93. Showing that the worker of Pristomyrmex punctatus (F. Smith) has a more or less depressed dorsum of alitrunk. present, respectively, bilaterally on the dorsums of petiole node and postpetiole, of which usually a pair shorter and the oth- er pair longer. First gastral tergite lacking erect or suberect hairs. A few pairs of long, forward-projecting hairs present near the anterior clypeal margin that are symmet- rical on the two sides of the midpoint. Scapes and tibiae with numerous erect to suberect short hairs. Color reddish-brown, but sometimes the gaster much darker or the appendages slightly lighter. Ergatoid Queen. ae, 3.60. 3972: FUE 0.86, 0.88: HW 0.94, 0.94; CI ude 109: SL 0.89, 0.91; SI 97, 97; EL 0.23, 0.23: PW 0.66, 0.68; AL 0.94, 0.96; PPW. 0.31, 0.32 PPE 0.23) 0324- PRT T3335) G7 = 2): Closely resembling the worker in the structure of mandibles, clypeus, petiole, postpetiole and gaster and also in sculp- ture, color, and pilosity. But the head with three ocelli; eyes larger; pronotum and propodeum narrower than those of work- er; Mesonotum more convex; an impres- sion present at the approximate positions of promesonotal suture and of metanotal groove, respectively; propodeal spines stronger than in worker. Wing absent, but the rudimentary vestige of a wing is pres- ent on the each lateral margin of the me- sonotum. Queen. Unions Male. TL 3.22, HL 0.60, HW 0.57, CI 95) SEAOSs Sie 325 EIWilis OO eerie, 0.32. PW 0.74, AL 1.04, PPW 0.26, PPL 0.17, PPI 153 (n = 1; one specimen [MCZC] collected from Nara, Japan, by Silvestri on July 21, 1925, was examined). Head, including the eyes, broader than long. Clypeus transverse, with a median short carina. Frontal area with a median longitudinal carina. Frontal carinae short, slightly beyond the posterior margins of — antennal sockets. Palp formula 5,3. On the mesoscutum, notauli pronounced, forming a Y shape; parapsidal furrows superficially _ impressed. Propodeum with a pair of teeth. Metapleural lobes subtriangular. Middle and hind tibiae each with a simple spur. Petiole node wedge-shaped, with a triangular apex; dorsum ae petiole pedun- cle a ming a declivity that reaches the top of the monies Postpetiole i in profile rounded dorsally, in dorsal view transverse-rectan- gular and distinctly broader than long. Dorsum of head smooth and shining, ex- cept for few short rugae present behind — the posterior margin of clypeus. Pronotum — REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang and mesoscutum smooth, except for those marked sutures, but mesoscutellum and propodeum sculptured with several longi- tudinal rugae. Sides of petiole with a few rugae. All dorsal surfaces with abundant erect or suberect hairs, but hairs on the legs and on the scapes somewhat ap- pressed. Colour reddish-brown; wing light-yellow. Comments and Discussion. The separa- tion of P. punctatus from both P. divisus and P. pulcher is summarized in the dis- cussions of both P. divisus and P. pulcher. The following characters can be used to separate the workers of P. punctatus from those of its closest relative, P. rigidus: P. punctatus Dorsum of petiole node with two or more pairs of hairs Tooth absent or inconspicuous from basal margin of mandible Dorsal surfaces of head and _alitrunk, and the sides of petiole and postpe- tiole more finely sculptured Propodeal spines relatively slender Clypeus with a median longitudinal ca- rina that meets the anterior clypeal margin Dorsum of alitrunk in dorsal view, mar- ginated, and more or less depressed Ventral surface of clypeus with a curved ruga but lacking distinct toothlike prominences P. rigidus Dorsum of petiole node only with a sin- gle pair of hairs In type specimens, a strongly prominent tooth present on basal margin of mandible Dorsal surfaces of head and alitrunk, and the sides of petiole and postpe- tiole more coarsely sculptured Propodeal spines relatively robust Median clypeal carina often not reach- ing the anterior clypeal margin Dorsum of alitrunk in dorsal view, con- vex Ventral surface of clypeus usually with two minute toothlike prominences 413 In addition, an ergatoid queen caste is present in P. punctatus but not seen in P. rigidus, while a normal queen caste exists in P. rigidus but has not been found in P. punctatus. Without a doubt, P. punctatus originat- ed in the subtropics or tropics of Asia, as its three close relatives, P. rigidus, P. pulcher, and P. divisus, are restricted to Sumatra, Malaysia, Brunei, Thailand, and the Philippines, respectively. But P. punc- tatus has a very large range, from New Guinea, Indonesia to Malaysia, Thailand, and then north to China and Japan, indi- cating its exceptional dispersal ability and tolerance of temperate climates. Pristo- myrmex punctatus has shown some ten- dency for dispersal by humens. Intercep- tion records from ports in North America suggest that humen commerce may have played a role in this species’ spread in tem- perate Asia. The chromosome numbers of this spe- cies, reported by Imai (1966) and Itow et al. (1984), are 2n (diploid) = 24 in the ce- rebral ganglion cells of the workers and n (haploid) = 12 in the spermatocytes of the males. The larva of this species was de- scribed by Wheeler and Wheeler (1954). Biology. It appears that P. punctatus oc- curs in open habitats. My impression stems from field experience in China as well as from data associated with speci- mens. This species lacks a normal queen caste; mature colonies usually contain sev- eral thousand workers and a few males, but ergatoid queens are rarely found. Eggs are normally laid by workers and can de- velop into workers or males. Ergatoid queens, when present, can also lay eggs. Brood production begins in April and lasts until the end of September. Young workers remain inside the nest and lay eggs. Older workers forage but lose the ability to lay eggs. Nests are often constructed in leaf litter from June to August but in the soil around trees from September to October. Sometimes arboreal nests are constructed on dead standing trees or in partially dead parts of living trees. The nest entrances of 414 the arboreal nests and those under rotten wood are often covered with soil particles. See the following references for informa- tion on these and other aspects of the bi- ology of P. punctatus (Itow et al., 1984; Mizutani, 1980, 1982; Tsuji, 1988a, 1988b, 1988c, 1990a, 1990b, 1994, 1995; Tsuji and Itd, 1986). Material Examined (ANIC, AMNH, BMHH, BMNH, IZAS, LACM, MCZC, NHMV, USNM). New Guinea: Trian Jaya Ransik (Shah); NETH. Vogelkop, Dano- waria (J. L. Gressitt); Vogelkop, Manok- wari, 75 m (D. Elmo Hardy); Vogelkop, Fak Fak, S. coast of Bomberai, 10 to 100 m (J. L. Gressitt). Borneo: Sarawak, Nanga Pelagus near Kapit, 180 to 585 m (T. C. Maa); Sarawak, Merirai Valley (T. C. Maa); Sarawak, Bau, Lake Area (T. C. Maa); North Borneo, Tawau, Quoin Hill, Cocoa Res. Sta. (K. J. Kuncheria); North Borneo, Forest Camp, 19 km N of Kalabakan (K. J. Kuncheria); North Borneo (SE), Forest Camp, 9.8 km SW of Tenom (Y. Hirashi- ma). Java: S. Coast, Sukawayani, 2 m, jun- gle remnant (J. L. Gressitt). Philippines: Samar (McGregor); Mindoro, San Jose (E. S. Ross); P.R. Mindanao, Talacogon, 8.32°N, 125.39°E, relict rainforest, on Agusan River (B. B. Lowery). Singapore (H. Overbeck). Thailand: Chiang Mai Prov., Chiang Rai, Fang Hort. Res. Sta.(D. G. Furth); Chiang Mai Prov., 18.70°N 98.82°E, Mae Wang Dist., Ban Huai Thong, 360 m, agricultural area (orchard), #96-116 (foragers on bamboo shoot) (R. R. Snelling and Saowapa Sonthichai). Viet- nam: Perfume Pagoda (P. Jolivet); Cuc Phuong Forest (P. Jolivet). China: Hainan Is. (J. L. Gressitt); Guangxi, Xingping (D. G. Furth); Guangxi, Guilin, Qixing Park (D. G. Furth); Hong Kong, N.T., Campus CGaUPAE Ke Shatines 2223 SeNe eA aS atseecar 20 m, #96-6 (R. R. Snelling); Hong Kong, in bank of mixed orchard (R. Winney); Tai- po (G. P. Tung); Shatin (Silvestri); Guang- dong Prov., 60 km W of Guangzhou, Ding- Hu Mts. (Boucek); Back Liang (S. F. Light); Fujian Prov., Jiangle (Minsheng Wang); Yenping (S. Ling); Zhejiang Prov., Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Mokanshan (N. Gist Gee); Shanghai (Sil- vestri); Soochow (N. Gist Gee); Nanking (G. P. Tung); Hubei Prov., Xiangfan (Min- sheng Wang); Guizhou, Leishan (Min- sheng Wang); Guizhou, Jiangkou (Min- sheng Wang); Szechwan Prov., Hsinching A F (W. L. Brown); Szechuen, near Mt. Ormel (D. C. Graham); Szechuen, Suifu (D. C. Graham); Szechuen, Jang Chen Pu near Mt. Omei (D. C. Graham); Taiwan, Suisha (R. Takahashi); Taiwan, Kosempo (H. Sauter); Taiwan, Bukai (L. Gressitt); Taiwan, Taihoku (J. Sonan, T. Shiraki); Tai- wan, Hassenzan (?); Taiwan, Nantou, Lan Wa Chu (D. G. Furth); Taiwan, Nantou, Sung Kang (D. G. Furth); Taiwan, Nantou, Hueisun (D. G. Furth); Taiwan, Nantou, Wushe (D. G. Furth); Taiwan, Taipei, Au- kung (D. G. Furth). Japan: Okinawa, Ku- nigami (Yonaha-dake), under chips (F. G. Werner); Ryukyu, Okinawa (S. M. Fuller- ton); Amami-Oshima, Loochoo Is. (R. Tak- ahashi); Kagoshima (Silvestri); Kumamotu (Silvestri); Kyushu, Amakusa, Tomioka (S. Murakami); Kyushu Is., Klyamacho, Kak- inohara (D. G. Furth); Kyushu Is., Mt. Hi- kosan, Soeda Notoge Pass (D. G. Furth); Kyushu Is., Mt. Hikosan, Biol. Sta. (D. G. Furth); Fukuoka Pref.,[zuka-machi, Joro- gahara (D. G. Furth); Shikoku Is., Toku- shima, Kawamata (M. Azuma); Hyogo, Mt. Rokko (M. Azuma); Hyogo, Takarazuka, Namaze (M. Tanaka); 8 mi N. of Kyoto (P. Hammond); Idzu (S. Akiyama); Kyoto (Sil- vestri); Mt. Maya (Silvestri); Michino-o | (Silvestri); Nara (Silvestri); Kuwana (?); Honshu, Toyama Pref., Toyama city (D. G. | Furth and K. Suzuki); Nagano Pref., Mat-_ (D. Gs Furth); Kamakura (H. Nagase); Kanagawa | Pref., Kamakura (H. Nagase); Kanagawa | Pref., Odawara (M. Kubota); Tokyo (L. | Gressitt); Tokyo Pref., Hachioji, Minami- | Asakawa (D. G. Furth); Yokohama (?); Chiba Pref., Ichikawa, Konodai (D. G. | Furth); Bonin Is., Chichi-jima (H. Ikeda); | Bonin Is., Chichi-jima, Omura, Camp | sumoto, Shimauchi-Shimoda beach (F. M. Snyder); Bonin Is., Chichi- jima, Yatsuse R., Gen.’s beach (F. M. Sny- | der); Bonin Is., Chichijima, Miyanchama, REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 415 yar RS Figures 94-97. Pristomyrmex rigidus sp. n. 94A: Worker head, full-face view; 94B: Showing two small denticles on the ventral clypeus; 95: Worker, lateral view; 96: Queen head, full-face view; 97: Queen, lateral view. Jack Wm’s beach (F. M. Snyder); Bonin Is., Chichijima, Sakai-ura, Bull beach (F. M. Snyder). I have also examined 11 workers col- lected at the two entry ports of the United States (USNM): nine of them, by Harley and Albrecht, on November 20, 1928, from Philadelphia, Pennsylvania, in lily bulbs imported from Japan; the other two specimens, by J. F. Byrnes, on September 25, 1967, from Anchorage, Alaska, on Ger- beria sp. imported from Japan. It appears, however, that P. punctatus has not yet be- come established in the United States (Cover, personal communication). In addition, this species has also been reported from North Korea and South Ko- rea (Collingwood, 1976, 1981; Kim and Kim, 1982: Kim and Kim, 1983). Pristomyrmex rigidus sp. n. Figures 94-97 Diagnosis (Worker). Antennal fossae reaching the anterior clypeal margin; eyes with six to eight ommatidia in the longest row; pronotum unarmed; dorsal surfaces 416 of head and alitrunk covered fully with well-developed coarse rugoreticulum; dor- sum of petiole node with a pair of hairs. Holotype Worker (BMNH). TL. 3.40, HL 0.98, HW 0.94, CI 96, SL 0.91, SI 97, EL 0.17, PW 0.64, AL 0.86. Paratypes: 11 workers (MHNG, BMNH, MCZC). Worker. TL 2.73-3.44, HL 0.75—0.98, HW 0.74—0.94, CI 93-102, SL 0.70—0.94, SI 91-103, EL 0.14—0.17, PW 0.50_-0.64, AL 0.70—0.86, PPW 0.24—0.30, PPL 0.16— O20VPEIMIA4=67 (r= 32): Mandibles with a few coarse longitudi- nal rugae that usually do not reach to the vicinity of the masticatory margin. Denti- tion of the masticatory margin of mandi- ble: the strongest apical tooth + the sec- ond strongest preapical + a long diastema + a truncated basal tooth (or two small denticles). A strongly developed tooth or a broad, subtriangular short prominence present about midway on the basal margin of mandible. Clypeus shield shaped, more or less depressed, with a median longitu- dinal carina that usually does not reach to the anterior clypeal margin; sometimes a few additional weak rugae are present on the clypeus. Anterior clypeal margin usu- ally with a median denticle and three oth- ers on each side, but sometimes with a lat- eral denticle indistinct or two lateral small denticles fused into a larger one. Lateral portions of clypeus reduced to margins, and antennal fossae reaching the anterior clypeal margin. Ventral surface of clypeus usually with two minute toothlike promi- nences (Fig. 94B), but sometimes the prominences are very weak. Palp formula 5,3. Frontal carinae extending to the level of the posterior margins of eyes. Antennal scrobes indistinct. Frontal lobes absent, so that the antennal articulations are entirely exposed. Antennal scapes, when lying on the dorsal head, surpassing the occipital margin of head by one-sixth to one-fifth of es length. Eyes containing six to eight Onninatidia in the longest row. Occipital margin rather straight or slightly concave. Dorsal surface of alerank serncan lia con- vex. Pronotum unarmed. Propodeal spines Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 robust, acute, and long, much longer than the distance between their bases. Meta- pleural lobes triangular and acute. Pedicel segments in profile as in Figure 95. In pro- file, anterior face of the petiole node, to- gether with the dorsal surface of petiole peduncle, forming a long declivity that reaches the top of the node. Petiole node in profile with an approximately right-an- gled apex. Dorsum of postpetiole in profile sometimes angulate but sometimes round- ed. In dorsal view, postpetiole transverse- rectangular, much broader than long and also broader than the petiole node. Dorsal surfaces of head and alitrunk as well as the sides of pronotum fully covered with coarse rugoreticulum. Sides of the rest of alitrunk with numerous irregular coarse rugae. Sides of petiole node and postpe- tiole usually with a few rims as illustrated in Figure 95, but sometimes some of the rims rather weak. Petiole and postpetiole in dorsal view, except for rims, very smooth and polished. Gaster unsculp- tured. Dorsal surfaces of head and alitrunk with numerous erect or suberect long hairs. A pair of similar hairs present on the dorsal petiole, and usually two pairs on the postpetiole as shown in Figure 95. Two or three pairs of long, forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with numerous erect or suberect hairs. First gastral tergite lacking erect or suberect hairs. Color uni- formly reddish-brown. Queen. TL 3.30-4.02, HL O;S5=15 OR HW 0.86—1.10, CI 100-102, SL 0.76—0.88, SI 80-91, EL 0.18—0.24, PW 0.66—0.86, AL 0.84—1.08, 0.22, PPI 140-164 (n = 6). Generally similar to worker, except for in addition, the sculp- | ture of mesoscutum is weaker and sparse. | caste differences: Male. Unknown. Comments. Pristomyrmex rigidus occurs in the Oriental region. It is closely related to P. punctatus. These two species consti- — tute a clade. The separation of the two | species is summarized under P. punctatus. — Characters separating P. rigidus from the — PPW 0.28-0.36, PPL 0.20- | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang P. divisus-P. pulcher clade are given under P. divisus. Discussion. The material that I have ex- amined may contain two species. Speci- mens from Thailand possess an anterodor- sally angulate postpetiole in profile, a strongly prominent tooth on the basal mar- gin of the mandible, and a truncated basal tooth on the masticatory margin of the mandible. Specimens from Indonesia, Ma- laysia, and Brunei have a rounded dorsal surface of the postpetiole, a weak tooth on the basal margin of the mandible, and two small basal denticles on the masticatory margin. But a single, variable species is maintained for the present because avail- able material is still limited. Further col- lecting and studying are needed. Holotype Worker. Thailand: 26. Kaeng Krachan NP., 19.xi.1985 (L6bl and Burcke hardt). Paratypes. 11 workers with same data as holotype. Additional Material Examined (ANIC, BMNH, MCZC, MHNG, NHMV). Thai- land: Khao Sabap NP. (L6bl and Burck- hardt). Brunei: Ulu Belalong, W. ridge, 500 ft (R. Levy). Malaysia: Sabah, Tawan Quoin Hill, 750 ft, rainforest, leaf mold, berlesate (R. W. Taylor); Sabah, Poring Hot Springs, 500 to 900 m (Burckhardt and Lébl); Sabah, Poring Hot Springs, Langanan Falls, 950 m (L6bl and Burck- hardt); Sabah, Batu Punggul Resort, pri- mary forest, sifting (P); Sarawak, 4th Di- vision, Gn. Mulu NP., limestone forest, 200 m, pitfall trap (H. Vallack); Sarawak, Gn Matang, 20 km W Kuching, 800 m, Prbimentane forest (L6bl and Bivekhar dt); Negri Sembilan, Pasoh For. Res., litter, primary forest (L. Ficken; M. Brendell, K. Jackson, and L. Ficken); Kuala, Lumpur (B. Bolton); Upper Gombak Val., near K. Lumpur, rainforest, berlesate, ca. 1,500 ft (R. W. Taylor); Kedah State, Gunong Jeral, 5.48N, 100.62E, 550 m, rainforest, berle- sate (R. W. Taylor and R. A. Barrett); Pa- hang, Batu Caves N Kuala, Lumpur (L6bl and Calame); Pahang Gombak, forest litter (L6bl and Calame). Indonesia: Sumatra, 417 Liwa, 5.04S, 104.03E, rainforest, litter (M. S. Harvey). Ecological Information. This species oc- curs in rainforest and has been taken in litter berlesates and pitfall traps. THE CRIBRARIUS GROUP Worker. Medium sized, with the follow- ing combination of characters. (1) Masticatory margin of mandibles with a long diastema between the preapi- cal and the basal tooth. (2) Palp formula 4,3 (Bolton, 1981). (3) Frontal carinae present. (4) Lateral portions of clypeus reduced to a margin, and the antennal fossae reach- ing the anterior clypeal margin. (5) Frontal lobes absent. (6) Lamella that encircles the base of antennal scape, entire. (7) Dorsum of alitrunk convex, not de- pressed; pro-mesonotal suture absent. (8) Pronotum armed with a pair of strong, acute, short spines. (9) Propodeal spines well developed and long; in dorsal view not forming a Stork (10) Petiole node thick in profile; its an- terodorsal, posterodorsal, and posteroven- tral corners showing right angles approxi- mately. This group contains one species, P. cri- brarius, from South Africa and Mozam- bique. Pristomyrmex cribrarius probably evolved from an Oriental ancestor of the punctatus group because P. cribrarius is similar, in the workers, to some Oriental species of the punctatus group in the fol- lowing characters: (1) a high palp formula; (2) well-developed, long propodeal spines; and (3) a few coarse longitudinal rugae present on the petiole and on the post- petiole, respectively. However, I erect the cribrarius group to accommodate this spe- cies instead of placing it in the punctatus group because it possesses a distinct pet- ole node in the workers and queens (i.e., in profile, the anterodorsal, posterodorsal, and posteroventral corners of the node 418 right-angled), which is not seen in the oth- er species groups. In addition, the workers of P. cribrarius possess a pair of robust pronotal spines that are absent in the punctatus group. Without a higher palp formula (4,3), P. cribrarius would fall into the quadridens group. The quadridens group now con- tains 25 species, but the palp formulae of 18 species examined are 2,3, 2,2, and 1,3. If P. cribrarius were placed in the quad- ridens group, the punctatus group (differ- ing from the quadridens group primarily by its high palp formula) would have no grounds to exist, as two species of the quadridens group, P. eduardi and P. orbi- ceps, also lack pronotal armaments. Pristomyrmex cribrarius is also some- what similar to the three members of the trispinosus group, from Mauritius. They all possess robust, acute, but short prono- tal spines; long, well-developed propodeal spines; and a very convex dorsum of the alitrunk in the workers. The similarity, however, is superficial. Pristomyrmex cri- brarius differs from the trispinosus group in the workers in several important char- acters: The frontal carinae are absent in the trispinosus group but extend to the level of the posterior margins of eyes in P. cribrarius; the promesonotal suture is present in the trispinosus group but absent in P. cribrarius; the propodeal spines in dorsal view are fused together at the base in the trispinosus group, as opposed to be separated at the base in P. cribrarius; palp formula is lower (1,2) in the trispinosus group but higher (4,3) in P. cribrarius; fo- veolate punctures on the dorsal surfaces of the head and the alitrunk are absent in the trispinosus group but dense in P. cribrar- ius; and numerous hairs are present on the first gastral tergite in the trispinosus group but not seen in P. cribrarius. Finally, P. cribrarius possesses (1) a long diastema on the masticatory margin ae mandible separating the preapical _ tooth from the basal tooth, (2) a higher palp for- mula (4,3), and (3) a pair of Tobuet spines on the pronotum in the workers, showing Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 it is not closely related to the levigatus, profundus, and umbripennis groups. Pristomyrmex cribrarius Arnold Figures 98-101 Pristomyrmex cribrarius Arnold, 1926: 281. Holotype queen, Mozambique: Amatongas Forest (G. Ar nold) (SAMC) [examined]. Diagnosis (Worker). See characters 8 and 10 under the group; additional char- acters including dorsal surfaces of head and alitrunk with foveolate-reticulate sculpture and postpetiole with a few coarse longitudinal rugae on each side. Worker. TL 3.10-3.54, HL 0.82-0.96, HW 0.89-1.02, CI 106—110, SL 0.72—0.80, SI7S—o2" eI, 0.19-0.22, PW 0.55-—0.64, AlE0FS0=0;905 BPVWV40:30=0S 65 RR 0:20= OV IAF ISO Go = 2). Mandibles with a few longitudinal basal rugae. Masticatory margin of mandible with an apical tooth + a preapical tooth + a long diastema + a broad and truncated basal tooth. Basal margin of mandible lack- ing a toothlike prominence or curved lobe. Clypeus with a strong median longitudinal carina. Anterior clypeal margin wath a me- dian truncated lobe and usually three to four denticles on each side. Ventral surface of clypeus with two weak toothlike prom- inences, or unarmed. Palp formula 4,3 (Bolton, 1981). Frontal carinae extending to the level of the posterior margins of — eyes. Antennal scrobes slightly concave. Antennal articulations entirely exposed. Antennal scapes, when lying on the dorsal : head, not reaching the occipital margin ofl head. Eyes with 9 to 10 ommatidia in the | longest row. Profile shape of alitrunk and | pedicel segments as in Figure 99. Dorsum of hla in profile, strongly convex. Pronotum with a pair of robust short | spines, ca. 0.06 to 0.08. Propodeal spines | well developed and long, ca. 0.25 to 0.30, much longer than the distance between their bases. Metapleural lobes small and | triangular. Petiole node high and thick in profile: its anterodorsal, posterodorsal, and | posteroventral corners showing right an- gles approximately. Postpetiole high ings REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 419 Figures 98-101. profile, curved dorsally; in dorsal view, transverse-rectangular and much broader than long. Dorsum of head, except for the scrobal areas, with foveolate-reticulate sculpture. Dorsum of alitrunk entirely sculptured with coarse longitudinal rugae and blunt foveolate punctures between ru- gae. Sides of alitrunk irregularly rugulose. Each side of petiole node and postpetiole with a few coarse longitudinal rugae. Gas- ter unsculptured. Dorsal surfaces of head and alitrunk with numerous erect or sub- erect hairs. Petiole and postpetiole each with a few pairs of hairs dorsally. First gas- tral tergite lacking hairs. Anterior clypeal margin with a row of forward-projecting hairs. Scapes and tibiae with erect or sub- Pristomyrmex cribrarius Arnold. 98A: Worker head, full-face view; 98B: Showing two minute denticles on the ventral clypeus; 99: Worker, lateral view; 100A: Queen head, full-face view; 100B: Showing two minute denticles on the ventral clypeus; 101: Queen, lateral view. erect short hairs. Color reddish-brown, but gaster darker. Queen. TL 3.84, HL 0.98, HW 1.04, Cl 1065 SE0:805 SI 7 EE O265 BW 0.325 ING WO sy IRA, O86, IPI OLR) IRE lay ~@ = |), Generally similar to worker, except for caste differences; in addition, pronotal ar- maments absent, eyes larger than in con- specific worker. Male. Unknown. Comments and Discussion. At first sight, P. cribrarius somewhat resembles two Australian species of the quadridens group, P. thoracicus and P. foveolatus: Their workers all possess a pair of short pronotal spines, a pair of long propodeal 420 spines, foveolate-reticulate sculpture on the dorsal head, and a transversely broad postpetiole in dorsal view. However, P. cri- brarius is rather different from P. thora- cicus and P. foveolatus. In the workers of P. cribrarius, palp formula is 4,3; the ven- tral surface of clypeus shows two minute teeth; the dorsum of alitrunk is very con- vex; the petiole and the postpetiole are sculptured with a few coarse longitudinal rugae; the eyes are larger, with 8 to 10 om- matidia in the longest row; and the petiole node, in profile, shows three right angles. But in the workers of P. thoracicus and P. foveolatus, palp formula is 2,3; the ventral surface of clypeus only has a single distinct tooth at the center; the dorsum of alitrunk is somewhat depressed; the petiole and the postpetiole are unsculptured; the eyes are smaller, with four to six ommatidia in the longest row; and the petiole node in pro- file lacks an acute posterodorsal angle. In the African Pristomyrmex fauna, P. cribrarius is easily recognized by the coarse longitudinal rugae on the dorsal al- itrunk anak on the des of the petiole and the postpetiole; its very convex dorsal ali- trunk; its well-developed, long propodeal spines; its distinct petiole node; and the numerous erect hairs on the alitrunk, pet- iole, and postpetiole. Distribution. South Africa, Mozambique (Bolton, 1981). Ecological Information. Specimens have been collected by W. L. and D. E. Brown in South Africa on sand, in coast vine for- est; by J. C. Faure, in South Africa (Zu- luland, St Lucia Lake), “by sifting the de- tritus and damp decaying leaves fond un- der bushes (Arnold, 1948)” ; and the ho- lotype by G. Amold on a tree trunk (Arnold, 1926). THE QUADRIDENS GROUP Worker. Usually medium- to large-sized ants (ble OM/3=1-46. SEW. O16S8=I"623 Ee 2.90-6.48) with the following combination of characters. (1) Masticatory margin of mandibles Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 with three to five teeth, which have one of the following three arrangements: a. the strongest apical + the second strongest preapical + a long diaste- ma + two small teeth of similar size (or one basal tooth, which is some- times formed by the fusion of the two small teeth) or b. the strongest apical + the second strongest preapical + a diastema + three small teeth of similar size or c. the strongest apical + the second strongest preapical + an intercalary small tooth + a very short diastema (or this diastema indistinct) + two small teeth of similar size (i.e., as shown in P. trachylissus). (2) Anterior clypeal margin with five or more denticles in most species, but several species having only three teeth or promi- nences. (3) Lateral portions of clypeus in front of antennal fossae reduced to a margin, and the antennal fossae reaching the an- terior clypeal margin. ae Palp formula 1 - a 11 species), o 2 (three species), or 2,3 (four specail a Frontal carinae nonre extending to the level of the posterior margins of eyes, with the exception in P. ert ythropt ygus, P. longispinus, P. trogor, and P. wilsoni. | (6) Frontal lobes indistinct or very weak. | (7) Antennal scrobes shallow or absent. | (8) Lamella, encircling the base of an- — tennal scape, entire. | (9) Dorsum of alitrunk lacking pro-me-_ sonotal suture. | (10) Pronotum armed with small teeth to well-developed spines, except in P. ed- | uardi and P. orbiceps. (11) Petiole node in profile usually. high, with the anterodorsal angle elevated above the posterodorsal, but sometimes | showing other forms. ; (12) Dorsum of head without sculpture, | with scattered foveolate punctures, or with — foveolate-reticulate sculpture or rugoreti- | culum. This group currently contains 25 spe-— REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang cies, accounting for almost half the genus. Of them, three occur in Africa, six in Aus- tralia and 16 in the Oriental region. In fact, like many Pristomyrmex, many spe- cies in this group have a restricted geo- graphic range. The quadridens group species are close- ly related to the cribrarius and punctatus groups but differ from them, most impor- tantly by their reduced palp formulae: 1,3, or 2,2, or 2,3 (4,3 in the cribrarius group and 4,3 or 5,3 in the punctatus group). The quadridens group differs from the trispinosus group because in the workers of the quadridens group, a promesonotal suture (or impression) is absent, the dor- sum of the propodeum in profile is not deeply concave, the propodeal spines in dorsal view do not form a fork, and regular striation is absent from the dorsal surfaces of the head and the alitrunk. The quadridens group differs from the levigatus, profundus, and umbripennis groups by possessing a distinct diastema on the masticatory margin of mandible in the workers and queens, with the excep- tion of P. trachylissus; but in the workers of P. trachylissus, five teeth are present on the masticatory margin, and the pronotum is armed with a pair of well-developed, long spines, which are not seen in the levigatus, profundus, and umbripennis groups. The evolution of mandibular dentition in the workers and queens is one of the reasons of the diversity of the quadridens group. The dental formula “apical tooth + a preapical + a long diastema + two small basal teeth of similar size”, possessed by many species of the group, is probably an “ancestral character”. It has given rise to three apomorphic dental formulae in the group: (1) an apical + a preapical + an intercalary small tooth + two small teeth of similar size, as in P. trachylissus; (2) an apical + a preapical + a diastema + three small teeth of similar size, as in P. eduardi, P. occultus, and P. quindentatus; and (3) an apical + a preapical + a long diastema + a basal tooth (which evolved through the 42] fusion of the two small teeth), as in P. er- ythropygus, P. quadridentatus, and P. wheeleri. The pronotal armaments in the workers of this group seem to show an evolutionary tendency to increase in size. For example, in P. brevispinosus, the pronotum is armed with teeth shorter than or about as long as propodeal armaments. In P. costatus, the pronotum is armed with a pair of short spines longer than the propodeal arma- ments, but much shorter than the distance between the bases of two pronotal spines. In P. bicolor, the pronotal spines are very long, much longer than the propodeal ar- maments and usually much longer than the distance between the bases of two pro- notal spines. In the workers and queens of the quad- ridens group, foveolate punctures on the dorsal head between the frontal carinae show continuous variation, from a com- plete absence to several punctures present to foveolate-reticulate sculpture (or dense assemblages similar to alveolate sculpture) or rugoreticulum. For example, P. collinus and P. flatus workers almost completely lack foveolate punctures on the dorsum of head between the frontal carinae. Pristo- myrmex quadridens workers have scat- tered foveolate punctures on the dorsum of head between the frontal carinae, but the spaces between foveolae are smooth. Pristomyrmex brevispinosus workers have rugoreticulum or foveolate-reticulate sculpture. Some populations, only with fo- veolate-reticulate sculpture behind the eyes, are considered intermediate forms and are grouped into P. brevispinosus. The males of eight species of this group are known (Figs. 261-266, 270-276). These males are more similar to those of the both trispinosus and levigatus groups than to the male of P. punctatus (punctatus group) in palp formula, propodeal arma- ments, and the shape of petiole or to those of the wmbripennis group in the size, pro- podeal armaments, and the shape and sculpture of petiole. Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 42. Pristomyrmex africanus Karavaiev Figures 102-108 Pristomyrmex africanus Karavaiev, 1931: 47. Holo- type worker, Kenya: Mabira (Dogiel) (UENC) [ex- amined]. Hylidris myersi Weber, 1941: 190. Syntype workers, Sudan: Equatoria, Aloma Plateau, Khor Aba, 3,700 ft, Lat. 3°47'N/Long. 30°37'E (N. A. Weber) [one syntype (MCZC) examined]. [Synonymy by Bolton, 1981}. Pieris myersi subsp. mbomu Weber, 1952: 19. Ho- lotype worker, Central African Republic: Ubangi- shari, Bas Mbomu, 5 mi W of Bangassau (N. A. Weber) (AMNH). [Synonymy by Bolton, 1981]. Hylidris myersi subsp. primus Weber, 1952: 19. Ho- lotype worker, Zaire: Stanleyville (N. A. Weber) (AMNH). [Synonymy by Bolton, 1981]. Hylidris myersi subsp. beni Weber, 1952: 20. Syntype workers, Zaire: 15 mi N of Beni (N. A. Weber) [one syntype (MCZC) examined]. [Synonymy by Bolton, 1981]. Diagnosis (Worker). Masticatory margin of Pemdible with a long diastema after the preapical tooth; ventral surface of clypeus with two toothlike prominences; eyes con- taining four to five ommatidia in the lon- gest row; dorsal surfaces of head between the frontal carinae and alitrunk with scat- tered foveolate punctures; petiole and postpetiole without hairs. Worker. TL 2.76—3.40, HL 0.74—0.90, HW 0.74—0.93, CI 98—104, SL 0.68—0.83, SI 86-95, EL 0.08—0.12, PW 0.48-0.58, AL 0.68—0.85, PPW 0.22-—0.27, PPL 0.18— Oe Rel BO=135) 7) = 27): Mandibles with a few longitudinal basal rugae. Masticatory margin of mandible with an apical tooth + a preapical tooth + a long diastema + a broad and truncated (or somewhat midconcave) basal tooth. Basal margin of mandible almost straight, lacking a foothlike prominence or curved lobe. Clypeus possessing or lacking a me- dian longitudinal carina. Anterior clypeal margin with a median denticle and two others on each side, but two lateral den- < is) ticles are usually fused into one promi- nence. Ventral surface of clypeus with two toothlike prominences. Palp formula 1,3. Frontal carinae extending to the level of the posterior margins of eyes. Antennal scrobes indistinct or very weak. Frontal lobes absent; thus, the antennal articula- tions are entirely exposed. Antennal scapes, when lying on the dorsal head, close to, or slightly surpassing, the occipi- tal margin of head. Eyes containing four to five Gruman in the longest row. Pro- mesonotum, in dorsal view, sometimes slightly concave. Profile shape of alitrunk and pedicel segments as in Figures 103— 104. Pronotum Fusually with a pair of teeth or short spines but rarely with a pair of tubercles. Propodeum with a pair of teeth or short spines. Metapleural lobes round- ed. Petiole node high in profile, higher than long, with a long anterior peduncle; its anterodorsal angle distinctly higher than the posterodorsal one that is usually rounded. Subpetiole with a narrow flange. Postpetiole in profile higher than long, rounded dorsally, in dorsal view broaden- ing from front to back and broader than long. Dorsum of head between the frontal carinae and the sides of head, with nu- merous scattered foveolate punctures; sometimes the punctures are shallow and sparse. Dorsum of alitrunk smooth, except for a few foveolate punctures shallow or conspicuous, present on each side of the mesonotum. Petiole, postpetiole, and gas- ter smooth and shining. Dorsum of ead between the frontal carinae, with some erect or suberect hairs. Promesonotum with a few pairs of hairs. Propodeum, dor- sal surfaces of petiole and postpetiole, and first gaster tergite lacking erect or suberect Thatins, A few pairs of forward. -projecting hairs present on the anterior clypeal mar- Figures 102-108. Pristomyrmex africanus Karavaiev. 102: Worker head, full-face view; 103: Worker, lateral view; 104: Showing that pronotal prominences are very weak in some specimens; 105A: Ergatoid queen, full-face view; 105B: Showing two small denticles on the ventral clypeus; 106: Ergatoid queen, lateral view; 107A: Queen head, full-face view; 107B: Showing two small denticles on the ventral clypeus; 108: Queen, lateral view. 424 gin. Scapes and tibiae with decumbent hairs. Color reddish-brown, but gaster darker. Ergatoid Queen. TL 3.70, HL 0.94, HW 1.00, CI 106, SL 0.85, SI 85, EL 0.16, PW 0.60, AL 0.92, PPW 0.30, PPL 0.20, PPI LEO Go = IL) General shape as in Figures 105—106. Similar to worker; color and pilosity as in worker. Sculpture, except for mesonotum where a few rugae present, as in worker. The head with three ocelli; eyes larger, with six to seven ommatidia in the longest row; pronotum armed with a pair of acute minute spines; propodeum with a pair of short spines; mesonotum more convex, and metanotal groove present. Flight sclerites and wings lacking, but a black speck is present on the each lateral margin of mesonotum. Queen. TL 3.40-3.82, HL 0.84—0.96, HW 0.86—1.02, CI 102—107, SL 0.74—0.88, SI (82-89) EL, 0202021, PW 07/0=0.74; AL 0.90—1.04, PPW 0.28-—0.30, PPL 0.20— O22 RPIMal3s2=14 35 — <>) General shape as in Figures 107—108, with normal caste differences from the conspecific worker. Eyes larger. Pronotum lacking teeth or spines but sometimes with a pair of blunt tubercles. Mesonotum with more hairs than in worker. Other charac- ters similar to worker. Male. Unknown. Comments. This African species is close- ly related to the Asian P. quadridens. Both worker and queen of P. africanus differ from those of P. quadridens in having two teeth on the ventral surface of clypeus and lacking erect or suberect hairs on both petiole and postpetiole. The separation of P. africanus, together with P. trogor, from the other two Oriental species, P. flatus and P. collinus, is provided under P. flatus. The workers of P. africanus and P. trogor differ from the workers of three Australian species, P. wheeleri, P. erythropygus, and P. quadridentatus as follows: P. africanus and P. trogor Dorsum and sides of both petiole and postpetiole lacking erect or suberect hairs Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Metapleural lobes rounded Dorsal alitrunk lacking a transverse ridge at the position of metanotal groove P. wheeleri, P. erythropygus, and P. quadridentatus Dorsum and sides of both petiole and postpetiole with erect or suberect hairs Metapleural lobes elongate-triangular, with an apex Dorsal alitrunk with a transverse ridge at the approximate position of matan- otal groove In addition, in P. africanus and P. trogor the denticles of the anterior clypeal margin are smaller but larger and stronger in P. wheeleri, P. erythropygus, and P. quadri- dentatus. Pristomyrmex africanus and P. trogor possess two toothlike prominences on the ventral clypeus that are not seen in P. wheeleri and P. erythropygus. Pristo- myrmex erythropygus has several short ru- gae on the juncture between the prono- tum and the mesonotum and has erect or suberect hairs on the first gastral tergite that are absent in P. africanus and P. tro- gor. Pristomyrmex trogor lacks a longitu- dinal median carina on the clypeus, pos- sessed by P. wheeleri, P. erythropy P. quadridentatus. The workers of P. africanus can be sep- arated from those of the other four African Pristomyrmex species as follows: (1) P. af- ricanus possesses numerous foveolate o 5 punctures on the dorsal head between the | frontal carinae that are not seen in P. tro- | gor and P. orbiceps; (2) P. africanus has | eyes containing four to five ommatidia in | the longest row, as compared with eight or more usually present in P. orbiceps, P. fos- sulatus, and P. cribrarius; (3) P. africanus has two teeth on the ventral clypeus that are absent in P. orbiceps and P. fossulatus; © (4) P. africanus lacks coarse longitudinal © rugae and erect or suberect hairs on the petiole and the postpetiole that are pres- ent in P. cribrarius. us, and — REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang Distribution. Ghana, Cameroon, Gabon, Kenya, Zaire, and Angola (Bolton, 1981). Some more records, including some eco- logical information, added here (MCZC, ANIC): Zaire: Ituri F. Beni-Irumu (N. A. Weber). Sudan: Equatoria, Imatong Mts. (N. A. Weber). Angola: Falls R. Chicapa, Saurimo, 9.39°S, 20.24°E, gallery forest, berlesate (Luna de Carvalho); R. Kahingo, 7.39°S, 20.51°E, gallery forest, berlesate (Mwaoka); Dundo, R. Mussungue, gallery for., berlesate (Luna de Carvalho); Dundo, dry forest (Luna de Carvalho); Dundo, Carrisso Park, R. Luachimo, 7.22°S, 20.50°E, gallery forest, berlesate (Luna de Carvalho); R. Camudembele, gallery for- est, berlesate (Luna de Carvalho); Ghana: E.R., Mt. Atewa, rainforest, Berlesate (R. W. Taylor); E.R., Nkwanda For., near En- yiresi, rainforest, Berlesate (R. W. Taylor); Tafo, Eastern Reg., rainforest, Berlesate (R. W. Taylor). Ecological Information. Weber (1941: 192; 1952: 18-20) noted that “workers were in rainforest of a luxuriant type re- ferred to as gallery forest”; they were among the leaf and humus cover on the forest floor and were slow moving in habit; “when disturbed they became motionless, ‘feigning death’ momentarily”. Pristomyrmex bicolor Emery stat. n. Figures 109-112 Pristomyrmex trachylissa var. bicolor Emery, 1900: 678. Syntype workers, Sumatra: Si-Rambé, xii. 1890-iii. 1891 (E. Modigliani) (MCSN, NHMV, USNM) [examined]. Pristomyrmex taurus Stitz, 1925: 120. Holotype worker, Philippines: N. Palawan, Binaluan, xi.— xii.1913 (G. Boettcher) (MNHU) [examined]. Syn. n. Note: The unique holotype of P. taurus differs from the syntypes of P. bicolor. In the former, the pronotal spine length is 0.58, and the anterior half of the dorsum of the alitrunk is smooth and shining. In the latter, the pronotal spine length is 0.36 to 0.40, and sometimes only a rather nar- row smooth area is present between the bases of two pronotal spines. Pristomyr- 425 mex taurus, however, is here regarded as a junior synonym of P. bicolor because, af- ter examining all the specimens available, I find that the pronotal spine length is con- tinuously variable in the range of 0.36 to 0.66, and sculpture on the anterior half of the dorsal alitrunk also shows continuous variation. I cannot use any lines to separate this composite at the present. The size of the holotype of P. taurus is HL 1.10, HW OS. Cl 9S. SE 114) ST 06) E0220" PV. O74 AIS) PSE 02 4s Mihensizessor three syntype workers of P. bicolor are HL 1 20=1228) IW 12421325 Cil03—lO7s sie 1.38—1.46, SI 108-111, EL 0.24—0.26, PW 0.84—0.90, AL 1.40—1.44, PSL2 0.10-0.12. Diagnosis (Worker). Masticatory margin of mandible with four teeth and a long di- astema after the preapical tooth; pronotal spines exceptionally long (usually 0.40— 0.66), much longer than propodeal arma- ments (teeth or short spines); dorsum of head sculptured with coarse rugoreticu- lum. Worker. TL 4.58-6.14, HL 1.04—1.42, HW 1.08-—1.46, CI 98—109, SL 1.14—1.58, SI 106-120, EL 0.20—0.26, PW 0.65—0.96, AL 1.18—1.64, PPW 0.33—0.40, PPL 0.38— 0.48, PPI 77-93 (n = 83). Mandibles usually with several longitu- dinal rugae, varying from superficial to rather coarse. Masticatory margin of man- dible with four teeth arranged as: two ad- jacent strong apical teeth + a long diaste- ma + two small basal teeth that are rough- ly the same size. Basal margin of mandible lacking a distinctly curved lobe or tooth. Clypeus with a strong median longitudinal carina. Anterior clypeal margin sometimes with seven denticles: a median denticle and three others on each side, but some- times one or a few are weak or indistinct, or two to three of the lateral denticles are fused into a larger one. Ventral center of clypeus usually with a transverse ruga of varying length, sometimes with a very weak prominence at middle and some- times without either of these characters. Palp formula 1,3. Frontal carinae strong, 426 CY SN RTS WZ S40 1) SE ~~ VO (cS Ne AO Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 SN Figures 109-112. Pristomyrmex bicolor Emery. 109A: Worker head, full-face view; 109B: Showing a weak transverse ruga on the ventral clypeus; 110: Worker, lateral view; 111A: Queen head, full-face view; 111B: Showing a transverse ruga on the ventral clypeus; 112: Queen, lateral view. extending to the level of the posterior mar- gins of eyes. Slightly concave scrobal areas present. Frontal lobes absent; thus, the an- tennal articulations are entirely exposed. Antennal scapes, when lying on the dorsal head, usually surpassing the occipital mar- gin by one-third to one-fourth of their length. Eyes usually containing 10 to 12 ommatidia in the longest row. Profile shape of alitrunk and pedicel segments as in Figure 110. Pronotal spines well devel- oped and exceptionally long, varying in length, usually 0.40 to 0.66, but in a few specimens they are 0.37 to 0.39. Propo- deum usually with a pair of acute short spines, which are 0.12 to 0.20, much more slender and much shorter than the pro- notal ones. Propodeal armaments occa- sionally reduced to a pair of teeth. Both pronotal and propodeal spines upward — pointed. Metapleural lobes subtriangular. In profile, petiole node high, with a long | anterior peduncle; its anterodorsal angle | elevated above the posterodorsal. In dorsal | view, crest of petiole node rounded. Post- ! petiole in profile convex dorsally, in dorsal | view longer than broad and broadening | from front to back. Dorsum of head with | well-developed coarse rugoreticulum. | Dorsal surface of alitrunk variably sculp- — tured: At one extreme, in a series from | North Borneo (SE, Forest Camp, 9.8 km | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 42 SW of Tenom), the dorsum of alitrunk is entirely rugoreticulate. In a few other se- ries, the rugoreticulum is absent between the bases On two pronotal spines, and some weak rugae or a smooth (broad or narrow) area are present there. At the other ex- treme, the sculpture is completely absent, and the area is smooth and shining on the anterior half of the dorsum, but the rest of the dorsal alitrunk is coarsely rugulose. Sides of pronotum with a rugoreticulum or many large coarse foveolate punctures. Sides of the rest of alitrunk irregularly coarsely rugulose. Dorsal surfaces of peti- ole node Pal postpetiole smooth and shin- ing, but a longitudinal ruga present on Gack side of petiole. Gaster unsculptured, smooth, and shining. Dorsal surfaces of head and alitrunk Sei numerous erect or suberect hairs. Petiole node with a few (usually two to three) pairs of hairs dor- sally. Dorsum of postpetiole with at least a pair of hairs. First gastral tergite lacking erect or suberect Hears: A row ae forward: projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect or suberect hairs. Color usually reddish-brown but sometimes black- brown. Queen. TL 6.54, 6.56; HL 1.34, 1.44; HW 1.38, 1.50; CI 103, 104; SL 1.48, 1.51; SI 99, 109; EL 0.32, 0.34; PW 1.20, 1.24: MG ei 2S 2 RP (OrA1e 0:43. PEEO0:438) 0.48; PPI 85, 90 (n = 2). General shape as in Figures 111-112, with normal caste differences from the conspecific worker; pronotum unarmed. Other characters similar to worker. Male. Unknown. Comments and Discussion. Pristomyr- mex bicolor occurs in the Oriental region. It possesses a pair of exceptionally long pronotal spines, which implies that this species may have evolved from the ances- tor of P. costatus. Only slight differences separate the workers of P. bicolor and of P. costatus, as follows: P. bicolor Pronotal spines well developed and ex- ceptionally long, usually 0.40 to 0.66 1 (rarely 0.37—0.39), and usually longer than the distance between their bases Dorsal surfaces of petiole node and postpetiole usually with one to three pairs of hairs Area between the bases of the pronotal spines smooth or sculptured; if sculp- tured, pronotal spine length is over 0.40 Larger species (HW usually 1.20—1.46, rarely 1.04—1.19; HL usually 1F20= EAA rarely 1.08—1.19) P. costatus Pronotal spines moderately long, usually 0.18 to 0.27 (rarely 0.32), and shorter than the distance between their bases Dorsal surfaces of petiole node and postpetiole with five or more pairs of hairs Entire dorsum of alitrunk with devel- oped rugoreticulum Smaller species (HW 0.90-1.16, HL 0.91—1.16) Another alternative is that P. bicolor may be derived from a P. curvulus—like an- cestor. Characters separating P. bicolor from P. curvulus are provided under P. curvulus. Pristomyrmex bicolor is also very similar in appearance to P. trachylissus. The two species are all from the Oriental region. The separation of the two species is sum- marized under P. trachylissus. It must be pointed out that P. bicolor is a highly variable species. I cannot separate any more sibling species from this mass at the present time. Further collecting and study will help clarify the situation. Material Examined (ANIC, BMHH, BMNH, LACM, MCZC, NHMV). Indo- nesia: W. Java, 9 km W Djasinga, Dungus Iwul, lowland rainforest (W. L. Brown): Java, Bali I, Tjanoi Kuning (J. Winkler); Borneo: Kaknntan Timur, ITCI Timber Camp, via Balikpapan, on fallen trees (N. Johnson); SE Borneo, 17 to 46 km W Ba- tulitjin, rainforest (W. L. Brown). Malaysia: North Borneo (SE), Forest Camp, 9.8 km SW of Tenom (Y. Hirashima); Sabah, Batu 428 Punggul Resort, primary forest, sifting (?); Sarawak, Genting Highlands (B. Bolton); Sarawak, 4th Div., G. Mulu Nat. Pk., RGS Expd., Long pala, lowland rainforest, on fallen tree and on rotten log (B. Bolton; P. M. Hammond and J. E. Marshall); Sara- wak, Mt. Penrissen, 4,500 ft (E. Mjdberg); Sarawak, Mt. Poi (E. Mjéberg); Malaya, Sq. Patani (G. H. Lowe); Selangor, Ulu Gombak For. Reserve, hill forest, Tree lookout area, ca. 450 m (R. Crozier); Se- langor, Genting Highlands, below Sri Lay- an, 900 m, hill forest (W. L. Brown); Neg. Sembilan, Pasoh For. Res. (M. Brendell, K. Jackson, and S. Lewis). Philippines: Lu- zon, Lagunas, Mt. Banahaw above Kina- buhayan, 600 to 700 m (J. Kodada and B. Rigova).? : Tjibodas, 1,500 m (?). Ecological Information. This species oc- curs in rainforest and has been collected on fallen trees and rotten logs. Pristomyrmex brevispinosus Emery Figures 113-118, 261, 270 Pristomyrmex brevispinosus Emery, 1887: 451. Syn- type workers and male, Sumatra: Mt. Singalang, Luglio, 1878 (O. Beccari) [syntype workers (MCSN, NMMV, USNM) examined]. *Pristomyrmex yaeyamensis Yamane and Terayama, 1999: 17. Holotype worker, Japan: Okinawa Pref., Yaeyama Is., Iriomote-jima, 7.viii.1985 (K. Kino- mura) (MNHA). Syn. n. Diagnosis (Worker). Masticatory margin of mandible with at most four teeth; pro- notal armaments toothlike, about long as propodeal armaments that are a pair of tri- * Note: I have seen photographs of Pristomyrmex yaeyamensis from Japanese Ants Image Database (which were placed under the name Pristomyrmex brevispinosus sulcatus). | propose Prystomyrmex yae- yamensis as a junior synonym of P. brevispinosus for the following reason: Although P yaeyamensis pos- sesses an ergatoid queen caste (Yamane and Teraya- ma, 1999), this condition, at this moment, is not enough to separate P. yaeyamensis from P. brevispi- nosus because the type specimens of P. brevispinosus contain only workers and male. In other words, it is not known, at the present, whether P. brevispinosus possesses a normal queen caste, an ergatoid queen caste, or both. (In Pristomyrmex, some species, e.g., P. wheeleri and P. africanus, contain both castes.) Thus, further ecological investigation is needed. Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 angular teeth or short spines; dorsum of head, at least behind the level of eyes, with foveolate-reticulate sculpture or rugoreti- culum. Workers DIE 3100-426 5s Hn On 3— 18045 HW 0.68—1.04, CI 93-102, SL 0.64—1.06, SI 94-108, EL 0.14—0.19, PW 0.48-0.68, AL 0.76—1.10, PPW 0.21—0.27, PPL 0.18— 0.30, PPI 86-113 (n = 100). Mandibles generally rather smooth but sometimes with a few longitudinal rugae. Dentition of the masticatory margin of mandible arranged as: the strongest apical + the second strongest preapical + a long diastema + two small teeth of similar size (or a broad tooth with two points). Basal margin of mandible almost straight, lack- ing a distinctly curved lobe or tooth. Clyp- eus with a median longitudinal carina that is sometimes interrupted. Anterior clypeal margin with a median denticle and two to three others on each side. Ventral center of clypeus usually with an acute tooth, but sometimes this tooth somewhat low and broad. Palp formula 1,3. Frontal carinae strong, extending to the level of the pos- terior margins of eyes. Antennal scrobes indistinct, but in some specimens shallow scrobal areas present lateral to the frontal carinae. Frontal lobes very weak; thus, the antennal articulations are almost entirely exposed. Antennal scapes, when lying on the dorsal head, slightly surpassing the oc- cipital margin of head. Eye usually con- taining seven to eight ommatidia in the longest row. Profile of alitrunk and pedicel segments as in Figures 114 and 116. Pron- otum armed with a pair of toothlike ar- maments that vary in length, approximate- ly from 0.06 to 0.10. Propodeum with a pair of triangular teeth or short spines varying from 0.04 to 0.12. Metapleural lobe subtriangular or with a somewhat rounded apex. Shape of petiole varying: In some populations, the anterior face of the petiole node in profile is almost insepara- ble from the upper surface of the pedun- cle (Fig. 114), but in other populations, the anterior face of the petiole node is dis- tinct from the upper surface of the pedun- 429 REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang Figures 113-118. Pristomyrmex brevispinosus Emery. 113A: Syntype worker head, full-face view; 114: Syntype worker, lateral view; 115A: Non-type worker head, full-face view; 116: Non-type worker, lateral view, showing some variation in the shape of petiole node and in the length of pronotal armaments; 117A: Queen head, full-face view; 118: Queen, lateral view; 113B, 115B and 117B: A tooth present on the ventral center of clypeus of syntype worker, non-type worker, and queen, respectively. 430 cle, as shown in Figure 116. Anterodorsal angle of petiole node higher than the pos- terodorsal. Postpetiole in profile convex dorsally, in dorsal view broadening from front to back. Density and intensity of ce- phalic sculpture variable: The dorsum of head is covered fully with coarse forveo- late-reticulate sculpture in type series, but with rugoreticulum in the some other se- ries and sometimes with foveolate-reticu- late sculpture only behind the eyes. Dor- sum of the alitrunk showing similar sculp- tural variation but usually possessing a few longitudinal coarse carinae. Petiole, post- petiole, and gaster smooth and shining. Dorsal surfaces of head and alitrunk with numerous erect or suberect hairs. Two pairs of hairs usually present bilaterally on the dorsum of petiole node. Usually, a pair, but sometimes two to three pairs, of hairs on the dorsum of postpetiole. First gastral tergite lacking erect or suberect hairs. A few pairs of forward-projecting hairs pre- sent near the anterior clypeal margin. Scapes and tibiae with some erect to sub- erect short hairs. Color reddish-brown to blackish-brown. Queen. TL 3.42-4.81, HL 0.82-1.09, HW 0.79-1.10, CI 93-106, SL 0.76—1.06, S93 S018 02020272 PWs0:66—0:39! AL 0.94—1.30, PPW 0.24—0.32, PPL 0.24— OBS), IPP GOI (py, = IS). General shape as in Figures 117-118, with normal caste differences from the conspecific worker, pronotum unarmed; other characters similar to worker. Ergatoid queen. This caste was reported (Onoyama, 1976; see also Yamane and Ter- ayama, 1999). Male (Figs. 261, 270). Two male speci- mens, together ath 31 workers and six queens, Sonnet. a series, collected in In- donesia (SE Celebes, 1-2 km E of Wolasi, 42 km S. Kendari, ca. 350 m), by W. L. Brown, and each of the two males was originally mounted, respectively, with a worker on the same pin: TL 3.10, 3.36; HL 0.60, 0.62; HW 0.61, 0.62; CI 102, 105; SL 0.21, 0.22; SI 35, 39; HWE 0.77, 0.80; EL OFS 1 OL32- RW OMS OFS 4 AE ae O Ge alaiG: Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 IRIE Ope, OpAalle IRL, O22), Owe IP }l Gls) 95 (n = 2). Head, including the eyes, distinctly broader than long. Clypeus convex, with- out a median longitudinal carina. Palp for- mula 1,3. Frontal carinae weak and short, just reaching the level of the posterior margins of antennal insertions. Maximum diameter of the median ocellus 0.10 to 0.11. On the mesonotum, notauli pro- nounced, forming a Y shape; parapsidal furrows indistinct. Scuto-scutellar sulcus with 12 to 13 narrow longitudinal ridges. Middle and hind tibiae without any spurs. Propodeum slightly tuberculate, lacking teeth or spines. Metapleural lobe with a somewhat rounded apex. Petiole node in profile low and rounded dorsally, with a fairly long anterior peduncle. Postpetiole in pr ofile slow and rounded dorsally and in dorsal view slightly longer than broad. Dorsum of head smooth and shining. Ali- trunk smooth and shining, except for those marked sutures. Petiole, postpetiole, and gaster smooth and shining. All dorsal sur- faces with abundant cieet or suberect hairs. Scapes and tibiae with numerous erect or suberect hairs. Body reddish- brown; funicular segments of antennae light-yellow and wings slightly infuscated. In general, the male of P. brevispinosus is extremely similar to the males of both P. quadridens and P. sulcatus. Comments and Discussion. Pristomyr- mex brevispinosus is the most widely dis- tributed species in the quadridens group. So far, it has been found in Sumatra, Cel- ebes, Malaya, Sarawak, Sabah, Thailand, the Philippines, Taiwan, and Japan. It is obvious that P. brevispinosus has evolved from a P. quadridens—like ances- tor. Pristomyrmex brevispinosus possesses densely assembled forveolate punctures (i.e., foveolate-reticulate sculpture) or ru- goreticulum on the cephalic dorsum in the workers and queens, on the dorsal alitrunk in the workers, and on the sides of the pronotum in the workers and queens. But P. quadridens has only scattered foveolate punctures in these areas. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang Some intermediate forms are present between P. brevispinosus and P. quadri- dens, but they may be easily assigned to either of the two species as follow Those populations possessing a few coarse lon- gitudinal carinae or some coarse rugae on the dorsum of the alitrunk or possessing foveolate-reticulate sculpture only behind the eyes should be assigned to P. brevispi- nosus, but those possessing only foveolate punctures and lacking coarse longitudinal carinae or rugae on the dorsal Falitrenk should be assigned to P. quadridens. The male of P. brevispinosus is indistin- guishable at present from that of P. quad- ridens; this strengthens the conclusion that these two species are closely related. An- other close relative of P. brevispinosus is P. sulcatus; their differentiation is dis- cussed under P. sulcatus. Characters sep- arating P. brevispinosus from the two Aus- tralian species, P. foveolatus and P. thora- cicus, are provided under P. foveolatus. Pristomyrmex brevispinosus may be split into two species in the future. One species would show that the anterior face of the petiole node, in profile, is insepa- rable from the upper surface of its anterior peduncle, and the other species would show that the anterior face of the petiole node, in profile, is distinct from the upper surface of the peduncle and that a concave shape is present between the two faces. However, at this moment, I cannot use this single line to separate the composite. Note: Lin and Wu (1998) elevated P. brevispinosus sulcatus var. formosae Forel to species rank. As an infrasubspecific tax- on, “formosae” is not an available name in the genus, according to the International Code of Zoological Nomenclature (also see Bolton, 1995: 365). If the specimens assigned to “formosae” represent a good species differing from P. brevispinosus, the correct pr ocean would be to describe it as new to science. I have, however, ex- amined three syntypes of “formosae”, and they fit comfortably with P. brevispinosus as defined in this revision. Material Examined (ANIC, BMHH, 431 BMNH, LACM, MCZC, MHNG, NHMYV, USNM). Indonesia: Sumatra, Pematang, Siantar (Mann); Si-Rambé (E. Modigliani); Sulawesi Tengah, near Morowali, Ranu River Area (M. J. D. Brendell); Sulawesi Utara, Dumoga-Bone N.P., lowland forest, 200 to 400 m, litter (?); Sulawesi, Dumo- ga-Bone N.P. (D. F. and A. K. Roche); S Celebes, Balampesoang Forest, 5 to 8 km NE Tanete, 400 m, degraded rainforest (W. L. Brown); SE @llebes. tor 2k of Wolasi, 42 km S Kendari, ca. 350 m, rainforest, rotten wood (W. L. Brown); N. Celebes, Mt. Tangkoko-Batuangus Res. 10 to 200 m, tropical evergreen forest, under bark log (W. L. Brown); N. Celebes, SW slope Mt. Klabat, 400 to 600 m, rainforest, rotten wood (W. L. Brown). Malaysia: Ma- laya, Genting, Highlands (B. Bolton): Se- langor, Ulu leona For. Reserve, Univ. Malaya Field Studies Center, ca. 260 m, rainforest (R. Crozier); Negri Sembilan, Sungei Menyala For. Res., near Port Dick- son, lowland rainforest (W. L. Brown and Tho Yow Pong); Perak, Sungei, Simei Falls, Cameron Hlds (T. Jaccoud and P. Marcuard); Trengganu, Kuala Buka near Trengganu, berlesate (T. Jaccoud and P. Marcuard); Sarawak, Semengoh For. Re- serve, 11 mi SW Kuching, rainforest, nest ex rotten log (R. W. Taylor); Sarawak, Gn Matang, 20 km E Kuching, 850 m, sub- montane forest (L6bl and Burckhardt); N Borneo (E. Mjoberg); N. Borneo, W. coast Residency, Ranau, 500 m (T. C. Maa); Sa- bah, Poring Hot Springs, 500 m (Burck- har dt and Lobl): Sabah, 7 km N Tambun- an, 700 m (L6bl and Burckhardt); Sabah, Batu Punggul Resort, Be forest, itt ing (?); S Sabah Crocker Range NP., Gg. eins Highland Res., 1,500 to "1.700 m (>): Neg. Sembilan Pasoh For. Res. (M. Bren- dell K. Jackson, and S. Levvis); Pahang, Genting, Highlands Awana, 1,150 m (Lobl and Calame); Pahang, Ringlet, ravine, 1,250 m (L6bl and Gulamey Pahang, Batu Gacs N Kuala, Lumpur (L6bl anid Cala- me). Philippines: P.R. Mindanao, 2 km N Malaybalay, 670 m, 8.09°N/125.05°E, re- forestation area (B. B. Lowery). Thailand: 432 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 119-122. Pristomyrmex collinus sp. n. 119: Worker head, full-face view; 120: Worker, lateral view; 121A: Queen head, full-face view; 121B: Showing a tooth on the ventral clypeus; 122: Queen, lateral view. Trang Prov.: Khao Chong Nature Educa- tion Center, lowland tropical rainforest, (07.35°N/99.46°E, misc stray foragers, sift- ed leaf in mixed forest; 07.55°N/99.58°E, misc stray foragers) (R. R. Snelling and Saowapa Sonthichari). Taiwan: Pilam (H. Sauter). Ecological Information. This species oc- curs in rainforest and has been collected on rotten logs. Pristomyrmex collinus sp. n. Figures 119-122 Diagnosis (Worker). Pronotum and pro- podeum each with a pair of short spines; dorsal surfaces of head and alitrunk un- sculptured, smooth, and highly polished; petiole node with one to two pairs of hairs, and with the anterior face of the node dis- tinct from the upper surface of peduncle; HW 0.77-0.94 and HL 0.82-0.94. Holotype Worker (MCZC). TL. 3.51, HL 0.88, HW 0.84, CI 95, SL 0.86, SI 102, PW 0.59, AL 0.91. Paratypes, 40 workers and nine queens (MCZC, BMNH, MHNG). Worker. TL 3.36—3.84, HL 0.82—0.94, HW 0.77-0.94, CI 93-103, SL 0.80—0.94, SI 98-110, EL 0.14—0.18, PW 0.54—0.62, AL 0.86-—1.00, PPW 0.25-0.26, PPL 0.25-— 0.28, PPI 89=100 (m= 40). Mandibles usually smooth and shining but sometimes with a few longitudinal ru- gae. Dentition of the masticatory margin of mandible: the strongest apical + the second strongest preapical + a long dia- REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang stema + two small teeth that are subequal in size. Basal margin of mandible some- what straight, lacking a distinct tooth. Clypeus depressed and smooth, usually unsculptured but very rarely with a longi- tudinal median carina. Anterior clypeal margin usually with a median denticle and two others on each side but sometimes with a lateral denticle indistinct or two lat- eral denticles fused into a larger one. Ven- tral surface of clypeus with a low, broad- based, central tooth. Palp formula 1,3. Frontal carinae extending to the level of the posterior margins of eyes. Antennal scrobes absent. Frontal lobes weak; thus, the antennal articulations are almost en- tirely exposed. Antennal scapes, when ly- ing on the dorsal head, slightly surpassing the occipital margin of head. Eyes mod- erate, usually containing six to seven om- matidia in the longest row. Pronotum with a pair of short but acute spines that are slightly variable in length. Propodeum armed with a pair of short spines that are about equal to or slightly longer than the pronotal ones. Metapleural lobes subtrian- gular. In both profile and dorsal view, the dorsum of alitrunk convex, that is, prono- tum plus mesonotum forming a convex dorsum. Petiole node in profile with a fair- ly long anterior peduncle, its anterodorsal angle higher than the posterodorsal. Post- petiole in profile rounded dorsally, in dor- sal view slightly longer than broad, or about as long as broad, but always slightly broadening from front to back. Dorsum of head, except for a few punctures border- ing the frontal carinae, smooth and highly polished. Dorsum of alitrunk unsculptured and highly polished. Very rarely, the dorsal surfaces of head and alitrunk with a few feeble punctures. Petiole, postpetiole, and gaster smooth and shining. A weak longi- tudinal ruga usually present on each side of the petiole but absent in few specimens. Dorsal surfaces of head and alitrunk with some sparse erect or suberect hairs. One or two pairs of hairs present on the dorsal surfaces of petiole node and postpetiole, respectively. First gastral tergite lacking 433 hairs. Three pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with erect to suberect short hairs. Color uniformly red- dish-brown. Queen. TL 4.04-4.38, HL 0.87-0.94, HW 0.85-0.96, CI 94—105, SL 0.84—0.94, SI 98-102, EL 0.22—0.26, PW 0.76-0.84, Ale, 0:92=1-268P PW. 0.270 S05 BEE O26= 0.29; PPI 100=111 (m = 8). Generally similar to worker, except for normal caste differences. In addition, pronotum unarmed; propodeal armaments toothlike, shorter than those of the con- specific worker. Male. Unknown. Comments. This species is known only from the Philippines so far. It has a num- ber of relatives. Two of them occur in Southeast Asia: P. flatus, also from the Philippines, and P. quadridens, from New Guinea, Indonesia, and Pohnpei. Of its more distant five relatives, two, P. african- us and P. trogor, occur in Africa; and the other three, P. quadridentatus, P. wheeleri, and P. erythropygus, are endemic to Aus- tralia. The following characters can be used to separate the workers of P. collinus from those of P. flatus: P. collinus Promesonotum in dorsal view showing a convex dorsum Anterior face of petiole node, in profile, distinct from the upper surface of its anterior peduncle Smaller species, with HW 0.77—0.94, HL 0.82-0.94, EL 0.14—0.18 P. flatus Promesonotum in dorsal view depressed or shallow-concave Anterior face of petiole node not dis- tinct from the upper surface of the peduncle Larger species, with HW 0.98—1.04, HL 0.94—-1.02, EL 0.22-0.24 A list of characters separating P. collinus and P. flatus from P. quadridens and from 434 TIPE fis 125 ai B Figures 123-126. Pristomyrmex costatus sp. n. 123A: Worker head, full-face view; 123B: Showing a transverse ruga on the | ventral clypeus; 124: Worker, lateral view; 125A: Queen head, full-face view; 125B: Ventral clypeus without a toothlike promi- | nence; 126: Queen, lateral view. two African and from three Australian spe- cies is provided under P. flatus. Holotype Worker. Philippines: Duma- guete (J. W. Chapman). Paratypes. 40 workers and nine queens with same data as holotype. Additional Material Examined (BMHH, USNM, NHMV, MCZC). Philippines: Du- maguete, Horns of Negros, 3,600 ft (J. W. Chapman); Dumaguete, Camp (J. W. Chapman); Los Banos (F. X. Williams): Luzon, Mt. Makiling (F. X. Williams; L. Quate and C. Yoshimoto); Luzon, Laguna, Mt. Makiling, 500 to 1,144 m (H. Zettel); Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Luzon, Laguna, Mt. Banahaw above Kina- | buhayan, 600 to 700 m (J. Kodada and B. | Rigova); Romblon Prov., Tablas, S. Agus- | tin, Dubduban, Busai Falls (H. Zettel); Pa- | nay Is., forest, 300 m (R. C. Mcqregor). | Ecological Information. This species oc- | cur in forest, according to the present re- — cords. Pristomyrmex costatus sp. n. Figures 123-126 Diagnosis (Worker). Ventral surface of — clypeus lacking a developed tooth, but | usually with a transverse ruga; pronotal REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang spines fairly long, ca. 1.5 to 2 times the length of propodeal armaments, but dis- tinctly shorter than the distance between the bases of two pronotal spines; dorsal surfaces of head and alitrunk sculptured with coarse rugoreticulum; petiole node lacking foveolate punctures; first gastral tergite lacking erect or suberect hairs. Holotype Worker (BMNH). TL 4.14, HL 0.99, HW 0.96, CI 97, SL 0.96, SI 100, BE O21 PW. O.6S:7 Al Tl2)) PPW-0:30: REM Os0O, Per 100: Paratypes, 3 workers (MCZC): TL 4.48-4.54, HL 1.04—1.08, ED O:98=1-02) (El 94-945 SI lOve. SI 108-110, EL 0.20—0.20, PW 0.68-0.72, Niels Awe ENO S0=Orsle ee EO 33— 0.34, PPI 91-94. Mandibles usually with several longitu- dinal rugae. Masticatory margin of man- dible een four teeth arranged as two ad- jacent strong apical teeth ie a long diaste- ma + two emia basal teeth of ernie size. Basal margin of mandible lacking a dis- tinctly caved lobe or tooth. Clypeus with a strong median longitudinal carina. An- terior clypeal margin ath a median den- ticle and two to three others on each side. Ventral surface of clypeus possessing or lacking a transverse ruga, never armed with a developed, acute tooth. Palp for- mula 1,3. Frontal carinae strong, extend- ing to the level of the posterior margins of eyes. Slightly concave scrobal areas pres- ent lateral to the frontal carinae. Frontal lobes weak so that the antennal articula- tions are almost entirely exposed. Antennal scapes, when lying on the dorsal head, slightly surpassing the occipital margin of head. Eyes containing 9 to 10 ommatidia in the longest row. Profile shape of alitrunk and pedicel segments as in Figure 124. Pronotum ar med with a pair of str ong and fairly long spines that are ca. 0.19 to 0. Die 1.5 to 2 enee the length of propodeal ar- maments, but distinctly shorter than the distance between the bases of two prono- tal spines. Propodeum with a pair of acute short spines that are ca. 0.10 to 0.16 and more slender than the pronotal ones. Me- tapleural lobes subtriangular. Petiole node 435 in profile, slightly higher than long, with a fairly long anterior ‘peduncle: is. antero- dorsal angle is on a higher level than the posterdorsal. Postpetiole in profile round- ed dorsally, in dorsal view broadening from front to back. Dorsum of head, except for the scrobal areas where there are only some transverse rugae, with well-devel- oped coarse rugoreticulum. Similar sculp- ture present on the sides of pronotum and the dorsum of alitrunk. Petiole with a coarse longitudinal ruga on each side, but dorsum es petiole mode unsculptured and smooth. Dorsum of postpetiole unsculp- tured and smooth. Gaster smooth and shining. Dorsal surfaces of head and ali- trunk with numerous erect or suberect long hairs. Sides and dorsum of petiole node and postpetiole with five or more pairs of hairs in the type specimens. First gastral tergite lacking erect or suberect hairs. Several pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect to suberect hairs. Color reddish-brown. Queen. Two queens, respectively, from N. Borneo and Sarawak, with the following measurements: TL 5.24, 5.26: HL 1. 12, 202 FW, Wil6: 122 Cl l02) L042 SIE es: 1.20: SI 98, 102: EL 0.28, 0.31: PW 1.00; NO AeA O2 SP PNWViOrsS 30:44 s Pils 0.40, 0.44, PPI 95, 100. General shape as in Figures 125-126, with normal caste differences from the worker, pronotum unarmed; other char- acters similar to worker. Male. Unknown. Comments and Discussion. Pristomyr- mex costatus is extremely similar in ap- pearance to the another Oriental species, P. sulcatus. The workers and queens of the two species can be separated as follows: In P. costatus, the ventral surface of the clyp- eus lacks a toothlike prominence, usually with a transverse ruga; in P. sulcatus, the ventral center of the clypeus has an acutely prominent tooth and no transverse ruga. The separation of P. costatus from P. modestus and from P. bicolor is summa- rized under the latter names. 436 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 @ oP. costatus * Ps SulleeEeus Figure 127. Distributions of Pristomyrmex costatus and Pristomyrmex sulcatus. Whether or not P. costatus and P. sul- Borneo, and the Philippines but P. sulcatus catus have a relationship of allopatric dis- in Pahang of Malaya, Thailand, Nepal, tribution needs further studies. According Burma, and China. Further, whether P. to the present records, as shown in Figure sulcatus, P. costatus, and P. brevispinosus 127, P. costatus occurs in Negri Sembilan evolved from the results of character dis- of Malaya, Singapore, Sabah, Sarawak, placement also needs further studies (P. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang costatus is more distant to sympatric IP, brevispinosus than to P. sulcatus). The following additional material shows some interesting variation: In nine work- ers, the dorsums of petiole node and post- petiole have five or more pairs of hairs, but the ventral surface of clypeus has only a short ruga. Four workers from Singapore have two pairs of hairs on the dorsum of petiole node and a pair on the dorsum of postpetiole. In a worker from Sabah, the anterodorsal angle of the petiole node in profile is not distinctly higher than the posterodorsal. In three workers, one from Sarawak and the other two from Sabah, the ventral center of clypeus lacks any transverse ruga but possesses a minute prominence instead of an acute tooth, and the dorsal surfaces of petiole and postpe- tiole have more than seven pairs of hairs. A single specimen, from the Philippines, shows reduced sculpture of the dorsal sur- faces of head and alitrunk. The measure- ments of the previously described speci- mens are TL 3.94-4.54, HL 0.91-1.18, HW 0.90-1.13, CI S9—103, SL 0.96—1.21, SI 100-112, EL 0.18—0.24, PW 0.60-0.76, AL 0.98—1.30 (n = 18). In addition, a series from Borneo ap- pears to be intermediate in size between P. costatus and P. bicolor by possessing the following worker measurements: TL 4.64— Se, IRE IOO SIZE JEW OR See Olt 931045 7S a4 22 SI 1038=113. EE 0:20-0:22, PW 0.70=0.78, AL 1.16—1.37, PSbLil 0:28-0:32, PSL2 0.10=0.116 (7 = 25). It is possible that the previously de- scribed material may comprise one or more sibling species. Further collecting and biological investigation are needed to resolve this possibility. Holotype Worker. Malaysia: Neg. Sem- bilan, Pasoh For. Res., xi.1994, (M. Bren- dell, K. Jackson, and S. Lewis). Paratypes. Three workers, N. Borneo (E. Mjéberg). Records of the Previously Examined Non-Type Material (ANIC, BMHH, BMNH, MCZC, MHNG, NHMV). Singa- pore: Nee Soon, Swamp forest, rainforest, 437 nest ex rotten log (R. W. Taylor); Bukit Ti- mah Nat. Res., degraded coastal hill forest, on granite (D. H. Murphy). SE Borneo: 17 to 46 km W Batulitjin, lowland rainfo- rest (W. L. Brown); Borneo: Pajan (E. Mjéberg). Sarawak: Gunong Matang 120 m (T. C. Maa); 4th Div., G. Mulu Nat. Pk., RGS Expd., Long pala, lowland rainforest, on rotten log and in leaf litter (B. Bolton); Semengoh NSG, 30 km S Kuching (H. Zettel). Sabah: Batu Punggul Resort pri- mary forest, sifting (P); 43 mi, labuk Rd. ex Sandakan (Lungmanis) (R. W. Taylor); 7 km N Tambunan, 700 m (L6bl and Burchhardt); N. Borneo, Tutu River (E. Mjéberg); N. Borneo, (SE) Forest Camp, 9.8 km SW of Tenom (K. J. Kuncheria). Philippines: Mindanao, Davao Province, Mt. McKinley, E. slope, 3,300 ft, under bark (F. G. Werner). Note: An unusual worker (BMNH), col- lected from Sabah (K. K.-Tambunan, Crocker Range, 1,600 m), having a smooth patch present between the bases of two shorter pronotal spines, is tentatively placed under P. costatus; the size of the specimen is TL 3.84, HL 0.96, HW 0.92, CI 96, SL 0.96, SI 104, EL 0.18, PW 0.64, AL 0.96, PSLI1 ca. 0.16, PSL2 ca. 0.08. Ecological Information. See the section “Records of the Previously Examined Non-Type Material”. Pristomyrmex curvulus sp. n. Figures 128-131 Diagnosis (Worker). Pronotal spines ex- ceptionally long, about 0.38 to 0.44; pro- podeal spine length ca. 0.12 to 0.16; an- terior face of petiole node distinctly sepa- rable from the upper surface of peduncle; dorsal surfaces of head and _ alitrunk smooth and shining. Holotype Worker (MCZC). TL5.24, HL eis; AW 1.20; Cl 102, Sk. 1.36, SP Hs: EL 0.22, PW 0.79, AL 1.26. Paratypes, 34 workers and one queen (MCZC, BMNH, LACM, MHNG). Worker. TL 4.62-5.30, HL 1.08—1.26, HW, 1_08=1-25,, Cl 97-105; SL, 122-4 SI 106-117, EL 0.20—0.26, PW 0.74—0.82, 438 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 128-131. Pristomyrmex curvulus sp. n. 128A: Worker head, full-face view; 128B: Showing a transverse ruga on the ventral clypeus; 129: Worker, lateral view; 130A: Queen head, full-face view; 130B: Showing a transverse ruga on the ventral clypeus; 131: Queen, lateral view. Mlle tO Me PNVOS0=0.34 Pel, Oloa— 0.40, PPI 78-94 (n = 20). Mandibles generally smooth and _ shin- ing, with a few basal longitudinal rugae. Masticatory margin of mandible with four teeth: an apical + a preapical + a long diastema + two small denticles that are roughly the same size. Basal margin of mandible lacking a toothlike prominence. Clypeus shining, with a median longitudi- nal carina; sometimes a few additional su- perficial rugae present. Ventral surface of clypeus usually with a long transverse ruga. Anterior clypeal margin usually with seven denticles (a median one and three others on each side), but in some specimens, one to two denticles weak or rudimentary. Palp formula 1,3. Frontal carinae just extending to the level of the posterior margins of eyes. Slightly concave scrobal areas pres- ent lateral to the frontal carinae. Frontal lobes weak so that the antennal articula- tions are almost entirely exposed. Antennal scapes long, surpassing the occipital mar- gin by one-fourth to one-third of their length. Eyes usually containing over 10 ommatidia in the longest row. Profile shape of alitrunk and pedicel segments as in Figure 129. Pronotum armed with a pair of exceptionally long spines that are about 0.38 to 0.44 and longer than the dis- tance between their bases. Propodeum REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang with a pair of acute short spines that are ca. 0.12 to 0.16, about as long as the dis- tance between their bases, and shorter than 0.5 times pronotal spine length. Both pronotal and propodeal spines directed upward. Metapleural lobes subtriangular. Petiole in profile nodiform, with a long an- terior peduncle; the anterior face of the node distinctly separable from the upper surface of its anterior peduncle, and its dorsum sloping somewhat downward pos- teriorly. Postpetiole in profile rounded dorsally, in dorsal view distinctly longer than broad and broadening from front to back. Dorsum of head generally smooth and shining, but gena with a few foveolate punctures, and frontal area usually with a few weak short rugae. Dorsal alitrunk, petiole, and postpetiole unsculptured, smooth, and shining. Gaster unsculptured. Dorsal surfaces of head and alitrunk with numerous erect or suberect hairs. Petiole node and postpetiole each with some hairs as shown in Figure 129. Antennal scapes and tibiae with numerous erect or sub- erect hairs. First gastral tergite without erect or suberect ais. A re pairs of for- ward-projecting hairs present near the an- terior clypeal margin. Color reddish brown but light yellow in a few specimens. Queen. TL 6.16, HL 1.18, HW 1.28, CI IOS, Sls ILS, SIMO IaIly O28, IPWY ILL Mle O2) PW 0:40> PPE O44. PPT SING = Il). General shape as in Figures 130-131, with normal caste differences from con- specific worker; pronotum unarmed; peti- ole with a lateral longitudinal ruga on each side. Other characters similar to worker. Male. Unknown. Comments and Discussion. This species is closely related to P. longispinus, also from Dumaguete, Philippines, but the workers of Hhieee two species can be sep- arated by the following characters: P. curvulus Anterior face of petiole node, in profile, distinct from the upper surface of pe- duncle 439 Clypeus with a median longitudinal ca- rina Frontal carinae extending to the poste- rior margins of eyes Propodeal spines shorter, about as long as the distance between their bases. Anterior clypeal margin usually with seven small denteles P. longispinus Anterior face of petiole node, in profile, not distinct from the upper surface of peduncle Clypeus lacking a median longitudinal carina Frontal carinae not extending to the posterior margins of eyes Propodeal spines longer, about two to three times the distance between their bases in length Anterior clypeal margin usually with three to five denticles Pristomyrmex curvulus may also have a close relationship with P. bicolor. The workers and queens of the two species can be separated as follows: In P. curvulus, the dorsal surfaces of the head and the ali- trunk are smooth and shining, with only a few shallow foveolate punctures present on the genae and a few short rugae on the frontal area, whereas in P. bicolor, the dor- sum of the head is entirely covered with coarse rugoreticulum, and the dorsum of the alitrunk is also strongly sculptured with coarse rugae. It is possible that P. curvulus is derived from a P. collinus—like ancestor. The work- er of P. curvulus, apart from the excep- tionally long pronotal spines, a transverse ruga on the ventral surface of the clypeus, and its larger size, is similar to that of P. collinus. The queen of P. curvulus is also similar to that of P. collinus, but in the for- mer it is larger, and the ventral surface of the clypeus ahas a transverse ruga, not a toothlike prominence as in P. cole Holotype Worker. Philippines: Duma- guete (J. W. Chapman). Paratypes. Philippines: two workers, Dumaguete, 7.vi.1942 (J. W. Chapman); 440 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 ZA EST S Figures 132-133. Pristomyrmex eduardi Forel. 132A: Worker head, full-face view; 132B: Showing a toothlike prominence on the ventral clypeus; 133: Worker, lateral view. two workers, Dumaguete, 14.vi.1942 (J. W. Chapman); 12 workers, Dumaguete, 10.v.1947 (J. W. Chapman); two workers, Dumaguete, 4.v.1948 (J. W. Chapman); one worker, Dumaguete, 1949 (J. W. Chapman); two workers and one queen, Dumaguete, 1950 (J. W. Chapman); 11 workers, Dumaguete, Horns of Negros, 3,600 ft (J. W. Chapman); two workers, Dumaguete, Horns of Negros, 3,600 ft (Domingo Empeso). Additional Material Examined (MCZC). Some specimens, also collected in Duma- guete, Philippines, by J. W. Chapman, are not included in the type series because they are badly mounted or damaged. Ecological Information. Unknown. Pristomyrmex eduardi Forel Figures 132-133 232 Pristomyrmex eduardi Forel, 1914: 232. Holotype worker, Sumatra Oriental, Bah Boelian (M. v. But- tel) (MHNG) [examined]. Diagnosis (Worker). Masticatory margin of mandible with five teeth; pronotum un- armed; eyes with three to four ommatidia in the largest row. Worker. TL 2.9, HL 0.77, HW 0.74, CI 96, SL 0.68, SI 92, EL 0.07, PW 0.50, AL 0.78) (= 1): Mandibles smooth and shining, except for a few longitudinal rugae. Masticatory margin of mandible with five teeth ar- ranged as the strongest apical + the sec- ond strongest preapical + a diastema + three small denticles of similar size; the length of diastema is about equal to the distance covered by three small denticles. Basal margin of mandible lacking a tooth- like prominence. Clypeus depressed, with a short median carina that does not reach the anterior clypeal margin but runs through the frontal area. Anterior clypeal margin with five toothlike prominences; the median three somewhat truncated. Ventral center of clypeus with a prominent tooth. Frontal carinae strong, extending to the level of the posterior margins of eyes and forming the dorsal margins of the shallow scrobes. Frontal lobes weak. Eye small, with three to four ommatidia in the longest row. Occipital margin in full-face view feebly concave. Profile of alitrunk and pedicel segments as in Figure 133. Pron- otum unarmed, lacking a pair of teeth or spines. Propodeum armed with a pair of acute short spines. Metapleural lobes prominent and rounded. Petiole in profile view with a fairly long anterior peduncle; the anterodorsal angle of the node high, and its dorsum sloping downward poste- riorly. Postpetiole in profile with a round- ed dorsum. Dorsum of head with coarse rugoreticulum, except for a smooth, me- REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 44] 152°25'E, in rotten logs, xi.1957 (Darlingtons) dian longitudinal strip. Dorsum of ali- trunk, as well as two sides of pronotum, with developed rugoreticulum. Petiole and postpetiole smooth and shining. Gaster unsculptured. Dorsal surfaces of head and alitrunk with numerous erect or suberect short hairs. Two pairs of the similar hairs present on the dorsum of petiolar node and three pairs on the dorsum of postpe- tiole as shown in Figure 133. First gastral tergite lacking erect or suberect hairs. A few pairs of forward. -projecting hairs pres- ent near the anterior clypeal margin. Scapes and tibiae with some erect to sub- erect short hairs. Color reddish-brown. Queen and Male. Unknown. Comments. This species is known only from the holotype. Its two close relatives are P. quindentatus, from Indonesia, and P. occultus, from Indonesia and Malaysia. Pristomyrmex eduardi can be separated from P. quindentatus and P. occultus be- cause it lacks pronotal armaments and pos- sesses smaller eyes (EL = 0.07, with three to four ommatidia in the longest row) in the workers. Though it was considered by Forel (1914) to be similar to P. punctatus (=P. pungens = P. japonicus), P. eduardi cannot be placed in the punctatus group because (1) it has five teeth present on the masti- catory margins of the mandibles, (2) its eyes are very small, (3) the ventral center of the clypeus is equipped with a promi- nent tooth, and (4) the petiole node has a distinct anterior face. In addition, its pro- podeal spines are much shorter than those in the four Oriental species of the punc- tatus group. Incidentally, the palp formula of P. eduardi cannot be determined from the unique holotype and thus remains un- known at this time. Ecological Information. Unknown. Pristomyrmex erythropygus Taylor Figures 134-137 Pristomyrmex erythropygus Taylor, 1968: 65. Holo- type worker, Australia: NE. New South Wales, Aca- cia Biareauh near Old Koreelah, ca. 28°24'S, (ANIC) [examined]. Diagnosis (Worker). Masticatory margin of fe aadible with three teeth: anterior clypeal margin with three strong teeth; propodeal armaments, ca. 0.13 to 0.20, usually slightly longer than pronotal spines; dorsum of head smooth, but dorsal alitrunk with several longitudinal rugae present at the juncture between the pron- otum and the mesonotum; first gastral ter- gite usually with erect or suberect hairs. Worker. TL 3.48-3.90, HL 0.90-1.08, HW 0.94—1.18, CI 104-110, SL 0.92—1.00, SI 85-97, EL) 0:16-0:20; PW 0:56—0.66; AL 0.84—0.98, PPW 0.24—0.30, PPL 0.19— 023) BRAS lO — 8): Mandibles generally smooth and _ shin- ing, but sometimes with a few basal lon- gitudinal rugae. Dentition of the mastica- tory margin rer mandible: an apical tooth + a preapical + a long diastema + a some- what truncated basal tooth. Basal margin of mandible lacking a distinctly curved lobe or tooth. Clypeus with a median lon- gitudinal carina. Anterior clypeal margin with three teeth: a median denticle and one on each side. Ventral surface of clyp- eus with a short transverse carina or with a low, broad prominence. Palp formula 2,2. Frontal carinae short, not extending to the level of the posterior margins of eyes. Antennal scrobes absent. Frontal lobes ab- sent; thus, the antennal articulations are entirely exposed. Antennal scapes, laid on the dorsal head, slightly surpassing the oc- cipital margin of head. Eyes containing eight to nine ommatidia in the longest row. Profile shape of alitrunk and pedicel seg- ments as in Figure 135A. Pronotum armed with a pair of moderately long spines, vary- ing in length from 0.08 to 0.13. Propodeal spines usually slightly longer than pronotal ones, varying in length from 0.13 to 0.20. Metapleural lobes triangular and much shorter than propodeal spines. Petiole node in profile with the anterodorsal angle higher than the posterodorsal. Anterior and dorsal faces of the postpetiole in pro- 442 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 SY Ss NY! mee oe OIA 137 Figures 134-137. Pristomyrmex erythropygus Taylor. 134A: Worker head, full-face view; 134B: Showing a short transverse carina or a low, broad prominence on the ventral clypeus; 135A: Worker, lateral view; 135B: Dorsum of the worker alitrunk, dorsal view; 136A: Queen head, full-face view; 136B: Showing a short transverse ruga, or a low, broad prominence on the ventral clypeus; 137: Queen, lateral view. file forming a single curved surface; in dor- sal view, postpetiole distinctly broader than long. Dorsum of head smooth and shining, except for a few short rugae pres- ent below the frontal carinae around the antennal fossae and on the genae. Dorsum of alitrunk possessing (1) several short ru- gae present approximately at the juncture between the pronotum and the mesono- tum (but weak in a smaller specimen), (2) a few transverse rugae present near the anterior pronotal margin, and (3) a trans- verse ridge present at the approximate po- sition of metanotal groove. Petiole, post- petiole, and gaster smooth and _ shining. Dorsal surfaces of head and alitrunk with numerous erect or suberect hairs. Dorsal surfaces of petiole node and postpetiole, respectively, with a pair of bilaterally dis- tributed long hairs; sometimes the crests of petiole node and postpetiole with ad- ditional one to two pairs of short hairs. First gastral tergite with numerous, evenly distributed, erect or suberect hairs. (Note: In three specimens placed under P. ery- thropygus, several longitudinal rugae are present at the juncture between the pron- otum and the mesonotum, but erect or REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 443 Figures 138-139. Pristomyrmex flatus sp. n. 138A: Worker head, full-face view; 138B: Showing a very short ruga on the ventral clypeus; 139: Worker, lateral view. suberect hairs are absent from the first gastral tergite. Are these hairs artificially erased? Further collecting is needed to clarify this question.) A few pairs of for- ward-projecting hairs present near the an- terior clypeal margin. Scapes and _ tibiae with some erect to suberect short hairs. Color reddish-brown to blackish-brown. Queen. TL 4.40, HL 1.14, HW 1.27, CI SO oly SON Ei nOl225 PW 40:s6: AL 1.20, PPW 0.34, PPL 0.24, PPI 142 (n = jl), General shape as in Figures 136-137, with normal caste differences from the conspecific worker; pronotum unarmed; pro-mesonotum lacking longitudinal ru- gae; propodeal spines distinctly shorter than those in conspecific worker; other characters similar to those in the conspe- cific worker. Male. Unknown. Comments. Pristomyrmex erythropygus is a sibling species of P. wheeleri and also related to P. quadridentatus. The three species are all from Australia. Pristomyr- mex erythropygus differs from P. wheeleri and P. quadridentatus because the former possesses numerous erect or suberect hairs on the first gastral tergite and several short longitudinal rugae at the juncture between the pronotum and the mesono- tum in the workers that are absent in the latter two species. In addition, the propo- deal spines are usually slightly longer than the pronotal ones in the workers of P. er- ythropygus but much shorter than the pronotal spines in P. quadridentatus. The differences between P. erythropy- gus and the two Oriental species (P. flatus Al P. collinus) and between P. erythro- pygus and the two African species (P. af- ricanus and P. trogor) are mentioned un- der P. flatus and P. africanus, respectively. Material Examined (ANIC, MCZC). Australia: New South Wales, Acacia Pla- teau, in rotten logs (Darlingtons); NE New South Wales, Nothofagus Mt., via Wood- enbong, Nothofagus forest, 1,100 m, sieved Titter, Q. M. Berlesale No. 414 (G. Monteith and G. Thompson); NE New South Wales, Gibraltar Range N.P., rain- forest, granite, sieved litter, QM. Berlesate No. 270) (Gab: Nionterth): New South Wales, Gibraltar Range Nat. Pk., 920 m, QO SileS M5222 2h) Beviecat ANIC 836, closed forest leer (L. Hill); New South Wales, 10.5 km W of Gibraltar Ra N.P. (HQ), rainforest, sieved litter, QM. Ber- lesate No. 213 (G. B. Monteith). Ecological Information. This species oc- curs in rainforest and has been collected in rotten logs and in litter berlesates. Pristomyrmex flatus sp. n. Figures 138-139 Diagnosis ( Worker). Pronotum and pro- podeum each with a pair of short spines; 444 dorsal surfaces of head and alitrunk smooth and unsculptured; petiole node with at least two pairs of hairs; anterior face of petiole node indistinguishable from the upper surface of its anterior peduncle; HW 0.98-1.04 and HL 0.94—1.02. Holotype Worker (MCZC). TL 3.94, HL 1.02, HW 1.04, CI 102, SL 1.10, SI 106, EL 0.24, PW 0.66, AL 1.03. Paratypes, three workers (MCZC, BMNB#H). Worker. TL 3.79-4.14, HL 0.94—1.02, HW 0.98—1.04, CI 102—106, SL 1.02—1.12, SI 104-108, EL 0.22—0.24, PW 0.64—0.67, AL 1.02-1.08, PPW 0.28-—0.30, PPL 0.28— 0.30, PPI 93-100 (n = 4). Mandibles generally smooth and shin- ing, with a few basal short rugae. Denti- tion of the masticatory margin of mandi- ble: the strongest apical tooth + the sec- ond strongest preapical + a long diastema + two small denticles that are about equal in size. Basal margin of mandible lacking a toothlike prominence. Clypeus de- pressed and smooth, but the frontal area with a median carina that extends a little to the clypeus. Anterior clypeal margin usually with a median denticle and three other small denticles on each side, but sometimes one of the lateral denticles very weak and indistinct. Ventral center of clyp- eus with a weak, toothlike prominence. Palp formula 1,3. Frontal carinae approx- imately reaching to the level of the pos- terior margins of eyes. Antennal scrobes absent. Frontal lobes very weak so that the antennal articulations are almost com- pletely exposed. Antennal scapes rather long, when lying on the dorsal head, sur- passing the occipital margin by one-fifth to one-fourth of their length. Eyes containing eight to nine ommatidia in the longest row. Pronotum armed with a pair of short ro- bust spines as in Figure 139. Propodeum with a pair of slender acute spines that are directed upward and slightly longer than the pronotal ones. Metapleural lobes sub- triangular. Pronotum and mesonotum in dorsal view slightly concave between the pronotal spines and between the two lat- eral margins of mesonotum, respectively. Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Petiole node massive; its anterior face in profile indistinguishable from the upper surface of petiole peduncle (Fig. 139). Dorsum of petiole node in dorsal view about rounded. Postpetiole in profile con- vex dorsally, in dorsal view slightly longer than broad or about as long as broad, broadening from front to back. Dorsum of head usually smooth and shining, but some small and shallow hair pits present. Dor- sum of alitrunk unsculptured, smooth and shining. Sides and dorsum of petiole and postpetiole smooth and shining. Gaster unsculptured. Dorsal surfaces of head with numerous erect or suberect hairs. Dorsum of alitrunk with sparse erect to suberect hairs. Two pairs of hairs present on the dorsum of petiole node, and two to three pairs on the dorsum of postpetiole. First gastral tergite lacking erect or suberect hairs. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with numerous erect to suberect short hairs. Color red- dish-brown but sometimes yellow-brown. Queen. Unknown. Male. See the following note. Comments. Like P. collinus, P. flatus is also known only from the Philippines. Pristomyrmex flatus is similar to P. colli- nus, P. quadridens, P. africanus, P. trogor, P. quadridentatus, P. wheeleri, and P. ery- thropygus. The workers of P. flatus can be immediately recognized; because in P. fla- tus, the anterior face of the petiole node is not distinct from the upper surface of its anterior peduncle, which is different in the previously mentioned species. A more detailed discussion of the sep- aration of P. flatus from P. collinus is pro- vided under P. collinus. Pristomyrmex flatus and P. collinus dif- fer from P. quadridens in the workers as follows: P. flatus and P. collinus Dorsum of alitrunk unsculptured, smooth, and shining | Dorsum of head smooth and shining, | | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang except for a few punctures bordering the frontal carinae P. quadridens Dorsum of alitrunk with scattered fo- veolate punctures; sometimes some short rugae present Dorsum of head with a few to many fo- veolate punctures, except for those bordering the frontal carinae Pristomyrmex flatus and P. collinus dif- fer from two African species, P. africanus and P. trogor, in the workers as follows: P. flatus and P. collinus Ventral center of clypeus with a short ruga or small toothlike prominence At least one to two pairs of hairs present on the petiole node and postpetiole Metapleural lobes triangular P. africanus and P. trogor Ventral surface of clypeus with two teeth Petiole and postpetiole lacking erect or suberect hairs Metapleural lobes rounded The other differences include that the anterior face of the petiole node is distinct from the upper surface of the peduncle in the workers of P. africanus and P. trogor but almost indistinct in P flatus; the dor- sum of the alitrunk in dorsal view is convex in the workers of P. collinus but distinctly shallowly concave in P. africanus and P. trogor; the dorsum of the head is sculp- tured with scattered foveolate punctures in the workers of P. africanus but is generally smooth and shining in P. flatus and P. col- linus; the frontal carinae extend to the lev- el of the posterior margins of eyes in the workers of P. flatus and P. collinus but not so in P. trogor. Pristomyrmex flatus and P. collinus dif- fer from three Australian species P. quad- ridentatus, P. wheeleri, and P. erythropy- gus in the workers as follows: P. flatus and P. collinus Anterior clypeal margin with five to sev- en smaller denticles 445 Clypeus unsculptured Masticatory margin of mandible with four teeth, consisting of an apical, a preapical, and two small basal denti- cles Palp formula 1,3. Alitrunk in dorsal view unsculptured P. quadridentatus, P. wheeleri, and P. erythropygus Anterior clypeal margin with three strong and larger teeth Clypeus sculptured with a strong me- dian carina Masticatory margin of mandible with three teeth, consisting of an apical, a preapical, and a basal tooth Palp formula 2,2 Alitrunk in dorsal view with a transverse ridge present at the approximate po- sition of metanotal groove In addition, the anterior face of the pet- iole node, in profile, is distinct from the upper surface of the peduncle in the work- ers of P. quadridentatus, P. wheeleri, and P. erythropygus but almost indistinct in P. flatus; the dorsum of the alitrunk is convex in the workers of P. collinus but almost flat or shallowly concave in P. quadridentatus, P. wheeleri, and P. erythropygus. Holotype Worker. Philippines: Luzon I., Bauqui; xi.1923. (R.C.Mcq.). Paratypes. Three workers with same data as holotype. Ecological Information. Unknown. Note: The following five male speci- mens, with same data as holotype and paratypes, may represent the male of this species. I tentatively place these males un- der P. flatus, which needs further confir- mation. Male (Figs. 262, 271). TL. 3.62—3.94, HL 0.64—0.68, HW 0.56-0.59, CI 85-91, SL 0.22-0.25, SI 39-43, HWE 0.98-1.04, EL 04820) 512 RWG074=OlS0n AealelG=1e 24s PPW 0.24-0:24" PPL 0.24—0.24 PPI 100= 100 (n = 5). Head, including the eyes, much broader than long; while excluding the eyes, dis- tinctly longer than broad. Eyes very large 446 and prominent; their length is about three- fourths of the head length. Clypeus con- vex, somewhat semicircular, its anterior margin straight and posterior one semicir- ular Palp for mula 1,3. Frontal carinae ab- sent or short, slightly beyond the anterior margins of antennal insertions. F rontal area usually with a median longitudinal ca- rina. Ocelli developed; maximum diameter of median ocellus 0.16 to 0.18. On the me- soscutum, notauli distinct, forming a Y shape, but usually without distinct ridges in them; parapsidal furrows absent. Scuto- scutellar sulcus usually with 9 to 10 narrow ridges. Propodeum lacking armaments. Metapleur: al lobes subtriangular. Middle and hind tibiae without any spurs. Petiole node in profile low, with a subtriangular apex and a rather long anterior peduncle; anterior face of the node, together with the dorsal surface of its anterior peduncle, forming a long declivity, which reaches the top of fhe nen Postpetiole in profile low, rounded dorsally, in dorsal view about as broad as long. Dorsum of head smooth and shining, except for a few short rugae present on ‘the posterior margin of clypeus. Alitrunk generally smooth antl shining, ex- cept for those marked sutures. Petiole, postpetiole, and gaster unsculptured, smooth, and shining. All dorsal surfaces with abundant long hairs. Scapes and tib- iae with numerous erect or suberect short hairs. Color reddish-brown; hairs reddish- brown; antennae sometimes yellow-brown; wings somewhat infuscated. Pristomyrmex foveolatus Taylor Figures 140-143 Pristomyrmex foveolatus Taylor, 1965: 38. Holotype worker, Australia: N. Queensland, west of Tully, Clump Point, rainforest, a few feet above sea level, 25.vi.1962 (R. W. Taylor) (MCZC) [examined]. Diagnosis (Worker). Pronotum with a pair of triangular short spines (ca. 0.06— 0.13); propodeal spines long (ca. 0.20— 0.30); dorsum of head, except for the an- tennal scrobes, with foveolate-reticulate sculpture; postpetiole unsculptured; PPI 133-150; SL 0.70-0.82 and SI 81-93. Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Worker. Tl 2.74—3.26, HL 0.78=0.92: HW 0.80—0.96, CI 100-108, SL 0.70—0.82, SI Si=93)) Ei O01 PW 0l50=0162% AL 0.68—0. 20, BRW.0:23=02273RREOAG= 0.20, PPI 133-150 (@ = 74). Mandibles usually with a few longitudi- nal rugae. Masticatory margin of aancable with three teeth: an apical + a preapical + a long diastema + a truncated basal tooth. Basal margin of mandible lacking a distinctly curved lobe or tooth. Clypeus with a strong median carina. Anterior clyp- eal margin with a median denticle and usually two to three others on each side; two or three lateral denticles are often fused into one prominence. Ventral center of clypeus with a low, broad, toothlike prominence. Palp formula 2,3. Frontal ca- rinae well developed, beyond the level of the posterior margins of eyes. Scrobal ar- eas shallow, present below the frontal ca- rinae. Frontal lobes almost completely ab- sent so that the antennal articulations are entirely exposed. Antennal scapes, when lying in the antennal scrobes, close to or just reaching the occipital margin of head. Eyes containing five to six Grae in the longest row. Profile shape of alitrunk and pedicel segments as in Figure 141. Pronotum armed with a pair of short spines (ca. 0.06—0.13). Propodeal spines long, ca. 0.20 to 0.30, usually straight but ROGUES slightly upcurved along their length. Metapleural lobe small- triangular, usually with an acute apex. Petiole hede in profile higher than long, with a long an- terior peduncle, its Lat ere dor sl angle forming an apex and its dorsum sloping downward posteriorly. Postpetiole in pro- file much higher than long, with a rounded dorsum; in der sal view, postpetiole trans- verse-rectangled, much broader than long, with the two sides subparallel. Dorsum of head, except for the antennal scrobes where there are only a few transverse ru- gae, with well-developed foveolate-reticu- late sculpture. Dorsum of alitrunk usually with foveolate-reticulate sculpture and a few coarse longitudinal rugae. Sides of REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang D» & 447 Figures 140-143. Pristomyrmex foveolatus Taylor. 140A: Worker head, full-face view; 140B: Showing a toothlike prominence on the center of ventral clypeus; 141: Worker, lateral view; 142A: Queen head, full-face view; 142B: Showing a toothlike prom- inence on the center of ventral clypeus; 143: Queen, lateral view. pronotum with a few foveolate punctures; sides of the rest of alitrunk with some ir- regularly superficial rugae. Petiole node and postpetiole smooth and shining. Gas- ter unsculptured. Dorsal surfaces of head and alitrunk with numerous erect or sub- erect long hairs. Dorsal surfaces of petiole node and postpetiole with a pair of long hairs, respectively, as shown in Figure 141. First gastral tergite lacking erect or sub- erect hairs. A few pairs of forward-pro- jecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect to suberect short hairs. Color reddish-brown. Queen. TL 3.20, HL 0.83, HW 0.86, CI 104, SL 0.74, SI 86, EL 0.16, PW 0.66, AL 0.87, PPW 0.25, PPL 0.16, PPI 139 (n = Il), General shape as in Figures 142-143, with normal caste differences from the conspecific worker; pronotum unarmed; 448 other characters similar to worker; propo- deal spine length 0.25. Male. Unknown. Comments. Pristomyrmex foveolatus is extremely similar to P. thoracicus, also from Australia, in many characters in the workers and queens, such as (1) the den- tition of the masticatory margin of man- dible, (2) palp formula, (3) structure and shape of the clypeus, (4) length and shape of the pronotal and propodeal spines, and (5) sculpture of the dorsal surfaces of the head and the alitrunk. The differences be- tween the workers of these two species are slight, as follows: P. foveolatus Antennal scapes shorter (SL 0.70—-0.82, SI 81-93) Postpetiole in dorsal view much broader than long, PPI 133-150, with the two sides subparallel, showing a_trans- verse rectangle P. thoracicus Antennal scapes longer (SL 0.86—0.98, SI 97-103) Postpetiole in dorsal view slightly broad- er than long, PPI 109-121, with the two sides not subparallel, showing a trapezoid Without doubt, P. foveolatus and P. thor- acicus are closely related to P. brevispino- sus, from the Oriental region. The workers of these three species all show the follow- ing characters: (1) a pair of short pronotal spines, (2) a toothlike prominence on the ventral center of the clypeus, (3) a similar structure and shape of petiole node in both profile and dorsal views, (4) a similar structure and shape of postpetiole in pro- file, (5) well-developed foveolate-reticulate sculpture on the dorsal head, and (6) dor- sal alitrunk with a few coarse longitudinal rugae. But P. foveolatus and P. thoracicus can be separated from P. brevispinosus as follows: The former two species have a pair of long propodeal spines (ca. 0.19— 0.30), two segments of maxillary palpi, and a truncated basal tooth on the masticatory margin of mandible in the workers and Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 queens, while P. brevispinosus has a pair of short propodeal spines or teeth (ca. 0.04—0.12), one segment of maxillary palp, and usually two small basal teeth on the masticatory margin of mandible. The separation of both P. foveolatus and P. thoracicus from the African P. cribrarius (a member of the cribrarius group) is pro- vided under P. cribrarius. Material Examined (ANIC, MCZC). Australia: N. Queensland, Clump Point, rainforest floor, ex small wood fragment (R. W. Taylor); Q., Clump Point, <20 m, berlesate (Taylor and Feehan); N. Q., NW of Daintree, Mt. Alexander, rainforest (P. F. Darlington); Q., Alexandra Bay, 16.12°S, 145.26°E, rainforest, <50 m, berlesate (Taylor and Feehan); Q. NE, Road, sum- mit on Alexandra, Ra. Daintree, 16.15°S, 145.26°E, rainforest, 250 m,_ berlesate, sieved litter (G. Monteith); Q., Kuranda, Black Mt. Rd. 360 m, 17.47°S, 145.39°E, rainforest, berlesate, sieved litter (G. Mon- teith); Q., Kuranda, Black Mt. Rd. 430 m, 16.45 X 145.35, rainforest, berlesate (Tay- lor and Feehan); Q., 4 km W of Kuranda, 450 m, 16.49 * 145.36, rainforest, berle- sate (Taylor and Feehan); Q., 1 km W of Kuranda, closed forest litter, berlesate (J. Doyen); N Q., Kuranda, RF, in log (B. B. Lowery); Q., NE, C. Tribulation, 16.08°S, 145.28°E, 20 m, rainforest, berlesate, sieved litter (G. Monteith); Q., NE, Cape Tribulation, Noah Ck, 5 m (G. B. Mon- teith); NE Q., 1.5 km W of Cape Tribu- lation (Site 3), 16.05°S, 145.28°E, 150 m, rainforest, berlesate, sieved litter (Mon- teith, Yeates, and Thomson); NE Q., 2.0 km WNW of Cape Tribulation (Site 2), 16.05°S, 145.28°E, 50 m, rainforest, ber- lesate, sieved litter (Monteith, Yeates, and Thomson); Q., near Cape Tribulation, 16.06°S, 145.28°E, 50 m, rainforest, ber- lesate (Taylor and Feehan); Q., Thorton Range, 16.14°S, 145.26°E, 100 to 150 m, rainforest, berlesate (Taylor and Feehan); Q., 20 km N of Cairns, rainforest (B. B. Lowery); NE Q., Lyons Lookout, Rex Hwy, Mossman, 400 m, rainforest, berle- sate, sieved litter (G. Monteith and D. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 449 T\\ RW ER SX CoN eae \) Figures 144-145. Pristomyrmex hirsutus sp. n. 144A: Worker head, full-face view; 144B: Showing a toothlike prominence on the ventral clypeus; 145: Worker, lateral view. Cook); NE Q., Hinchinbrook Is., Gayun- dah Creek, 18.22°S, 146.13°E, 10 to 80 m, rainforest, berlesate, sieved litter (Mon- teith, Davies, Thompson, and Gallon); Q., NE, Mossman Gorge, 16.25°S, 145.20°E, rainforest, berlesate, sieved litter (G. Mon- teith); NE Q., Bakers Blue Mt., 17 km W Mt. Molloy, 1,000 m, rainforest, berlesate, sieved litter (G. Monteith and D. Cook): Q., Mt. Cook, Nat. Pk., 15.29°S, 145.16°E, rainforest, berlesate (A. Calder and J. Fee- han); Q., Gap Creek, 15.50°S, 145.20°E, 5 km ESE of Mt. Finnigan, rainforest, ber- lesate (A. Calder and J. Feehan); Q., N Pingin Hill (J. Holt); Q., Mt. Windsor Ta- bleland, ca. 850 m, 16.18°S, 145.05°E, rainforest, berlesate (R. W. Taylor); N Q., 28 km NNW Mt. Carbine, Windsor Table- land, 900 m, rainforest, berlesate, sieved litter (Monteith, Yeates, and Cook); Q., ME, Cannon Vale, 20.16°S, 148.43°E, 10 m, dry rainforest, berlesate, stick brushing (G. Monteith); Q., ME, Mt. Dryander, 20.15°S, 148.33°E, 500 to 650 m, rainfo- rest, berlesate, stick brushing (G. Mon- teith); Q., ME, Brandy Ck Rd, Conway SF, 20.20°S, 148.42°E, 60 m, rainforest, ber- lesate, sieved litter and stick brushing (G. Monteith); Q., Finch Hatton Gorge, 21.04°S, 148.38°E, 470 m, mesophyll no- tophyll vine forest, berlesate (A. Gillison); OF Hinch Hatton) Gorges 21305 ¢ss 148.38°E, 200 m, rainforest, berlesate (R. W. Taylor and T. A. Weir); N Q., 15.50°S, 145.20°E, 12-mi scrub Gap Creek, com- plex mesophyll vine for. (Davies and Ra- ven); N Q., 20 km N of Cairns, lowland RF, creek between rocks (B. B. Lowery). Ecological Information. This species oc- curs only in rainforest in North Queens- land, Australia, and has been collected in litter berlesates; its nests are constructed in rotting logs (Taylor, 1965). Pristomyrmex hirsutus sp. n. Figures 144-145 Diagnosis (Worker). Masticatory margin of mandible with a long diastema; dorsal head with a well-developed rugoreticulum; petiole node in profile high, with a single evenly blunt-rounded apex; first gastral tergite with numerous erect or suberect short hairs. Holotype Worker (BMHH). TL 5.78, Ee SS EW eles OleOle Seale COm Sil A SEE OL2 SPW 090s Alea too RENV 0.38, PPL 0.42, PPI 90. Mandibles smooth and shining. Masti- catory margin of mandible with four teeth arranged as two adjacent strong apical teeth + a long diastema + two basal den- ticles of similar size. Basal margin of man- dible lacking a toothlike prominence. Clypeus somewhat uneven, with a few weak short rugae. Anterior clypeal margin with a median denticle and two others on each side (but one of them appears to be fused by two small denticles). Frontal ca- 450 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 146-147. Pristomyrmex longispinus sp. n. 146: rinae extending to the level of the poste- rior margins of the eyes. Antennal scrobes shallow, approximately ending at the level of the posterior margins of eyes. Frontal lobes weak so that the antennal articula- tions are almost entirely exposed. Antennal scapes long, when laid on the dorsal head, surpassing Ache occipital margin by about one-third of their length. eraaile shape of alitrunk and pedicel segments as in Figure 145. Pronotum armed with a pair of spines (ca. 0.19). Propodeum with a pair of some- what elongate-triangular teeth (ca. 0. 09). Metapleural lobes uben iangular. Petiole node in profile high (ca. 0. 76), with a sin- gle evenly blunt- rounded apex. Postpetiole in profile rounded dorsally, in dorsal view longer than broad and broadening from front to back. Dorsum of head, except for the scrobal areas, with well-developed coarse rugoreticulum. Similar sculpture present on the dorsum of the alitrunk and the two sides of the pronotum. Sides of the rest of the alitrunk with irregular coarse rugae. Petiole, postpetiole, and gaster SOoEl and shining. All dorsal surfaces of body, including hea alitrunk, petiole node, postpetiole, and gaster, with numer- ous erect or suberect hairs, as shown in Figure 145. A few pairs of forward-pro- jecting hairs present near the anterior clypeal margin. Scapes and tibiae with erect or suberect hairs. Color reddish- brown. Worker head, full-face view; 147: Worker, lateral view. Queen and Male. Unknown. Comments. Though only a single speci- men is available so fir it undoubtedly rep- resents a good species because it has nu- merous erect or suberect hairs on the first gastral tergite, which is possessed only by fis new species in the Oriental fauna of the quadridens group. In addition, this species has an uneven clypeus and a dis- tinct petiole node (bearing a single evenly blunt-rounded apex) that is not seen in the other Oriental species of the group. Holotype Worker. Philippines: Misamis or Mt. Balatukan, 15 km SW of Gingoog, 1,000 to 2,000 m, 1—5.v.1960 (H. Torrev- illas). Note: The holotype of this species is an old specimen, and many hairs have obvi- ously been removed from its first gastral tergite. Therefore, the figure can aot ac- curately show this character. Ecological Information. Unknown. Pristomyrmex longispinus sp. n. Figures 146-147, 263, 272 Diagnosis (Worker). Frontal carinae short, usually not extending to the level of the posterior margins oe eyes; pronotal spines exceptionally long (0. 42-0, 50); pro- podeal spines moderately long (0. 18—0.26); anterior face of petiole node, in profile, in- distinguishable from the upper surface of its anterior peduncle. Holotype Worker (MCZC). TL5.30, HL REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang IRA SIV 23 Cll06, Siw 345 Si 1205s EL 0.24, PW 0.80, AL 1.26. Paratypes, 73 workers and three males (BMNH, LACM, MCZC, MHNG, USNM). Worker. TL 4.52-5.62, HL 1.09—1.26, EVV 15135) Cl 103=109) SE 126=1-40; SI 103-113, EL 0.21—0.26, PW 0.76—0.86, AL 1.14—1.32, PPW 0.31—0.35, PPL 0.36— 0.40, PPI 84—89 (n = 20). Mandibles generally unsculptured, smooth, and shining. Masticatory margin of mandible with four teeth: the strongest apical + the second strongest preapical + a long diastema + two Saal teeth that are roughly the same size. Basal margin of mandible lacking a toothlike prominence. Clypeus depressed, unsculptured, and shining; its anterior margin with a median denticle and usually two others on each side, but sometimes two lateral denticles are fused into one prominence. Ventral center of clypeus with a short transverse ruga or a broad-based weak prominence. Palp formula 1,3. Frontal carinae short, usually not extending to the level of the posterior margins of the eyes. Antennal scrobes absent. Frontal lobes very weak so that the antennal articulations are almost entirely exposed. Antennal scapes long, when lying on the dorsal head, surpassing the occipital margin by about one-fifth to one-fourth of their length. Head in full- face view subglobal. Profile shape of ali- trunk and pedicel segments as in Figure 147. Pronotum armed with a pair of ex- ceptionally long spines that are about 0.42 to 0.50 and longer than the distance be- tween their bases. Propodeum with a pair of moderately long spines that are 0.18 to 0.26 and over two times the distance be- tween their bases. Both pronotal and pro- podeal spines acute and directed upward. Metapleural lobes subtriangular. Dorsum of alitrunk sometimes slightly concave. An- terior face of petiole node, in profile, in- distinguishable from the dorsal surface of its anterior peduncle (i.e., the anterior face of petiole node and the dorsal surface of peduncle forming a long declivity from the base of peduncle to the top of petiole 451 node). Postpetiole in profile rounded dor- sally, in dorsal view longer than broad. Dorsum of head generally smooth and shining, but a few weak short rugae pres- ent on the frontal area and sometimes a few foveolate punctures on the genae. Al- itrunk smooth and polished. Petiole, post- petiole, and gaster unsculptured, smooth, and shining. Dorsal surfaces of head and alitrunk with some erect or suberect long hairs. Usually two to three pairs of snag hairs present on the dorsal surfaces of pet- iole node and_ postpetiole, respectively. First gastral tergite usually lacking erect or suberect hairs, rarely with few hairs. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect or suberect hairs. Color reddish-brown. Queen. Unknown. Male (Figs. 263, 272). Three male spec- imens, together with 48 workers, collected in Dumaguete, Philippines, by J. W. Chap- man, Gonstiuite a series; and two of the three males were originally mounted, re- spectively, with a worker on the same pin: TL 4.42-4.54, HL 0.67-0.73, HW 0.64— 0.66, CI 88-96, SL 0.20-0.22, SI 31-34, HWE 0.94—0.98, EL 0.43—-0.46, PW 0.8S— 0.93, AL 1.32-1.44, PPW 0.28-—0.30, PPL 0.242026. PP 5=1l7 Gi — 3): Head, including the eyes, distinctly broader than long. “Clypeus convex, with- out a median longitudinal carina. Palp for- mula 1,3. F rental carinae sometimes ab- sent but sometimes present, just reaching the level of the posterior margins of an- tennal insertions. Maximum length of the median ocellus 0.12 to 0.12. On the me- sonotum, notauli pronounced, forming a Y shape; parapsidal furrows absent. Scuto- scutellar sulcus with 12 to 13 narrow ridg- es. Middle and hind tibiae without any spurs. Propodeum slightly tuberculate, lacking spines and teeth. Metapleural lobes prominent and subtriangular. Petiole node in profile low, with a subtriangular apex and a rather long anterior peduncle; anterior face of the Rode together with the dorsal surface of the peduncle, form- 452 ing a long declivity that reaches the top of the node. Postpetiole in profile low, round- ed dorsally, and in dorsal view broader than long. Dorsum of head smooth and shining, but frontal area usually with a me- dian longitudinal carina. Alitrunk smooth and shining, except for those marked su- tures. Petiole, postpetiole, and gaster un- sculptured, smooth, and shining. All dorsal surfaces with abundant erect or suberect hairs. Scapes and tibiae with numerous erect or suberect short hairs. Color red- dish-brown; hairs reddish-brown; wings somewhat infuscated. Comments and Discussion. Pristomyr- mex longispinus is closely related to P. cur- vulus. The former is very similar in the workers to the latter in the shape of pro- notal and propodeal spines as well as in the size, sculpture, hair, and color of body. The differences between the two species are provided under P. curvulus. Pristomyrmex longispinus may have evolved from a P. flatus—like ancestor. Ex- cept for the well-developed pronotal spines, its larger size, and shorter frontal carinae, the workers of P. longispinus are similar to those of P. flatus. It is possible that a P. collinus—like an- cestor may have split into the four species P. collinus, P. flatus, P. curvulus, and P. lon- gispinus because morphological characters show that (1) P. collinus and P. flatus are a pair of sibling species, (2) P. curvulus and P. longispinus are another pair of sibling species, and (3) PB. curvulus seems to be derived from a P. collinus—like ancestor (see the discussion under P. curvulus). This hypothesis also obtains support from biogeographic data. The four species are all endemic to the Philippines. Pristomyr- mex curvulus and P. longispinus are found only in Dumaguete and P. flatus only in Luzon. But P. collinus has a larger range; it occurs sympatrically with P. curvulus and P. longispinus in Dumaguete and with P. flatus in Luzon. Holotype Worker. Philippines: Duma- guete (J. W. Chapman). Paratypes. Philippines: 48 workers and Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 three males with same data as holotype; four workers, Dumaguete, Horns of Ne- gros (J. W. Chapman); two workers, Du- maguete, 1942 (J. W. Chapman); one worker, Dumaguete, 7.xi.43 (J. W. Chap- man); two workers, Dumaguete, v.1947 (J. W. Chapman); one worker, Dumaguete, 13.v.1947 (J. W. Chapman); one worker, Dumaguete, Silliman University, 1948 (Domingo Empeso); 13 workers, Duma- guete, 1949 (J. W. Chapman); one worker, Dumaguete, 18.vi.49. (J. W. Chapman). Additional Material Examined. More than two dozen specimens from Camp, Dumaguete, Philippines, are treated as non-type material because of in a poor sit- uation. Ecological Information. Unknown. Pristomyrmex modestus sp. n. Figures 148-149 Diagnosis (Worker). Pronotum with a pair of moderately long, robust spines; dorsal surfaces of head and _ alitrunk, as well as the two sides of pronotum, with a coarse rugoreticulum; petiole node in pro- file somewhat transrectangular, slightly longer than high, with seven to eight fo- veolate punctures. Holotype Worker (BMNH). TL 4.00, HL 0.99, HW 0.92, CI 93, SL 0.95, SI 103, EL O19) PW 0.64-AL 104) RPRW.0)35% PRIEO:355 REIOO! Mandibles with a few longitudinal ru- gae. Masticatory margin of mandible with two adjacent strong apical teeth + a long diastema + one broad, somewhat concave basal tooth. Basal margin of mandible lack- ing a distinctly curved lobe or tooth. Clyp- eus with a strong median carina. Anterior clypeal margin with a median denticle and three others on each side. Ventral surface of clypeus with a short transverse ruga. Frontal carinae strong, extending to the level of the posterior margins of the eyes. | Antennal scrobes indistinct. Frontal lobes | weak so that the antennal articulations are almost entirely exposed. Antennal scapes, laid on the dorsal head, slightly surpassing _ the occipital margin of head. Eyes rather — REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 453 Figures 148-149. Pristomyrmex modestus sp. n. 148A: Worker head, full-face view; 148B: Showing a short transverse ruga on the ventral clypeus; 149: Worker, lateral view. large. Profile shape of alitrunk and pedicel segments as in Figure 149. Pronotal spines robust, ca. 0.14, shorter than the distance between their bases. Propodeal spines acute, slender, ca. 0.11. Metapleural lobes developed, prominent, ca. 0.14, each with a rounded apex. In profile view, petiole node slightly longer than high, somewhat transrectangular, with the anterodorsal an- gle on approximately the same level as or weakly higher than the posterodorsal; in dorsal view, petiole node longer than broad. Postpetiole in profile rounded dor- sally; in dorsal view, approximately quad- rate and about as long as broad. Dorsal surfaces of head and alitrunk, as well as the two sides of pronotum, entirely sculp- tured with coarse rugoreticulum. Dorsum and the sides of petiole node with seven to eight large foveolate punctures. Post- petiole with a few shallow foveolate punc- tures. Gaster unsculptured, smooth, and shining. Dorsal surfaces of head and ali- trunk with numerous erect or suberect hairs. Petiole node and postpetiole each with a few pairs of hairs, as shown in Fig- ure 149. First gastral tergite lacking erect or suberect hairs. A few pairs of forward- projecting hairs present near the anterior clypeal margin. Scapes and tibiae with sparse erect or suberect hairs. Color red- dish-brown. Queen and Male. Unknown. Comments. This species must have evolved from the ancestor of P. costatus. It differs from P. costatus in the workers as follows: P. modestus Petiole node in profile longer than high, somewhat rectangular, with the anter- odorsal angle on approximately the same level as the posterodorsal Dorsum and sides of petiole node with seven to eight foveolate punctures P. costatus Petiole node in profile higher than long; its anterodorsal angle is distinctly higher than the posterdorsal Dorsum and sides of petiole node with- out foveolate punctures Holotype Worker. Malaysia: Sarawak, 4th Division, Gn. Mulu N.P., Kerangas for., leaf litter, 19.iii.1978 (H. Vallack). Ecological Information. The holotype is collected from a forest. Pristomyrmex nitidissimus Donisthorpe Figures 150-151 Pristomyrmex nitidissimus Donisthorpe, 1949: 411. Holotype worker, New Guinea: Maffin Bay, ix.1944 (E. S. Ross) (CASC) [examined]. 454 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 150-151. ruga on the ventral clypeus; 151: Worker, lateral view. Diagnosis (Worker). Pronotum armed with a pair of teeth; dorsal surfaces of head and alitrunk with numerous scattered fo- veolate punctures; ventral surface of clyp- eus with a coarse transverse carina; larger size (HL 1.10=1.16, HW 1.22-1.24 and EE tOh24=022'5)) Worker. TL 4.58, 4.69: HL 1.10, 1.16: FIV 2 2124 CialO al Sipe A eG: SI 93, 94: EL. 0.24, 0.25: PW 0.75, 0.78; IN IAD TO) Ge SD). Mandibles with a few longitudinal ru- gae. Dentition of the masticatory margin of mandible: the strongest apical + the second strongest preapical + a short dia- stema + a broad basal tooth showing two minute points (which is formed by the fu- sion of two basal denticles). Basal margin of mandible with a central, broadly curved lobe. Clypeus with a median longitudinal carina. Anterior clypeal margin with a me- dian denticle and two others on each side. Ventral surface of clypeus with a trans- verse ridge. Frontal carinae extending to the level of the posterior margins of eyes. Antennal scrobe indistinct, but a smooth area present below the frontal carina. Frontal lobes very weak so that the anten- nal articulations are almost entirely ex- posed. Antennal scapes, laid on the dorsal head, slightly surpassing the occipital mar- gin of head. Eyes large. Pronotum armed with a pair of teeth. Propodeum with a Pristomyrmex nitidissimus Donisthorpe. 150A: Worker head, full-face view; 150B: Showing a short transverse pair of subtriangular short spines that are slightly longer than the pronotal teeth. Metapleural lobes each with a subtrian- gular apex. Petiole node in profile as in Figure 151, with a fairly long anterior pe- duncle, in dorsal view longer than broad. Postpetiole in profile rounded dorsally, in dorsal view slightly longer than broad. Dorsum of head with numerous rather large, scattered foveolate punctures; space between foveolae usually smooth. Similar foveolate punctures present on the dorsal surface of alitrunk, but promesonotum with a smooth, unsculptured median strip. Petiole, postpetiole, and gaster smooth and shining. Dorsal surfaces of head, ali- trunk, petiole node, and postpetiole with numerous erect or suberect hairs. First gastral tergite lacking erect or suberect hairs. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect to suberect short hairs. Color blackish- brown. Queen and Male. Unknown. Comments. At first glance, P. nitidissi- mus appears to resemble P. quadridens very much, but after being compared in detail, the workers of the two species are separable as follows: P. nitidissimus Ventral surface of clypeus with a coarse transverse ruga, lacking a toothlike | prominence REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang l0a"Ges— 16)! Mandibles smooth and shining, except for a few longitudinal rugae. Masticatory margin of mandible with five teeth ar- ranged as the strongest eee + the sec- eine strongest preapical + a diastema + three sell denticles of similar size (some- times three small denticles are fused to- gether so that they are not clearly visible; or, one of them is weak or worn down, but the length of diastema is slightly shorter than the distance covered by these three teeth). Basal margin of mandible lacking toothlike prominences. Clypeus depressed and smooth, but the median carina of the frontal area extending a little to the clyp- eus. Anterior clypeal margin usually with a broad, truncated median denticle and two to three others on each side. Ventral center of clypeus with a prominent tooth. Palp formula 1,3. Frontal carinae strong, extending to the level of the posterior mar- gins of eyes. Slightly impressed scrobal ar- eas present lateral to the frontal carinae. Frontal lobes weak; thus, the antennal ar- ticulations are almost entirely exposed. An- tennal scapes, laid on the dorsal head, just surpassing the occipital margin of head. Eyes moderately sized. Occipital margin feebly concave. Pronotum with a pair of triangular teeth. Propodeum with a pair of short spines that are about two to three times the length of pronotal armaments. Metapleural lobes somewhat rounded. 468 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 167-170. Pristomyrmex quindentatus sp. n. 167A: Worker head, full-face view; 167B: Showing a tooth on the center of ventral clypeus; 168: Worker, lateral view; 169A: Queen head, full-face view; 169B: Showing a tooth on the center of ventral clypeus; 170: Queen, lateral view. Petiole node in profile with a fairly long anterior peduncle; its anterodorsal angle is on a higher level than the posterodorsal (Fig. 168). Postpetiole in profile rounded dorsally, in dorsal view about quadrate. Dorsum of head, except for the shallow scrobes, with numerous scattered foveo- late punctures. Dorsum of alitrunk pos- sessing scattered foveolate punctures, usu- ally with a smooth and unsculptured me- dian strip. Petiole smooth and _ shining, with a weak longitudinal ruga on each side. Postpetiole and gaster unsculptured. Dor- sal surfaces of head and alitrunk with nu- merous erect or suberect short hairs. Two pairs of hairs usually present on the dor- sum of petiole node, and a few pairs on the dorsum of postpetiole, as illustrated in Figure 168. First gastral tergite lacking or bearing few suberect hairs. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with numerous erect to suberect short hairs. Color reddish-brown. | Queen. TL 3.64, 3.70; HL 0.83, 0.83; | HW 0.82, 0.83; CI 99, 100; SL 0.72, 0.78; | SI 88, 94; EL 0.18, 0.19; PW 0.64, 0.64; | AL 0.97, 1.00: PPW 0.23, 0.23: PPI 0.228 | O23 ERE LOOMO05a¢7— 2): | Generally similar to worker, except for — normal caste differences. In addition, | pronotum unarmed; propodeal armaments toothlike, shorter than those in the con-— specific worker, and the dorsum of alitrunk | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang lacking an unsculptured median longitu- dinal strip. Male. Unknown. Comments. Pristomyrmex quindentatus is closely related to P. occultus. The two species occur in the Oriental region. Their workers share a critical character, that is, the masticatory margin of mandible with an apical tooth + a preapical tooth + a diastema + three small denticles of similar size; this dentition is not seen in all other Pristomyrmex species, except in P. eduar- di. But P. quindentatus and P. occultus can be separated from P. eduardi in the work- ers in having a pair of pronotal teeth. The worker of P. quindentatus differs from that of P. occultus as follows: P. quindentatus Dorsal surfaces of head and _alitrunk only with scattered, shallow foveolate punctures; dorsum of alitrunk with an unsculptured median longitudinal strip Anterior clypeal margin with a truncat- ed median tooth A lateral carina lacking or vestigial on each side of the petiole node SL 0.76—0.80, SI 94—100 P. occultus Dorsal surfaces of head and alitrunk en- tirely covered with coarse rugoreti- culum Median tooth on the anterior clypeal margin not truncated Petiole node with a lateral longitudinal carina on each side SieO 72-0 os Sl o7—93 Pristomyrmex quindentatus may have evolved from the ancestor of P. quadri- dens. Pristomyrmex quindentatus can be separated from P. quadridens as follows: the masticatory margin of the mandible possesses five teeth in the workers and queens of P. quindentatus but at most four teeth in P. quadridens; in addition, the propodeum is armed with a pair of short spines in the workers of P. quindentatus but a pair of teeth in P. quadridens. Holotype Worker. Indonesia: Seram, 469 above Haruru, near Masohi, rainforest, 50 to 150 m, 18.iii. 1981 (W. L. Brown). Paratypes. Five workers and two queens with same data as holotype. Additional Material Examined (MCZC). A worker collected in Indonesia (Blawan, Idjen, Dammerman, 950 m) has the fol- lowing measurements: TL 3.50, HL 0.97, HW 0.95, CI 99, SL 0.84, SI 88, EL 0.14, PW 0.62, AL 0.94. It shows some varia- tion: (1) The propodeum is armed with a pair of triangular teeth, (2) the dorsum of the head possesses dense foveolate punc- tures, (3) a longitudinal carina is present on each side of the petiole, and (4) three small denticles on the masticatory margin of the mandible are worn down and are not clearly visible. Ecological Information. This species has been collected in rainforest. Pristomyrmex sulcatus Emery stat. n. Figures 171-176, 266, 276 Pristomyrmex brevispinosus sulcatus Emery, 1895: 464. Syntype workers, queen, Burma: Carin Cheba, 500 to 1,000 m, xii.1887 (L. Fea) [syntype workers (MCSN, NHMYV) examined]. Diagnosis (Worker). Ventral surface of clypeus with a strongly prominent tooth at the center; pronotum with a pair of mod- erately long spines (0.14—0.20), usually longer than propodeal armaments (0.07— 0.13); dorsum of head with foveolate-retic- ulate sculpture or rugoreticulum; petiole node lacking foveolate punctures; first gas- tral tergite lacking erect or suberect hairs. Worker. TL 3.98—4.74, HL 0.98-1.16, HW 0.96—-1.10, CI 93-105, SL 1.02—1.15, SI 101—11t, El 0,140.19, Pw 0:64-0:75, AL 0.98—1.18, PPW 0.26—0.30, PPL 0.26— 0.32, PPI 90-100 (n = 30). Mandibles usually with some longitudi- nal rugae, varying from superficial to dis- tinct. Masticatory margin of mandible with four teeth arranged as the strongest apical + the second strongest preapical + a long diastema + two small basal teeth of similar size that are fused together. Basal margin of mandible almost straight, lacking a dis- tinctly convex lobe or tooth. Clypeus with Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 A470 Aas ie “es oF —176. Pristomyrmex sulcatus Emery. 171A: Syntype worker head, full-face view; 172: Syntype worker, lateral view; | 173A: Non-type worker head, full-face view; 174: Non-type worker, lateral view; 175A: Queen head, full-face view; 176: Queen, | lateral view; 171B, 173B and 175B: Showing that a prominent tooth is present on the ventral center of the clypeus in the syntype | worker, non-type worker and queen, respectively. Figures 171 | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang a median longitudinal carina. Anterior clypeal margin with a median denticle and two to three others on each side; but sometimes two lateral denticles are fused into a larger tooth. Ventral center of clyp- eus with a strongly prominent tooth. Palp formula 1,3. Frontal carinae strong, ex- tending to the level of the posterior mar- gins of eyes, forming the dorsal margins of the shallow antennal scrobes. Frontal lobes very weak; thus, the antennal artic- ulations are almost entirely exposed. An- tennal scapes, when lying on the dorsal head, slightly surpassing the occipital mar- gin. Eyes usually containing six to seven Bemacdia in the longest row. Profile shape of alitrunk and pedicel segments as in Figures 172 and 174. Pronotmn with a pair of spines, varying in length from 0.14 (in the type series) to 0.20. Propodeum with a pair of teeth or short spines varying from 0.07 to 0.13. Metapleural lobes sub- triangular or somewhat rounded. Petiole Ride. in profile high, with a fairly long an- terior peduncle; its anterodorsal angle is on a higher level than the posterodor sal. Postpetiole in profile rounded dorsally, in dorsal view broadening from front to back, about as long as or slightly longer than broad. Dorsum of head, except for the scrobal areas where rugae are somewhat weak, with course rugoreticulum or dense alveolate punctures. samulae but slightly sparse sculpture present on the dorsum of alittunk and often on the two sides of pronotum. Petiole, postpetiole, and gaster smooth and _ shining. head and alitrunk PRP l00;Ga—s)) Head, including the eyes, distinctly broader than long. Clypeus convex, with- out a median longitudinal carina. Palp for- mula 1,3. Frontal carinae weak and short, just reaching the level of the posterior margins of antennal insertions. Maximum diameter of the median ocellus 0.10. On the mesonotum, notauli strongly marked, forming a Y shape; parapsidal AunGuis ab- sent. Salo: scutellar sulcus rather broad, with 10 ridges that expand at the upper end. Propodeum slightly tuberculate, lack- ing teeth or spines. Metapleural lobes sub- tr iangular. Middle and hind tibiae without any spurs. Petiole node in profile low, with a fairly long anterior peduncle. Postpetiole low, in dorsal view about as long as broad. Dorsum of head generally oot and shining but with a median longitudinal ca- rina present on the frontal area fand several short rugae on the posterior clypeal mar- gin. Alieeani smooth and shining, except for those marked sutures. Petiole, post- petiole, and gaster smooth and shining. All dorsal surfaces with abundant erect or suberect hairs. Scapes and legs with erect or suberect short hairs. Body reddish- brown; funicular segments of antennae white and wings slightly light-yellow. At the present, dhe male of P. Ruleatis is al- most indistinguished from the males of P. brevispinosus and P. quadridens. Discussion and Comments. Taxonomic status of “sulcatus” is somewhat compli- cated. The syntype workers of P. brevispi- nosus sulcatus differ from those of P. bre- vispinosus by possessing a pair of fairly long pronotal spines (0.15—0.16), as com- pared with a pair of teeth (0.06) in the latter. However, after examining all avail- able material, I find that the length of the pronotal armaments is continuously vari- able (from 0.06, as in the syntypes of P. brevispinosus, to 0.20, as in the specimens from Khao Yai Nat. Park, Thailand). The syntype workers of “sulcatus”, in fact, are intermediates between the two extreme ends (i.e., in one extreme, the pronotal teeth are slightly shorter than or about as long as the propodeal teeth; in the other extreme, the pronotal spines are two to three times as long as the propodeal ar- maments; see Figs. 114, 172, 14D), Te keep “sulcatus” as a valid name (i.e., raise it to the rank of species instead of assigning it as a junior synonym of P. brevispinosus) because more ecological work must be done before the status of “sulcatus” be- comes clarified. With this tentative pro- posal, P. sulcatus comprises those popula- tions with pronotal spines (ca. 0.14—0.20), distributed in Pahang (1,300-1,720 m), Malaya, and in northwest and central Thai- land, Burma, Nepal, and southwest China. Pristomyrmex brevispinosus comprises populations with toothlike pronotal arma- ments (ca. 0.06—0.10), occurring in Pahang (1,250 m), Malaya, and Trang Province of South Thailand (07.55°N) and in the Phil- ippines, Taiwan, and Japan. In other words, P. sillectans may be a northerly re- placement OEE. brevispinosus along the Malay peninsula. The separation of P. sulcatus from P. costatus is provided under P. costatus. The worker of P. sulcatus differs from that of P. hirsutus by its petiole node with the an- terodorsal angle higher than the postero- dorsal and its first gastral tergite lacking erect or suberect hairs. The worker of P Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 f sulcatus differs from that of P. modestus by its petiole node higher than long and | lacking distinct foveolate punctures. The | worker of P. sulcatus differs from that of | P. occultus by its masticatory margin of the | mandible possessing at most four teeth | and its propodeal armaments not longer than the pronotal spines. The queen of P. sulcatus differs from that of P. bicolor by. possessing a strongly prominent tooth on | the ventral center of the clypeus. The | worker and queen of P. sulcatus differ | from those of the Australian P. foveolatus and P. thoracicus as follows: | P. sulcatus Pronotal spines longer than or about as_ long as the propodeal armaments | (worker) | Propodeum with a pair of teeth (Fig. 176) (queen) Maxillary palp with one segment (work-_ er and queen) | P. foveolatus and P. thoracicus Pronotal spines much shorter than the propodeal spines (worker) Propodeum with a pair of fairly long spines (Figs. 143, 180) (queen) Maxillary palp with two segments (work-_ er and queen) Material Examined (ANIC, BMNH, | IZAS, LACM, MCZC, MHNG, NHMV,} USNM). Thailand: Nakhon Ratchasima Prov., Khao Yai Nat. Park, 700 to 750 m,| hill forest, rotten wood (I. Burikam and W. | L. Brown); Khao Yai Nat. Park (L6bl and! Burckhardt); Chiang Mai (54 km NE Chiang Mai), Mae Nang Kaeo, 900 m) (Burckhardt and L6bl); Chiang Mai Pr. st Doi Suthep NP (7), near Ruesse Care, 900 | to 1,000 m (Zettel); Chiang Mai Prov., | Chiang Dao Wildlife Sanctuary, 19. 29°N | 98. 97°, 470 to 500 m elevation (Saowana_ Sontintclnet). Nepal: Khandbari District, Arun River at Num 1,500 to 1,600 m (A. / and Z. Smetana). Burma: Carin Cheba (L. 7 Fea). China, Yunnan Prov. (?). W. Malay- | sia: Pahang, Cameron Highls, Gunung Ja-_ sar, 1,720 m (L6blI and Calame); Malaysia: Pahang, Cameron Highlands, Tanah Rata | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 473 Figures 177-180. Pristomyrmex thoracicus Taylor. 177A: Worker head, full-face view; 177B: Showing a tooth on the center of ventral clypeus; 178: Worker, lateral view; 179A: Queen head, full-face view; 179B: Showing a toothlike prominence on the center of ventral clypeus; 180: Queen, lateral view. Umg., Gn. Jasar, 1300 m (Schuh and Lang)?: Kabu, 400 ft (Abor Exped.). Ecological Information. This species has been collected in forests. Pristomyrmex thoracicus Taylor Figures 177-180 Pristomyrmex thoracicus Taylor, 1965: 41. Holotype worker, Australia: N. Queensland, Lake Eacham National Park, rainforest, 2,500 ft, 6.vi.1962 (R. W. Taylor) (MCZC) [examined]. Diagnosis (Worker). Pronotum with a pair of triangular short spines (ca. 0.06— 0.08); propodeal spines long (ca. 0.19- 0.24); dorsum of head, except for the an- tennal scrobes, with foveolate-reticulate sculpture; postpetiole unsculptured; PPI 109-121; SL 0.86—0.98 and SI 97-103. Worker. TL 3.22-3.72, HL 0.86—0.96, HW 0.86-—0.96, CI 97-101, SL 0.86—0.98, SI 97-103, EL 0.10—0.13, PW 0.54—0.60, AL 0.84—0.96, PPW 0.23—0.26, PPL 0.20— O22 SE RIGhOS= 24) Mandibles with a few longitudinal ru- gae. Masticatory margin of mandible with three teeth arranged as an apical + a preapical + a long diastema + a truncated basal tooth. Basal margin of mandible lack- ing a distinct curved lobe or tooth. Clyp- eus with a strong median longitudinal ca- rina. Anterior clypeal margin with a me- dian denticle and usually two others on A474 each side, but sometimes two lateral den- ticles are fused into a larger tooth. Ventral center of clypeus with a low, broad, tooth- like prominence. Palp formula 2,3 . Frontal carinae well developed, beyond i level of the posterior margins of the eyes. Scro- bal areas shallow, present lateral to the frontal carinae in full-face view. Frontal lobes almost completely absent so that the antennal articulations are entirely exposed. Antennal scapes usually slightly surpassing the occipital margin of head when lying in the antennal scrobes. Eyes with five to six ommatidia in the longest row. Profile shape of alitrunk and pedicel segments as in Figure 178. Pronotum with a pair of tri- angular short spines, ca. 0.06 to 0.08. Pro- podeal spines long, ca. 0.19 to 0.24, usually slightly upeurved at their apices. Meta- pleural lobe small-triangular with a rather acute apex. Petiole node in profile higher than long, with a long anterior peduncle, its adicrodoteal angle forming an apex and its dorsum sloping dower posteriorly. In dorsal view, crest of petiole node strongly convex. Postpetiole in profile rounded dorsally, in dorsal view broader than long and broadening from front to back. Dorsum of head between the frontal carinae, as well as the two sides of the dor- sal head, with foveolate-reticulate sculp- ture. Antennal scrobes rather smooth, with only a few weak rugae. Dorsum of anten- nal scape with a longitudinal carina. Dor- sum of alitrunk with a rugoreticulum. Pet- iole and postpetiole smooth and _ shining. Gaster unsculptured. Dorsal surfaces of head and alitrunk with numerous erect or suberect hairs. Dorsal surfaces of petiole node and postpetiole each with a pair of hairs. First gastral tergite lacking erect or suberect fires A few pairs of fee pro- jecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect to suberect short hairs. Color reddish-brown. Queen. TL 3.844.10, HL 0.94—0.96, HW 0.94—0.96, CI LOO—100, SL 0.94—0.96, SI 98-100, EL 0.16—0.18, PW 0.70_-0.72, Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Al e061 4S PPWa0.25=0)27 SE Rin 022= O225P RIAA 23h os General shape as in Figures 179-180, with normal caste differences from the conspecific worker; pronotum unarmed; crest of petiole node in dorsal view feebly | convex; other characters similar to worker. | Male. Unknown. | Comments. Pristomyrmex thoracicus is | so far known only from North Queensland, | Australia. It is a sibling species of P. fov- | eolatus, also from North Queensland. | Their differences are given under P. fov- eolatus. Characters separating P. thoraci- | cus from the Asian P. brevispinosus, and from the African P. cribrarius (a member - of the cribrarius group) are provided un- der P. foveolatus and under P. cribrarius, | respectively. | Material Examined (MCZC, ANIC). | Australia: Queensland, Lake Eacham, - rainforest, 2,500 ft, nest ex rotten log (R. | W. Taylor); N.Q., Kuranda, Tainiorest | 1,100 ft, stray floor (R. W. Taylor); N.Q., | Cravtonds Lookout, Beatrice River (Dar-. lingtons); N.Q., Malanda, rainforest, rotten | log (W. L. Brown); N.Q., 3.2 km E of Lake Barrine, ca. 700 m, rainforest, ex rotten. log (R. W. Taylor and J. Feehan); N.Q.,} Iralce Barrine Nat. Pk., 760 m, 17.l5°Sm 148.38°E, rainforest, ex rotting log (R. W. Taylor and T. A. Weir); N.Q., Palmerston | N.P.. ca. 1,000 ft, rainforest, nest in soil | under log (R. W. Taylor). | Ecological Information. This species oc- | curs in rainforest nesting in rotting logs. and in soil under logs (Taylor, 1965, 1968). | | Pristomyrmex trachylissus (F. Smith) Figures 181-184 Myrmica trachylissa F. Smith, 1858: 126. Holotype | queen, Borneo (A. R. Wallace) (BMNH) [exam- | ined]. Pristomyrmex trachylissus (F. Smith) Mayr, 1886: | 359. Diagnosis (Worker). Large size (HL = | 1.36, HW Ses): masticatory margin of mandible with five teeth; pronotum Heh a | pair of exceptionally long spines that are | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang OS J Sy ey, L 3 ZEN A475 Figures 181-184. Pristomyrmex trachylissus (F. Smith). 181A: Worker head, full-face view; 181B: Showing a short ruga on the ventral clypeus; 182: Worker, lateral view; 183A: Queen head, full-face view, antennae missing; 183B: Showing a short ruga on the ventral clypeus; 184: Queen, lateral view. about as long as the distance between their bases. Worker. TL 5.58-6.48, HL 1.36-1.46, IVY 1-45—1.62, Cl 107-114, SL 1.40=1.50, ol 90-98, EL 0.22-0.24 PW 0.92-1.02, AL 1.50-1.64, PPW 0.35—0.39, PPL 0.44— 0.48, PPI 76-83 (n = 6). Mandibles generally smooth and shin- ing, except for a few superficial rugae. Masticatory margin of mandible with five teeth arranged as the strongest apical + the second strongest preapical + a small third tooth + a short diastema (or this di- astema indistinct) + two small basal teeth. Basal margin of mandible with a central, broadly curved lobe. Clypeus with a strong median longitudinal carina. Anterior clyp- eal margin with a median denticle and a broad prominence on each side. Ventral surface of clypeus generally smooth and shining but sometimes with a weak, short, transverse ruga at the center. Frontal ca- rinae strong, extending to the level of the posterior margins of the eyes. Antennal scrobes shallow, present lateral to the fron- tal carinae. Frontal lobes weak; thus, the antennal articulations are almost entirely exposed. Antennal scapes, laid on the dor- sal head, slightly surpassing the occipital margin of head. Eyes with 10 to 11 om- matidia in the longest row. Profile shape of alitrunk and pedicel segments as in Figure 182. Pronotum armed with a pair of ro- bust, exceptionally long spines, ca. 0.40 to 0.50, about as long as the distance be- tween their bases. Propodeum with a pair of short spines, ca. 0.12 to 0.18, shorter than or at most about as long as the dis- 476 tance between the bases of two propodeal spines. Metapleural lobes each with a somewhat blunt-rounded apex. Petiole node in profile high, with a long anterior peduncle; its anterodorsal angle is on a higher level than the posterodorsal. Post- petiole in profile rounded dorsally, in dor- sal view longer than broad and broadening from front to back. Dorsum of head en- tirely sculptured with well-developed coarse rugoreticulum. Similar sculpture present on the dorsum of alitrunk and on the two sides of pronotum, except for the space between the bases of two pronotal spines, which is rather smooth. Petiole smooth and shining but with a lateral lon- gitudinal carina on each side. Postpetiole and gaster unsculptured, smooth, and shining. Dorsal surfaces of head arial ali- trunk moth numerous erect or suberect hairs. A few pairs of hairs present on the dorsum of petiole node and at least a pair on the postpetiole, as shown in Figure 182. First gastral tergite lacking ATE or sub- erect “Thetin, A ian pairs ae forward-pro- jecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect or suberect hairs. Color red- dish-brown, but the masticatory margins and the basal margins of mandibles and the funiculi of antennae black-brown. Queen. TL 7.55, HL 1.84, HW 2.14, CI 116, SL- (antennal scapes missing), SI-, PW 52) Ale 2:10) (2 = 1): General shape as in Figures 183-184, with normal caste differences from the conspecific worker; pronotum unarmed; other characters similar to worker. Male. Unknown. Comments. Pristomyrmex trachylissus must have evolved from the ancestor of P. bicolor. It is extremely similar in appear- ance of the workers and queens to P. bi- color. The two species may occur sympat- rically in Sarawak and in Sabah. Their workers and queens can be separated by the following characters: P. trachylissus Masticatory margin of mandible with five teeth; diastema indistinct or very short Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Basal margin of mandible with a central, broadly curved lobe Anterior clypeal margin with a median tooth and a broad, short pone on each side SI < 100 (only in worker). P. bicolor Masticatory margin of mandible with. four teeth; a long diastema present between the preapical and the third tooth Basal margin of mandible lacking a dis- tinctly curved lobe Anterior clypeal margin usually with seven denticles, but in some speci- mens, two or three lateral denticles fused into a broadly convex lobe SI > 105 (only in worker). Material Examined (BMNH, MCZC, ANIC). East Malaysia: Sarawak, 4th Div., G. Mulu Nat. Pk., RGS Expd., Long pala, lowland Pintorese on log and on rotten log (B. Bolton); North Borneo (SE), For- est Camp, 19 km N of Kalabakan 180 m (Y. Hirashima). Ecological Information. This species oc- curs in rainforest and has been collected on a rotten log. Pristomyrmex trogor Bolton Figures 185-186 Pristomyrmex trogor Bolton, 1981: 287. Holotyped worker, Zaire (B. Congo on the label): S slope of. Mt. Kahuzi, 1,900 m, 5.ix.1957 (E. S. Ross and R. E. Leech) [CASC (Bolton, 1981)]; [five paratype | workers (MCZC) examined]. Diagnosis (Worker). Frontal carinae ab-- sent; ventral surface of clypeus with two. toothlike prominences; pronotum with a pair of short spines; dorsal surfaces of head | and alitrunk smooth and shining; petiole and postpetiole lacking hairs. Worker. TL 3.42-— 3.98, HL 0.88-1. 00, HW 0.92-1.02, CI 102-105, SL 0.96—1.06, | SI 100-104, EL 0.14—0.16, PW 0.56—0.62, | AL 0.84—0.94, PPW 0.23-0.26, PPL 0.22—| OLR6SRPI100=106KGis—ao): Mandibles smooth, with only a few weak. longitudinal basal rugae. Masticatory mar-_ REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang AT ~l Figures 185-186. Pristomyrmex trogor Bolton. 185A: Worker head, full-face view; 185B: Showing two toothlike prominences on the ventral clypeus; 186: Worker, lateral view. gin of mandible possessing the strongest apical tooth + the second strongest preap- ical tooth + a diastema + two small basal denticles that are often fused into a broad, short tooth. Basal margin of mandible lacking a toothlike prominence or curved lobe. Clypeus lacking a median longitudi- nal carina. Ventral surface of clypeus with two strongly prominent teeth. Anterior clypeal margin with a median tooth and two lateral denticles on each side; some- times two small lateral denticles are fused into one prominence. Palp formula 2,3 (Bolton, 1981). Frontal carinae absent. Antennal scrobes absent. Frontal lobes in- distinct. Antennal scapes, when lying on the dorsal head, just reaching or slightly surpassing the occipital margin of head. Eyes containing five to six ommatidia in the longest row. Promesonotum in dorsal view weakly concave. Pronotum and pro- podeum each with a pair of short spines (Fig. 186). Metapleural lobes rounded. In profile view, petiole node high, higher than long, with a long anterior peduncle, its an- terodorsal angle higher than the postero- dorsal. In dorsal view, petiole node about as broad as long. Postpetiole in profile higher than long, rounded dorsally, in dor- sal view broadening from front to back. All dorsal surfaces unsculptured, smooth, and shining. Dorsum of head with some fine long hairs. Dorsal surface of alitrunk with only one to two pairs of hairs that arise from the lateral margins of the mesono- tum. Petiole node, postpetiole, and first gastral tergite lacking erect or suberect hairs. A row of fine, forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some fine, appressed or decumbent, or suberect hairs. Color reddish-brown. Queen. Unknown. Male. One paratype male has the same data as holotype and was originally mount- ed together with two paratype workers on the same pin: TL 2.92) HL 0155, HW 0.57, CHAl04 SEO S05 a3. FIVE O94 Re 0:28, PW. 0:82; Aly 16 @ = 1): Head, including the eyes, distinctly broader than long. Clypeus narrow and convex, its anterior margin rather straight. Frontal carinae absent. Maximum diame- ter of the median ocellus 0.10. Scapes lon- ger than the other antennal segments, ex- cept for the apical ones. On the meso- scutum, notauli distinct, forming a Y shape; parapsidal furrows absent. Scuto- scutellar sulcus with about 10 narrow short ridges. Propodeum weakly tuberculate, lacking teeth or spines. Metapleural lobes subtriangular. Middle and hind tibiae with- out any spurs. Waist abnormally shaped, with a huge segment formed by the fusion 478 of petiole and postpetiole; entire posterior face of the waist attached to first gastral segment. Dorsum of head smooth and shining, but frontal area with a median longitudinal carina; a short ruga present below each antennal socket. Alitrunk gen- erally smooth and shining, except for those marked sutures. Petiole, postpetiole, and gaster smooth and shining. All dorsal sur- faces with abundant long haus. Scapes and tibiae with numerous erect or suberect short hairs. Color somewhat blackish- brown; wings white. Con menis Pristomyrmex trogor is sim- ilar to another African species, P african- us, in many characters of the workers, such as dentition of the masticatory margin of mandible; two teeth present on the ventral clypeus; promesonotum in dorsal view shallowly concave or flat; pronotum and propodeum each with a pair of short spines; the structure and shape of petiole, postpetiole, and metapleural lobes; as well as the distribution of hairs. But P. trogor is easily distinguished from P. africanus and from the other African species of the genus because P. trogor is the only species lacking frontal carinae in the workers. In Added. the dorsum of the head is smooth and shining in the workers of P. trogor but has foveolate punctures in P. africanus, P. fossulatus, and P. cribrarius; the pronotum possesses a pair of short spines in the workers of P. trogor that is not seen in P. fossulatus and P. orbiceps. The separation of P. trogor from the two Asian species (P. flatus and P. collinus) and from the three Australian species (P. whee- leri, P. erythropygus, and P. quadridenta- tus) is given under P. flatus and under P. africanus, respectively. Distribution. Zaire (known only from the type series). Ecological Information. Unknown. Pristomyrmex wheeleri Taylor Figures 187-192 Pristomyrmex wheeleri Taylor, 1965: 48. Holotype worker, Australia: SE Queensland, ca. Binna Burra, Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 | rainforest, 2,800 to 3,000 ft, nest under stone, 21—| 25.v.1962 (R. W. Taylor) (MCZC) [examined]. i Diagnosis (Worker). Masticatory margin | of mandible with three teeth; antenet™ : clypeal margin at most with three teeth; propodeal spine length ca. 0.18 to 0.28,) about equal to or slightly longer than pro-| notal spines; dorsum of alitrunk smooth, but with a transverse ridge present at the approximate position of Perot groove; petiole node and postpetiole dor sally witht some hairs; first gastral tergite lee hairs. Worker, Wii 326-454. Hi 0.92-1" 17, HW 0.97—1.34, Cl 103-116, SL 0:94=1 iss | SI 88-100, EL 0.17—0.24; PW 0.56-0.74, | AL 0.88—1.08, PPW 0.24—0.31, PPL 0.22— 0.28, PPI 104-122 (7 = 40). Mandibles usually smooth and shining, except for some small hair pits. Mastica- tory margin of mandible with three teeth arranged as an apical + a preapical + a| long ate + a somewhat truncated Racal tooth. Basal margin of mandible lack- ing a curved lobe or roan. Clypeus with a Lace longitudinal carina. Anterior clyp-. eal margin usually with three strong teeth: a Sateen tooth and one on each side, but sometimes the median tooth weak or ves-. tigial. Ventral surface of clypeus usually with a short transverse carina. Palp for- mula 2,2. Frontal carinae not or just ex-| tending to the level of the posterior mar- gins of the eyes. Antennal scrobes absent. Frontal lobes nearly completely absent; thus, the antennal articulations are almost) entirely exposed. Antennal scapes, laid on the dorsal head, slightly surpassing the oc-| cipital margin. Hye. usually containing: eight to nine ommatidia in the longest row. | Profile shape of alitrunk and pedicel seg- ments as in Figure 188. Pronotal spines: varying in length, from 0.12 to 0.28. Pro-| podeal spines Ree 0.18—0.28) equal to or} slightly longer than the pronotal spines. | Metapleural lobes small, usually triangular. Petiole node in profile cath a long anterior. peduncle; its anterodorsal angle is on a higher level than the posterodor sal: some-| | | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang A479 wie ; \ a Figures 187-192. Pristomyrmex wheeleri Taylor. 187A: Worker head, full-face view; 187B: Showing a short ruga on the ventral clypeus; 188: Worker, lateral view; 189A: Ergatoid queen, full-face view; 189B: Showing a short ruga on the ventral clypeus;: 190: Ergatoid queen, lateral view; 191A: Queen head, full-face view; 191B: Showing a short ruga on the ventral clypeus; 192: Queen, lateral view. 480 times posterodorsal angle indistinct, show- ing a single curved surface. Postpetiole in profile as in Figure 188, in dorsal view slightly broader than long and broadening from front to back. Dorsum of head smooth and shining. Dorsum of alitrunk smooth, but with a transverse ridge pre- sent at the approximate position of metan- otal groove. Petiole, postpetiole, and gaster smooth and shining. Dorsal surfaces of head and alitrunk with numerous erect or suberect long hairs. A pair of long hairs bilaterally distributed on the dorsum of petiole node and on the postpetiole, re- spectively; sometimes the crests of both petiole node and postpetiole with addi- tional one to two pairs of short hairs. First gastral tergite lacking erect or suberect ours. Usually, three pairs of forward-pro- jecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect to suberect short hairs. Color: head deep reddish-brown; alitrunk, pedi- cel segments, and gaster yellow-brown to reddish-brown. Ergatoid Queen. TL 4.06, HL 1.08, HW WANG 1 WOW SE Le SIMS TRIE OPAL EY OOSEPAL, 106, PST (0584 BSE2 0:25: RPWi80:28)) PPPs 0265 PEN 108 @ =o). Note: This is a paratype). General shape as in Figures 189-190. Similar to worker; color and pilosity as in worker, but head with three ocelli and pro- mesonotum convex. Flight sclerites and wings lacking, but a black speck is present on each lateral margin of the mesonotum. Queen. TL 4.62—5.06, HL 1.09-1.28, EValelS Ae ClO ald Silzel-O06=122.0) SI 82-90, EL 0.22-0.28, PW 0.80-0.90, AL 1.14-1.34, PPW 0.30-0.36, PPL 0.26— OSOSBPl lO s3G — to)! General shape as in Figures 191-192, with normal caste differences from the conspecific worker; a pair of acute minute spines present on the humeral angles of the pronotum; propodeal spines rather long, ca. 0.20 to 0.24; other characters sim- ilar to worker. Male. Unknown. Comments. Pristomyrmex wheeleri oc- Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 curs in East Australia. Its two close rela- tives are P. erythropygus and P. quadri- dentatus, also from East Australia. Char- acters separating P. wheeleri from P. ery- thropygus and from P. quadridentatus are provided under P. erythropygus and under P. quadridentatus, respectively. The differ- ences between P. wheeleri and the two Asian species (P. flatus and P. collinus) and between P. wheeleri and the two African species (P. africanus and P. trogor) are giv- en under P. flatus and P. africanus, re- spectively. | Material Examined (ANIC, MCZC). Australia: SE Queensland: Mt. D’Aguilar. Range, 2,000 ft, rainforest, ex rotten log (R. W. Taylor); Cunningham's Gap, 3,000 ft, rainforest floor, ex small wood fragment (R. W. Taylor); Cunningham’s Gap, rain- forest, 2,500 ft, nest under stone (B. B.. Lowery); National Pk. (H. Hacker); Tam- borine Mt., rotting leaves (A. M. Lea);; Tamborine Mt., S side, Curris Falls, rain-. forest, berlesate leaf mold (T. E. Wood- ward); Tamborine Mt., rainforest, 2,000 ft, nest between stones (B. B. Lowery); ca. Binna Burra, rainforest, 2,600 to 3,000: ft, nest under stone (R. W. Taylor); M’Pheraon Rge, v. Binna Burra, rainforest, 2,600 to 3,600 ft (P. F. Darlington); Binna’ Burra, Lamington Nat. Pk., leaf and log litter (J. and N. Lawrence); Lamington| Nat. Park (O’Reillys), 28.14°S, 153.08°E,; rainforest, ca. 920 m, ex snl fragment rotten wood (R. W. Taylor and R. Kohout ll Mt. Chinghee, 12 km SE Rathdowney, 28.19°S, 152. 58°E, 720 m, rainforest, stick brushing (Monteith, Yeates, and Thomp-| son). New South Wales: Unumgar Forest (Darlingtons); Woodenbong, rainforest (Darlingtons); Tooloom Range, ca. 2,000 ft (Darlingtons): Mt. Warning, Painforeem 800 to 3,500 ft, under and heaven rocks/ nest in wat rotten log but 2 inches below ground (B. B. Lowery); Mt. Warning, 10 fa from Murwillumbah, RF, ca. 3,000 ft, between rocks (B. B. Lowery); Whianj Whian S. F., 28.39°S, 153.20°E, rainforest, 200 m, under stone, ace. no. 16993 16.vi.1976 (P. Ward); Tomewin, rainforest. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 1500 ft, under stone (B. B. Lowery); Blue Knob Mt., Nightcap Ranges, rainforest, 2,800 to 3,000 ft (B. B. Lowery); Bilambil, N of Tumbulgum, rainforest (B. B. Low- ery); Hills above Tumbulgum, 800 ft, near Murwillumbah, rainforest (B. B. Lowery); Bonalbo, Sandy Crk., RF, 3000 ft, under rock (E. G. Kearney). Ecological Information. This species oc- curs in rainforest, nesting in the soil, usu- ally under or between rocks, often in a tan- gle of small plant roots; it probably re- stricts its foraging activity to the soil and leaf litter (Taylor, 1965, 1968). An additional 11 specimens, including nine workers, a queen, and an ergatoid queen, are examined here. They show the following some differences from the above “examined material”: (1) The ventral cen- ter of the clypeus possesses a_toothlike prominence in these 11 specimens, but a short transverse carina in the previous “Material Examined” section; (2) in the nine workers, the juncture between the pronotum and the mesonotum bears sev- eral short longitudinal rugae, varying from superficial to rather distinct (but is smooth and shining in P wheeleri); (3) in the queen, the petiole node in profile is wedge-shaped, and the propodeal spines (ca. 0.13) are shorter than those (ca. 0.20— 0.24) in P. wheeleri; and (4) in the ergatoid queen, only one ocellus is present; PSL1 and PSL2 are not shorter than 0.08 and 0.13, respectively (in P. wheeleri, three ocelli are distinct; PSL1 and PSL2 are ca. 0.18 and ca. 0.25, respectively). Further collecting and studying will help deter- mine whether these differences are signif- icant or not. These 11 specimens have the following measurements: Worker: TL 3.50—4.76, HL 0.90-1.05, HW 0.94—1.12, CI 104—108, SL Woo—l.01, SI 89=-93. EL 0.16—0.20, PW 0.60-0.68, AL 0.8S—0.96, PSL1 0.10—-0.14, ES2 0.16-0.20, PPW 0.24-029 PPL 0.21—0.23, PPI 104-126 (n = 9). Queen: Wi 3:84, Mi 1.22, HW 1.32) CI 108, SL Ov, SL 8, Bi 0:25; PW 0:74, AL. 1.18, PSL2 0.13, PPW 0.36, PPL 0.28, PPI 129 481 (a = IL), laigseicorel Giueeing INL, Sic4) Is0b, Oo, Jay IOs) (Gib Ike}, Sib; Oyetss, SIL UL EL 0.20, PW 0.64, AL 0.94, PSL1 0.08, IE SIERA (OS). IRIAN (O75) IRIEL, (OL PARAL JIS) (n = 1). Collecting Data for the 11 Specimens (ANIC). Australia: New South Wales, Tuckers Knob, 21 km SW Coffs. Hbr., rainforest, 760 m, ANIC Berlesate No. 201 (N. I. Mitchell); New South Wales, Bellan- gry, F'st, rainforest, ca. 3000 ft, iv.1958 (Darlingtons); New South Wales, Up. Al- lyn R., near Eccleston, 32.08°S, 151.29°E, rainforest, 400 m, acc. no. 316 (P. Ward): New South Wales, Upr. Allyn Val., near Eccleston, rainforest, ca. 2,000 ft, fallen epiphyte masses, ANIC Berlesate 45, 11— 14.xii.1967 (Taylor and Brooks); New South Wales, Comboyne plat., 2 to 2,800 ft, under rocks, x.1957 (Darlingtons). Pristomyrmex wilsoni Taylor Figures 193-194 Pristomyrmex wilsoni Taylor, 1968: 63. Holotype worker, Australia: N. Queensland, Mt. Lewis, ca. 3,000 ft, near Julatten, rainforest, ground strays, 30—31.x.1966 (R. W. Taylor) [ANIC (Taylor, 1968)]; [one paratype worker (MCZC) examined]. Diagnosis (Worker). Pronotal spines and propodeal spines exceptionally long (ca. 0.66—0.88). Worker. TL 4.14-4.92, HL 0.96-1.11, HW 0.96-1.12, Cl 96—104, SL 1.21—1.46, SM 2a sos OM9=02 2 PVWE0:65—0N76: AL Ada 3s05 PRPWr02S=0:32] PRE 030= ORIEL 9 300K — 3) Mandibles usually smooth and shining but sometimes with one to two longitudi- nal rugae. Masticatory margin of mandible with three teeth arranged as an apical + a preapical + a long diastema + a broad, truncated basal tooth. Basal margin of mandible lacking a distinctly curved lobe or tooth. Clypeus usually with a median longitudinal carina, but sometimes this median carina is interrupted or indistinct. Anterior clypeal margin with a median denticle and two others on each side, but sometimes two lateral denticles are fused into one prominence. Ventral surface of 482 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 193-194. Pristomyrmex wilsoni Taylor. 193A: Worker head, full-face view; 193B: Showing a ruga on the ventral clypeus; - 194: Worker, lateral view. clypeus with a short transverse carina. Palp formula 2,3. Frontal carinae short, not be- yond, or just reaching to the level of the posterior margins of eyes; sometimes fron- tal carinae absent Antennal scrobes ab- sent. Frontal lobes absent; thus, the anten- nal articulations are entirely exposed. An- tennal scapes long, when laid on the dorsal head, surpassing the occipital margin by about one-third of their length. Eyes con- taining 10 to 11 Ommnatichnk: in the longest row. Eronile shape of alitrunk and pedicel segments as in Figure 194. Pronotal spines exceptionally long (ca. 0.66—0.88), curved at their apices, diver ging toward the out- sides in dorsal view. Propodeal spines ex- ceptionally long (ca. 0.66—0.82), as illus- trated in Figure 194; in dorsal view, they are somewhat joined together at the base but are divergent posteriorly, Metapleural lobes small-triangular, each with an apex. Petiole with a fairly long anterior pedun- cle. Petiole node and postpetiole in profile higher than long, in dorsal view each with a somewhat aontenl apex. Postpetiole in dorsal view usually longer than broad, broadening from front to Thal Dorsum of head usually smooth and shining, except for a few foveolate punctures present on the genae and sometimes bordering the frontal carinae. Dorsum of alitrunk un-! sculptured and highly polished. Petiole, postpetiole, and gaster smooth and shine ing. Dorsal surfaces of head and alitrunk with numerous erect or suberect long. hairs. A pair of similar long hairs bilaterally distributed on the dorsum of petiole node and on the postpetiole, respectively. First gastral tergite lacking erect or suberect: hairs. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect to suberect short hairs. Color reddish- brown to blackish-brown. Queen and Male. Unknown. Comments and Discussion. Pristomyr- | mex wilsoni occurs only in North Queens-’ land, Australia. It is a unique species in the: genus that possesses both exceptionally: long pronotal and propodeal spines and. Thee can be immediately recognized. | Pristomyrmex wilsoni may abe derived from the common ancestor of the clade consisting of P. curvulus and P. longispinus of the Philippines. The workers of P. wil-) soni are similar to those of P. curvulus and. P. longispinus in possession of long pro-| notal spines and long antennal scapes and in the size and sculpture of body. Pristo-| myrmex wilsoni differs from P. curvulus: REVISION OF THE ANT GENUS PRISTOMYRMEX *® Wang and P. longispinus in the workers as fol- lows: P. wilsoni Propodeal spines exceptionally long, ca. 0.66 to 0.82; in dorsal view, they are close to each other at the base Maxillary palp with two segments Masticatory margin of mandible with a broad, truncated basal tooth Petiole node and postpetiole, in dorsal view each with a somewhat conical apex A pair of hairs present near the apex of petiole node P. curvulus and P. longispinus Propodeal spines short to moderately long, ca. 0.12 to 0.26; in dorsal view, they are separated at the base Maxillary palp with one segment Masticatory margin of mandible with two small basal teeth Petiole node and postpetiole in dorsal view each lacking a conical apex Two or more pairs of hairs present on the dorsal surface of petiole node An alternative is that P. wilsoni might evolved from the ancestor of the Austra- lian P. wheeleri because of the following characters being similar: (1) masticatory margin of mandible with three teeth, (2) maxillary palp with two segments, (3) body smooth, and (4) ventral surface of clypeus with a short transverse carina. However, the workers of P. wilsoni obviously differ from those of P. wheeleri as follows: P. wilsoni Pronotal and propodeal spines excep- tionally long (ca. 0.66—0.88) Antennal scapes relatively long (SL 1.21—1.46; SI 125-133) Petiole node and postpetiole in dorsal view each with a conical apex Dorsum of alitrunk unsculptured Labial palp with three segments P. wheeleri Pronotal and propodeal spines moder- ately long (ca. 0.12—0.28) 483 Antennal scapes relatively short (SL 0.94—1.18; SI 8S—100) Petiole node and postpetiole in dorsal view lacking a conical apex A transverse ridge present at the ap- proximate position of metanotal groove Labial palp with two segments Material Examined (ANIC, MCZC). Australia: N. Queensland, Mt. Lewis, 1,000 m, RF (R. W. Taylor); Queensland, Mt. Lewis, 960 m, 16.35°S, 145.17°E, rain- forest, acc. no. 76.349 (R. W. Taylor and T. A. Weir); NE Queensland, 2.5 km N Mt. Lewis via Julatten, 1,040 m, RF, Py- rethrum knockdown (D. K. Yeates and G. I. Thompson); N. Queensland, 2 km SE Mt. Spurgeon via Mt. Carbine, 1,100 m, Pyrethrum tree logs (Montelth and Thompson); NE Queensland, McDowall Ra, 17 km N Daintree, 16.06°S, 145.20°E, rainforest, 520 m, sieved litter, QM Ber- lesate No. 684 (G. Montelth); NE Queens- land, Table Mt., 10 km S of Cape Tribu- lation, 16.09°S, 145.26°E, rainforest, 320 m, sieved litter, QM Berlesate No. 542 (G. B. Montelth and D. Cook). Ecological Information. Taylor (1968) reports that this species occurs in rainfo- rest; it has been collected at an elevation of about 915 m in little berlesates and on the surfaces of rocks and logs. “Collections were made on two overcast days but not on a third, which was brightly sunny. Strays were not seen during several hours night collecting (to about four hours after sunset) at the same locality” (Taylor, 1968). THE TRISPINOSUS GROUP Worker. Medium-sized ants (HL 0.8S— JED OFSP ILS, Wi, B24) satin the following combination of characters: (1) Masticatory margin of mandibles with four teeth that have one of the fol- lowing two arrangements: a. the strongest apical + the second strongest preapical + a relatively short (first) diastema + a small den- ticle + a relatively long (second) di- 484 Figures 195-196. Pristomyrmex bispinosus (Donisthorpe). 195: Propodeal spines, dorsal view. astema + a small basal denticle, as in P. bispinosus and P. trispinosus (in P. trispinosus, sometimes the first di- astema is somewhat indistinct) or b. the apical + the preapical + a rela- tively long (first) diastema + a small denticle + a relatively short (second) diastema + a small basal denticle, as in P. browni; sometimes the second diastema is indistinct. (2) Anterior margin of the median por- tion of clypeus with at least five denticles. Two ends of the anterior clypeal margin each with a developed subtriangular tooth. Lateral portions of clypeus reduced to a margin; thus, the antennal fossae reach the anterior clypeal margin. (3) Palp formula 1,2. (4) Frontal carinae absent. (5) Frontal lobes almost completely ab- sent. (6) Antennal scrobes absent. (7) Lamella that encircles the base of antennal scape entire. (8) Dorsum of alitrunk with a prome- sonotal suture or impression. (9) Alitrunk in profile with a convex promesonotum and a deeply concave pro- podeal dorsum. (10) Pronotum with a pair of robust, short to moderately long spines. Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 | Worker head, full-face view; 196A: Worker, lateral view; 196B: | (11) Propodeal spines long, in dorsal ! view joined together at the base so that they form a fork. | (12) Petiole with a long anterior pedun- cle that is longer than the length of the node. | (13) Foveolate punctures completely absent, but regular striations are present on the dorsal surfaces of the head and the alitrunk in P. trispinosus and P. browni and present on the genae and around the an-_ tennal sockets in P. bispinosus. (14) First gastral tergite with numerous | hairs. This is a monophyletic group because it | possesses many autapomorphic characters, such as characters 3, 4, 8, 9, 11, and 13. This group contains three native Mauritian — species. Pristomyrmex bispinosus (Donisthorpe) Figures 195-196 Dodous bispinosus Donisthorpe, 1949: 272. Lecto- type worker, Mauritius: Le Pouce Mt., 2.xi.1948 (R. Mamet) (BMNH), here designated, [examined]. Pristomyrmex bispinosus (Donisthorpe) Brown, - 1971: 3. Diagnosis (Worker). Dorsal surfaces of | head and alitrunk mostly unsculptured; mesonotum unarmed, at most weakly tu- — berculate; alitrunk in profile with a convex | REVISION OF THE ANT GENUS PRISTOMYRMEX ° Wang promesonotum and a deeply concave pro- podeal dorsum; propodeal spines in dorsal view forming a divergent fork. Worker. TL 4.36—4.82, HL 1.14—1.20, IW 1.08—1.16, Cl 90=100, SL 1.32=—1.40, Sl 18=130), EL 0:22=0'245 PW 0:70=0.75; AL, 1.14—1.20; PPW 0:28-0:32, PPI 0.26— ODS TARO syC a Lib)e Mandibles smooth and shining. Masti- catory margin of mandible with four teeth arranged as the strongest apical + the sec- ond strongest preapical + a relatively short diastema + a small denticle + a relatively long diastema + a small basal denticle. Basal margin of mandible lacking a tooth- like prominence. Anterior margin of the median portion of clypeus with at least five denticles, but sometimes two lateral den- ticles are fused into a broad, truncated lobe. Two ends of anterior clypeal margin each with a strong, subtriangular tooth. Ventral surface of clypeus usually with a transverse ruga. Clypeus usually unsculp- tured, but frontal area usually with three to four short carinae that often extend to the posterior clypeal margin. Palp formula 1,2. Frontal carinae absent. Antennal scro- bes absent. Frontal lobes nearly complete- ly absent so that the antennal articulations are almost entirely exposed. Antennal scapes long, when lying on the dorsal head surpassing the occipital margin by about one-fourth of their length. ‘Eyes usually containing 11 to 12 ommatidia in the lon- gest row. Occipital margin slightly concave. Alitrunk in profile with a convex pro-me- sonotum and a deeply concave propodeal dorsum. Pronotum armed with a pair of fairly long, acute spines. Propodeum with a pair of long spines that, in dorsal view, are joined together at the base but are di- vergent along their length so that they form a fork (Fig. 196B). Mesonotum un- armed but sometimes with tubercles. Pro- mesonotal suture or impression present. Metapleural lobes dentiform. Petiole in profile with a long anterior peduncle; an- terodorsal angle of the node is higher than the posterodorsal. Postpetiole in profile usually with a curved anterior and dorsal 485 surface, in dorsal view slightly broader than long. Dorsum of head mostly smooth and shining but with some regular short rugae present around the antennal fossae, on genae and sometimes around the cen- trical disc. Dorsum of alitrunk as well as the sides of pronotum usually smooth and shining, but the sides of the rest of alitrunk usually with some regular short rugae. Pet- iole unsculptured. Dorsum of postpetiole smooth and shining. Gaster unsculptured. All dorsal surfaces with numerous erect or suberect hairs. A row of forward-project- ing hairs present near the anterior clypeal margin. Scapes and tibiae with some short hairs. Color yellow-brown, but sometimes reddish-brown. Queen and Male. Unknown. Comments. This species occurs in Maur- itius. It can be easily separated from its two relatives, P. trispinosus and P. browni, in the workers, as follows: In P. bispinosus, the dorsal surfaces of the head and the al- itrunk are smooth and shining, except for some short rugae around the atennal fos- sae, on the genae, and sometimes around the cenineal disc of the head: but in P trispinosus and P. browni, the dorsal sur- faces of the head and the alitrunk are en- tirely sculptured with regular striations. Material Examined (MCZC, ANIC, MNHN). Mauritius: Le Pouce Mt., 700 to SOO m, native forest (W. L. Brown); Le Pouce, 700 m, 20°12’S, 57°31'E, ex closed forest, on low vegetation (P. S. Ward); Le Pouce Mt. (Ray Mamet). Ecological Information. This species oc- curs in a forest and has been collected on the trees and on the main path (Brown, 1971). Pristomyrmex browni sp. n. Figures 197-198 Diagnosis (Worker). Dorsal surfaces of head and alitrunk with dense regular stri- ations; propodeal spines in dorsal view joined together at base and subparallel along their length; HW 0.82—0.90 and HL 0.88—1.01. 486 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 197-198. Pristomyrmex browni sp. n. 197: Worker head, full-face view; 198A: Worker, lateral view; 198B: Propodeal spines, dorsal view. Holotype Worker (MCZC). TL. 3.50, HL 0.92, HW 0.84, CI 91, SL 0.88, SI 105, PW 0.46, AL 0.90. Paratypes, 11 workers and one male (MCZC, ANIC, BMNH). Worker. TL 3.42-3.78, HL 0.92-1.00, HW 0.82-0.90, CI 87-94, SL 0.86—0.97, SIR9S= Ee Os14= O19 RW 04420550; AL 0.84—0.96, PPW 0.26—0.28, PPL 0.22— Oy IRIE IZ ae ip = ILL). Mandibles smooth and shining. Masti- catory margin of mandible with four teeth arranged as the strongest apical + the sec- ond strongest preapical + a relatively long (first) diastema + a small denticle + a rel- atively short (second) diastema + a small basal denticle: sometimes the second dia- stema indistinct. Basal margin of mandible almost straight, lacking a toothlike promi- nence. Anterior margin of the median por- tion of clypeus with at least five toothlike prominences, but sometimes two lateral denticles are fused into a broad lobe. Two ends of the anterior clypeal margin each with a strong subtriangular tooth. Ventral surface of clypeus lacking toothlike prom- inences. Clypeus usually with a few weak longitudinal rugae. Palp formula 1,2. Frontal carinae absent. Antennal scrobes absent. Frontal lobes nearly completely absent so that the antennal articulations are almost entirely exposed. Antennal scapes, when lying on the dorsal head, slightly surpassing the occipital margin by about one-eighth to one-seventh of their length. Eyes moderately sized. Occipital margin medially deeply emarginate. Ali- trunk in profile with a convex pro-meson- otum and a deeply concave propodeal dor- sum. Pronotum with a pair of acute short spines. Mesonotum lacking spines or teeth but usually with three blunt small tuber- | cles that are present on the posterior end | and on the two sides, respectively. Propo-- deal spines in profile well developed, long, bent at about a right angle near the base. In dorsal view, propodeal spines somewhat | laterally compressed; they are joined to- gether at the base but subparallel along_ their length (Fig. 198B). Metapleural | lobes small, triangular. Petiole node in | profile nodiform with a long anterior pe- | duncle; its anterodorsal angle is higher than the posterodorsal. Postpetiole in pro- | file with a single curved anterior and dor- | sal surface, in dorsal view slightly broader | than long and also broader than the petiole / node. Dorsum of head entirely covered | with regular coarse striations that consist of longitudinal rugae and a few rugae | around the antennal fossae. Sides and dor- | sum of alitrunk sculptured with coarse cir- cular striations evenly, but the center of REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang mesonotum with a few coarse longitudinal rugae. Sides of petiole and postpetiole usually with a few superficial rugae. Gaster smooth and shining. Dorsal surfaces of head, alitrunk, petiole, and postpetiole with some erect or suberect hairs. First gastral tergite with numerous recumbent hairs. A row of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with numerous suberect short hairs. Color reddish-brown. Queen. Unknown. Male. One paratype male, collected in Mauritius by W. L. Brown, was originally mounted together with a worker on the same pin: TL 3.04, HL 0.58, HW 0.48, CI So. OL) 0.20) SI 42, HWE 0:76, EL 0:38, PW 0.73, AL 1.08, PPW 0.24, PPL 0.20, RPT 120m = 1). Head, including the eyes, broader than long. Clypeus convex without a median longitudinal carina. Anterior clypeal mar- gin transverse. Frontal carinae absent. Scapes only slightly longer than the first funicular segments but distinctly shorter than the rest of the funicular segments. On the mesoscutum, notauli distinct, showing a V shape; parapsidal furrows very super- ficial. Scuto-scutellar sulcus with six nar- row ridges. Propodeum weakly tubercu- late, lacking teeth or spines. Metapleural lobes triangular. Middle and hind tibiae without any spurs. Petiole node in profile with a fairly long anterior peduncle; ante- rior face of the node, together with the dorsal surface of the peduncle, forming a declivity. Postpetiole in profile low and rounded dorsally and in dorsal view broad- er than long. Dorsum of head smooth and shining, but frontal area with a median longitudinal carina. Pronotum and mesos- cutum rather smooth and shining, except for those marked sutures, but mesoscutel- lum with some longitudinal and reticulate rugae. Petiole smooth and shining but with a longitudinal carina present on each side of the dorsal surface. Postpetiole and gas- ter smooth and shining. All dorsal surfaces with abundant erect or suberect hairs. Scapes and tibiae with numerous erect or 487 suberect short hairs. Color reddish-brown: wings slightly smoky. Comments. This new species occurs in Mauritius and Reunion Island. It is closely related to the Mauritian P. bispinosus and P. trispinosus. Characters separating P. browni from P. bispinosus and P. trispino- sus are provided under P. bispinosus and P. trispinosus, respectively. The following additional characters should also be men- tioned: In the workers of P. browni, the sculpture of the dorsal head consists most- ly of longitudinal rugae, except for a few rugae around the antennal fossae; the two sides of the mesonotum each has a small tubercle. In P. trispinosus, many striations present on the dorsal head spread out from the center to the two sides; the me- sonotum possesses a pair of strong digitlike prominences. In the type worker speci- mens of P. browni, the first gastral tergite possesses only recumbent hairs, and the occipital margin of the head in full-face view is deeply concave, which are different may sz bispinosus and P. trispinosus. One non-type specimen shows the fol- lowing variations: (1) The first gastral ter- gite possesses some erect or suberect hairs, (2) the occipital margin of the head in full-face view is feebly concave, (3) the sides of the mesonotum lack any tubercles, (4) the two small basal denticles on the masticatory margin of the mandible are in- distinct, (5) the dorsum of the petiole pe- duncle in profile is obviously curved, and (6) the rugae are very superficial on the clypeus and on the sides of the petiole. This specimen has the following measure- ments: TL 3.24, HL 0.88, HW 0.84, CI 95, SIO SOs SIN 9os HIE OMA ENV Ox4 05s AE: OM ave Holotype Worker. Mauritius: Le Pouce Mt., 700 to SOO m, native forest, 1.iv.1969 (W. L. Brown). Paratypes. Mauritius: Le Pouce Mt., 700 to 800 m, native forest, 4.iii. 1977 (W. L. Brown). Additional Non-Type Worker Examined. Indian Ocean, Reunion I., Mare Longue, 488 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 199-200. Pristomyrmex trispinosus (Donisthorpe). 199: Propodeal spines, dorsal view. near St. Philippe, primitive forest, 580 m, 15.i.1975 (D. Schauenberg) (MCZC). Ecological Information. This species oc- curs in native forest. Pristomyrmex trispinosus (Donisthorpe) Figures 199-200 Dodous trispinosus Donisthorpe, 1946: 145. Syntype workers and males, Mauritius: Cocotte Mt., 27.xii.1941 and 6.ii.1943 (R. Mamet) (BMNH, LACM, MCZC) [examined]. Pristomyrmex trispinosus (Donisthorpe) Brown, 1971: 3. Diagnosis ( Worker). Pronotum, meson- otum, and propodeum each with a pair of strong prominences; dorsal surfaces of head and alitrunk with dense regular stri- ations; propodeal spines in dorsal view di- vergent; HW and HL > 1.10. Worker. TL 4.50-4.68, HL 1.16—1.22, EIW. eS ess Cl Sl=97 Sie 133 18445 SI17/=122 EL 020201245, PW 0:66=0570: AL 1.16—1.26, PPW 0.27-0.30, PPL 0.24— 0:26) RPI 108=115 Ga — 6). Mandibles smooth and shining. Masti- catory margin of mandible with four teeth arranged as the strongest apical + the sec- ond strongest preapical + a short diastema (sometimes, this diastema is not distinct) + a small denticle + a long diastema + a small basal denticle. Basal margin of man- dible lacking a toothlike prominence. An- terior margin of the median portion of Worker head, full-face view; 200A: Worker, lateral view; 200B: | clypeus with at least five denticles, but sometimes two lateral denticles are fused into a broad lobe. Two ends of the anterior - clypeal margin each with a developed prominence. Ventral surface of clypeus lacking toothlike prominences. Clypeus— usually with a few longitudinal rugae. Palp— formula 1,2. Frontal carinae absent. An- tennal scrobes absent. Frontal lobes ab- | sent so that the antennal articulations are completely exposed. Antennal scapes long, - when lying on the dorsal head surpassing | the occipital margin by one-fourth to one-_ third of their length. Eyes usually contain-— fi ing 11 to 12 ommatidia in the longest row. | Occipital margin in full-face view slightly | concave. Alitrunk in profile with a convex pro-mesonotum and a deeply concave pro- podeal dorsum. Pronotum armed with a pair of moderately long acute spines. Me- sonotum with a pair of thick, blunt, digit- like short prominences. Propodeum with a_ pair of developed long spines that, in dor- sal view, are joined together at the base but divergent along their length so that they form a fork (Fig. 200B). Metapleural | lobes dentiform. Petiole node in profile | nodiform with a long anterior peduncle. i Postpetiole in profile rounded anterodor- sally but usually with a distinct postero- dorsal angle; in dorsal view, postpetiole - slightly broader than long. Entire dorsum — REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang of head with regular coarse striations: many striations spread out from the center to the two sides and to the occipital mar- gin; some are around the antennal fossae, the rest are some longitudinal rugae pre- sent on the sides of the head. Sides and the dorsum of alitrunk with numerous evenly distributed, circular coarse stria- tions. Center of mesonotum with a few coarse short rugae. Petiole and postpetiole rather smooth and shining, but sometimes their sides with a few superficial short ru- gae. Gaster unsculptured. All dorsal sur- faces with numerous erect or suberect hairs. Scapes and tibiae with numerous erect or suberect short hairs. A row of for- ward-projecting hairs present near the an- terior clypeal margin. Color yellow-brown, but sometimes reddish-brown. Queen. Unknown. Male. One syntype male (BMNH), to- gether with a number of syntype workers, constitutes a series (see Donisthorpe, 1946): TL 4.46, HL 0.72, HW 0.70, CI 97, SL 0.20, SI 29, EL 0.22, PW 0.90, AL 1.42 im = 1]. Head, including the eyes, broader than long. Clypeus somewhat transverse, con- vex in middle. On the mesoscutum, notauli indistinct. Scuto-scutellar sulcus wide, sep- arated into small cells by narrow ridges. Propodeum weakly tuberculate, lacking teeth or spines. Metapleural lobes sub- triangular. Petiole node in profile low with a fairly long anterior peduncle. Postpetiole in profile rounded dorsally. Dorsum of head smooth and shining. Mesoscutum smooth and shining, but mesoscutellum with some rugae and a few foveolate punc- tures. Petiole node rather smooth. Post- petiole and gaster unsculptured, smooth, and shining. All dorsal surfaces with abun- dant erect or suberect long hairs. Scapes and tibiae with numerous erect or sub- erect short hairs. Color reddish-brown: wings somewhat dusky. Comments. Pristomyrmex trispinosus is known only from Mauritius. It differs from P. bispinosus in the workers in having reg- ular coarse striations on the entire dorsal 489 surfaces of the head and the alitrunk and a pair of strong, digitlike prominences on the mesonotum. The workers of P. trispi- nosus and P. bispinosus are separable from those of P. browni by the following char- acters: P. trispinosus and P. bispinosus Propodeal spines in dorsal view diver- gent, not laterally compressed; in pro- file rather straight A relatively short diastema present be- tween the preapical and the third tooth on the masticatory margin of the mandible Larger species with HW > 1.08, HL > ae Sie oe NN OO Sale = 4.36 P. browni Propodeal spines in dorsal view subpar- allel, somewhat laterally compressed; in profile, bent at about a right-angle near the base A relatively long diastema present be- tween the preapical and the third tooth on the masticatory margin of the mandible Smaller species with HW 0.82-0.90, HL 0.88—1.01, SL 0.80—0.97, PW 0.40— ODO 324= 3.05 Material Examined (MCZC). Mauritius: Cocotte Mt. (R. Mamet). Ecological Information. A nest of P. tri- spinosus was found under a flat stone, and about 30 to 40 workers, two males, larvae, and pupae were collected; this species, when disturbed, simulates death (Donis- thorpe, 1946). THE LEVIGATUS GROUP Worker. This group shows the following combination of characters: (1) Usually small-sized species: HW: 0.60—0.96 in 11 species, 0.98—1.26 in one species (P. lucidus). HL: 0.60-0.90 in 11 species, 0.92—1.16 in P. lucidus. TL: 2.20- 3.49 in 11 species, 3.71—4.84 in P. lucidus. (2) Masticatory margin of mandible with four teeth arranged as the strongest apical + the second strongest preapical + 490 the smallest third + an acute basal tooth that is larger than the third tooth but smaller than the apical and preapical teeth; masticatory margin lacking a distinct dia- stema. (3) A tooth that is short and broad or prominent, present about midway on the basal margin of mandible. (4) Lateral portions of clypeus in front of antennal insertions reduced to a narrow margin. (5) Lamella that encircles the base of antennal scape entire. (6) Anterior clypeal margin with at most three denticles (i.e., a median den- ticle and one on each side), but sometimes the median tooth is indistinct so that only two denticles are seen on the margin. (7) Ventral surface of clypeus smooth or bearing a weak transverse ruga but lacking toothlike prominences. (8) Palp formula 1,3 in 10 species ex- amined. (9) Frontal carinae extending to the lev- el of the posterior margins of eyes. (10) Well-developed scrobes absent. (11) Dorsum of alitrunk in profile more or less arched, in dorsal view without any sutures. (12) Pronotum usually unarmed but with a pair of small teeth in one species (P. minusculus). (13) Petiole node in profile high, higher than the length of the node, usually with a distinct anterior face, but in P. inermis, the anterior face of the petiole node is in- separable from the dorsal surface of the peduncle. (14) Dorsal surfaces of head and ali- trunk smooth or with scattered foveolate punctures or with foveolate-reticulate sculpture. This is a monophyletic group, contain- ing 12 species. They are endemic in the Oriental region, except for one species (P. minusculus) entering in North Queens- land, Australia. Further, most of the spe- cies of this group occur in Papua New Guinea and some nearby islands. Obviously, this group, together with the Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 profundus group, constitutes a monophy- letic lineage because the workers and queens of the two groups possess a syna- | pomorphy, that is, the masticatory margin of the mandible with four teeth (the stron-— gest apical + the second strongest preap- ical + the smallest third + an acute basal | tooth) but lacking a diastema. The workers of the group are easily distinguished from those of the profundus group by lacking well-developed antennal scrobes and pos- sessing (1) two to three teeth on the an- terior clypeal margin, (2) a more or less | arched dorsum of the alitrunk, and (3) a tooth present about midway on the basal margin of the mandible but not adjacent to the basal tooth of the masticatory mar- gin. The dentition of the masticatory margin of the mandible in the workers and queens of the levigatus group is a critical character separating the levigatus group from other five (i.e., cribrarius, punctatus, quadri-— dens, and umbripennis) groups. In the workers and queens of the levi- gatus group, foveolate punctures show trispinosus, continuous variation on the dorsum of the | head between the frontal carinae. I treat this case as follows: (1) P. levigatus almost | completely lacks distinct foveolate punc- tures on the dorsum of the head between the frontal carinae; (2) P. simplex assem- bles those populations with some scattered foveolate punctures, but spaces between foveolae are usually smooth; and (3) P. cog- vii shows foveolate-reticulate sculpture; some populations, only with foveolate-re- ticulate sculpture behind the eyes, are considered intermediate forms and also grouped into P. coggii. In fact, this similar continuous variation also occurs in the oth- er two (i.e., guadridens and umbripennis) groups. An ergatoid queen caste has been found in P. mandibularis of the group. This caste — is also present in the punctatus and quad- ridens groups. Pronotal armaments, widely occurring in the cribrarius, quadridens, and trispinosus groups, appear in the - REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 49] Figures 201-202. Pristomyrmex acerosus sp. n. 201: Worker head, full-face view; 202: Worker, lateral view. workers of one species (P. minusculus) of the levigatus group. These facts indicate that ergatoid queens and pronotal arma- ments have arisen several times in Pristo- myrmex, respectively. Pristomyrmex acerosus sp. n. Figures 201-202 Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four teeth, of which the third tooth, counting from the apex, smallest; subpetiole with a pinlike long process. Holotype Worker (BMHH). TL. 2.80, HL 0.76, HW 0.75, CI 99, SL 0.66, SI 88, EL 0.12, PW 0.48, AL 0.76, PPW 0.22, REE OM. PPI W29) Mandibles smooth and shining. A broad-based short tooth present about midway on the basal margin of the man- dible. Clypeus depressed, unsculptured, its anterior margin with three denticles: a median tooth and one on each side. Fron- tal carinae just extending to the level of the posterior margins of eyes. Antennal scrobes absent. Frontal lobes weakly ex- panded basally. Eyes moderately sized. Occipital margin feebly concave in full- face view. Dorsum of alitrunk in profile convex. Pronotum unarmed. Propodeum with a pair of subtriangular short spines. Metapleural lobes rounded. Petiole node in profile high, with the anterodorsal angle higher than the posterodorsal, its anterior face subparallel to the posterior one, an- terior peduncle of the node about as long as the node. Subpetiole with a semitrans- lucent pinlike long process. Postpetiole with a rounded dorsum. In dorsal view, petiole node subrounded; postpetiole broader than long, somewhat transrectan- gular. Dorsal surfaces of head and alitrunk smooth and shining, except for some small, shallow hair pits. Petiole, postpetiole, and gaster unsculptured, smooth, and shining. Dorsal surface of head with numerous erect to suberect hairs. Dorsum of alitrunk with sparse hairs. Two pairs of hairs pre- sent on the dorsum of petiole node. A few on the dorsum of postpetiole and on the first gastral tergite, respectively. Three pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some suberect short hairs. Color lightly yellow-brown. Queen and Male. Unknown. Comments and Discussion. This is the only species in the genus that has so far been found to have a semitranslucent, pin- like, long process on the ventral surface of the petiole. However, further collecting is required. If this long process is present in other specimens, this holotype represents a good species; otherwise, this holotype 492 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 203-204. Pristomyrmex boltoni sp. n. 203: Worker head, full-face view; 204: Worker, lateral view. would be an aberrant specimen, and P. ac- erosus would become a junior synonym of P. levigatus. Holotype Worker. New Hebrides: Ma- lekoula I. (N), Vao Isl, 0 to 20 m, 7.ix.1979 (W.C. Gagne, G. M. Nishida, and G. A. Samuelson). Ecological Information. Unknown. Pristomyrmex boltoni sp. n. Figures 203-204 Diagnosis ( Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four teeth, of which the third tooth, counting from the apex, smallest; eyes very small, with two to three omma- tidia in the longest row; dorsal surfaces of head between frontal carinae and alitrunk smooth and shining. Holotype Worker (LAMN). TL 2.48, HL 0.66, HW 0.66, CI 100, SL 0.52, SI 79, EL 0.08, PW 0.42, AL 0.62, PPW 0.18, PPL 0.16, PPI 113. Paratypes, two workers (LAMN, MCZC). Worker. TL 2.40, 2.40; HL 0.65, 0.66; HW 0.66, 0.66; CI 100, 102; SL 0.55, 0.55: SI 83, 83; EL 0.08, 0.08; PW 0.44, 0.44; AL 0.60, 0.61; PPW 0.18, 0.18; PPL 0.16, (IS IPI WIE TLS Go =D). Mandibles smooth and shining but with few longitudinal rugae in the paratypes. A | broad-based short tooth present about — midway on the basal margin of the man- | dible. Frontal area concave with a median carina. Clypeus flat, unsculptured, smooth, — and shining, its anterior margin with two — lateral teeth, but in the two paratypes, an additional weak median tooth present. | Frontal carinae distinct, extending to the level of the posterior margins of eyes. Scrobal impressions shallow, present lat- | eral to the frontal carinae in full-face view. — Frontal lobes weak so that the antennal ar- — ticulations are almost entirely exposed. An- | tennal scapes, when lying on the head, | close to the occipital margin. Eyes very small, with two to three ommatidia in the longest row. Pronotum unarmed. Propo- | deum with a pair of triangular teeth. Me-_ tapleural lobes rounded. Petiole node in — profile high with the anterodorsal angle higher than the posterodorsal, its anterior — peduncle about as long as the node. Post- | petiole in profile higher than long with a rounded dorsum. In dorsal view, petiole — node subrounded, about as broad as long; postpetiole broader than long. Dorsum of — head between the frontal carinae smooth | and shining but with some foveolate punc- — tures present around the eyes. Dorsum of | REVISION OF THE ANT GENUS PRISTOMYRMEX ° Wang alitrunk smooth and shining. Petiole and postpetiole smooth and shining. Gaster unsculptured. Dorsal surfaces of head and alitrunk with numerous erect or suberect hairs. Two pairs of hairs present on the dorsum of petiole node, three pairs on the dorsum of postpetiole, and a few on the base of the first gastral tergite. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect or suberect short hairs. Color reddish-brown. Queen and Male. Unknown. Comments. Pristomyrmex boltoni is a sibling species of P. longus and P. coggii. The three species are all from New Guin- ea. Their workers possess very small eyes. Pristomyrmex boltoni differs from P. lon- gus and P. coggii as follows: The dorsum of the petiole node in dorsal view is slight- ly broader than long or about as broad as long in the workers of P. boltoni, but long- oval and distinctly longer than broad in P. longus. The dorsal surfaces of the head be- tween the frontal carinae and the alitrunk are smooth in the workers of P. boltoni but are covered with numerous foveolate punctures in P. coggii. The workers of P. boltoni are also similar in appearance to those of P. levigatus, but they can be separated by the following characters: P. boltoni Eyes smaller, with the maximum diam- eter 0.08, containing two to three om- matidia in the longest row P. levigatus Eyes larger, with the maximum diame- ter 0.12 to 0.16 (rarely 0.10), contain- ing five to seven ommatidia in the longest row Holotype Worker. New Guinea: Gulf Prov., Ivimka Camp, Lakekamu Basin, 7.73°S, 146.76°E, 120 m, #96-235, lowland wet forest, ex sifted leaf litter, 28.x.1996 (R. R. Snelling). Paratypes. Two workers, New Guinea: Gulf Prov., Ivimka Camp, Lakekamu Ba- sin, 7.7°S, 146.8°E, 140 m elevation, #96- 493 280, lowland wet forest, ex sifted leaf litter, 6.xi.1996 (R. R. Snelling). A non-type specimen (ANIC), collected in New Guinea (Brown R., lowland RF, under log) by B. B. Lowery, has the fol- lowing measurements: HW 0.66, HL 0.66, SL 0.56, EL 0.08, PW 0.44, AL 0.60. Ecological Information. This species oc- curs in lowland forest and has been col- lected in litter samples. Pristomyrmex coggii Emery Figures 205-208 Pristomyrmex coggii Emery, 1897: 584. Lectotype worker, New Guinea: Montes Hansemanni et Ber- linhafen (Bird) (MCSN), here designated, [exam- ined]. Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four teeth, of which the third tooth, counting from the apex, smallest; eyes with two to four ommatidia in the longest row; dorsal surface of head, except for the scrobal areas, with foveolate-retic- ulate sculpture; dorsum of petiole node in dorsal view about as long as broad or slightly broader than long; first gastral ter- gite with only a few hairs. Worker. TL 2.20-2.86, HL 0.60-0.74, HW 0.60-0.76, CI 98-104, SL 0.48—0.64, SI 74-86, EL 0.05—0.09, PW 0.40-0.48, AL 0.60-0.77, PPW 0.16—0.20, PPL 0.12— Of SRP aS Sr33e(n— 2.0). Mandibles usually smooth and shining but sometimes with a few superficial lon- gitudinal rugae. A broad and short or strongly prominent tooth present about midway on the basal margin of mandible. Clypeus flat, its anterior margin sometimes with three denticles: a median denticle and one on each side, but sometimes the median denticle absent or weak so that only two teeth are present there. Frontal area concave, with a median carina that usually extends to the clypeus. Ventral center of clypeus lacking any rugae or prominences. Palp formula 1,3. Frontal ca- rinae distinct, extending to the level of the posterior margins of eyes. Scrobal impres- sions smooth and shallow, present lateral 494 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 205-208. Pristomyrmex coggii Emery. 205: Worker head, full-face view; 206: Worker, lateral view; 207: Queen head, full-face view; 208: Queen, lateral view. to the frontal carinae. Frontal lobes weakly expanded so that the antennal articulations are almost completely exposed. Eyes small, usually with two to three, rarely four, ommatidia in the longest row. Occip- ital margin straight or feebly concave in full-face view. Pronotum unarmed. Pro- podeum with a pair of triangular short spines. Metapleural lobes rounded. Petiole node in profile high, with the anterodorsal angle higher than the posterodorsal, its an- terior peduncle about as long as the node. In dorsal view, dorsum of petiole node subrounded, about as long as broad, or transoval, slightly broader than long. Sub- petiole with a narrow, long, semitranslu- cent lamella. Postpetiole in profile round- ed dorsally, in dorsal view somewhat trans- | verse-rectangular and broader than long. — Dorsum of head, except for the scrobal ar- _ eas, with dense foveolate punctures that | form foveolate-reticulate sculpture; some- _ times the punctures are almost aligned so — that it seems that the several longitudinal — rugae appear between the frontal carinae. Dorsum of alitrunk with sparse foveolate _ punctures. Petiole, postpetiole, and gaster — unsculptured, smooth, and shining. Dorsal _ surfaces of head and alitrunk with numer- ous erect to suberect hairs. Dorsal surfaces of petiole node and _ postpetiole usually — with two pairs of hairs, respectively. A few | REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 495 hairs present on the base of the first gastral tergite. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with numerous erect to suberect short hairs. Color red- dish-brown. Queen. TL 2.94-3.16, HL 0.72-0.78, HW 0.72-0.82, CI 100—105, SL 0.56—0.66, Si Sone en Ola —OslivmueNViOlaa—OKO4- ANlEnOM 80.96, BPW 021-024" PPL.OllW— OalSs PPL 1222133 (n= 3). Generally similar to worker, except for normal caste differences. In addition, fo- veolate punctures shallow on the meson- otum, propodeal armaments slightly short- er than those in conspecific worker. Male. Unknown. Comments and Discussion. Pristomyr- mex coggii is closely related to P. boltoni and P. longs. The three species occur in New Guinea. Characters separating P. cog- gii from P. boltoni are provided unde: fie latter name. Pristomyrmex coggii differs from P. longus because the deisuin of the petiole Roden in dorsal view is about as long as broad or broader than long in the wor ia ers of P. coggii but longer than broad in P. longus. Pristomyrmex coggii differs from P. obe- sus of Solomon Islands as follows: The workers of P. coggii possess only a few hairs on the first gastral tergite and have smaller eyes containing two to three, rarely four, ommatidia in the longest row. But in the workers of PB. obesus, the entire first gastral tergite is evenly covered with erect or suberect hairs, and the eyes contain five to seven (rarely four) ommatidia in the longest row. Pristomyrmex coggii differ from P. simplex of New Guinea and the Philippines because the dorsum of the head between the frontal carinae bears fo- veolate-reticulate sculpture in the workers of P. coggii but only scattered foveolate punctures in P. simplex; in addition, the eyes usually contain five ommatidia in the longest row in the workers of P. simplex Pit - usually two to three in P. coggii. It must be pointed out that then material I have examined may resolve into two spe- cies with further study. Additional collect- ing will help clarify the situation. Material Examined (MCZC, ANIC, LAMN, BMNH, NHMYV). Papua New Guinea: Gulf Prov., Ivimka Camp, Lake- kamu Basin, lowland wet forest, #96-291 @273"S. TA6./67E) 120) msex sifted leafilit= ter), #96-350 (7.7°S, 146.8°E, 400 m ele- vation, sifted leaf litter and debris from rotten log) (R. R. Snelling); W. Highlands, Baiyer R., 4,000 ft (S. Peck); Rinona (R. Vane-Wright); New Guinea (Bird); Nad- zab, dry evergreen forest (E. O. Wilson); Huon Pen., lower Busu R., lowland rain- forest (E. O. Wilson); Wau, Kilolo Creek (J. Balogh); Lae, rainforest, leaf mold (T. E. Woodward); Lae, Busu R., lowland rainforest, in rotten wood on ground (B B. Lowery); NW Lae, “Timber Track” ca. 16 km, rainforest, ca. 220 m, berlesate (R. W. Taylor); near Popondetta, <50 m (R. W. Taylor); 8 km S of Kokoda, 800 m, rain- forest (R. W. Taylor). Ecological Information. This species oc- curs in rainforest and has been collected in litter samples and in rotten wood on the ground. Note: I have examined additional 25 specimens, including 19 workers and 6 queens. In these workers, the eyes are slightly larger than those in P. coggii; they contain four to five ommatidia in the lon- gest row. Further collecting and studying are needed. These 25 specimens have the following measurements: Worker: TL 2.76—3.00, HL 0.73—0.80, HW 0.73-0.82, CI 100-106, SL 0.58-0.64, SI 74-82, EL 0.09-0.10, PW 0.48—0.52, AL 0.68—0.82 (n = 19). Queen: 2) 92235345) Elle OM2—O0FS 0S EIN OM/2= 0.82, CI 100-103, SL 0.56—0.62, SI 73-82, I CLIO IEW O50. G2. ALL O.SO= 0.92 (n = 6). Collecting Data for the 25 Specimens (WI GAG: ANIC. BMNH). New Guinea: Morobe Dist., Bulolo (B. B. Lowery); Bul- olo, rainforest, 3,500 ft (B. B. Lowery); NE New Guinea, 6 km NE of Wau, Bulolo R. valley, rainforest, 1,100 m, berlesate, leaf mold (R. W. Taylor); Wau, Bishop Mu- 496 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 (2 KM Figures 209-212. Pristomyrmex inermis sp. n. 209: Worker head, full-face view; 210: Worker, lateral view; 211: Queen head, full-face view; 212: Queen, lateral view. seum Station, 1,200 m, rainforest, rotten log (R. W. Taylor); Wau, 4,000 ft, forest litter (S. Peck); N. Wau, on Bulolo Rd., 650 m (S. Peck); Wau, Kunai Creek, rain- forest, ca. 1,400 m, berlesate (R. W. Tay- lor); Tapini, 1,000 to 1,200 m, rainforest, rotten log (R. W. Taylor). Pristomyrmex inermis sp. n. Figures 209-212 Diagnosis (Worker). Propodeum lacking a pair of teeth or spines. Holotype Worker (LAMN). TL 2.86, HL 0.72, HW 0.70, CI 97, SL 0.64, SI 91, EL 0.14, PW 0.50, AL 0.72. Paratypes, 17 workers and three queens (ANIC, LACM, MCZC, BMNH). Worker. TL 2.65-—3.36, HL 0.70-0.84, HW 0.69-0.84, CI 96-103, SL 0.62—0.76, SI 83-92, EL 0.12-0.16, PW 0.46-0.58, Al? 0.63-0:80; PPW 0:23-027 Reb Ove O21 PRI 12821425(n— ls) Mandibles smooth and shining but sometimes with a few superficial small punctures. Masticatory margin of mandi- ble with four teeth arranged as the stron- gest apical + the second strongest preap- ical + the smallest third + a basal tooth; a distinct diastema lacking. A broad-based triangular short tooth present about mid- way on the basal margin of mandible. Frontal area concave. Clypeus flat, smooth, and shining, its anterior margin with three denticles: a weak median tooth REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang and one on each side, but sometimes the median tooth indistinct or lacking so that only two teeth are present there. Ventral surface of clypeus lacking any distinct ru- gae or teeth. Palp for vile 1,3. Frontal ca- rinae just extending to the ewe of the pos- terior margins of fhe eyes. Antennal scro- bes absent. Frontal lobes very weak. Eyes moderately sized. Occipital margin feebly convex in full face view. Dorsum of ali- trunk in profile arched. Pronotum un- armed. Propodeum lacking a pair of teeth or spines but usually with a pair of blunt small tubercles. Metapleural lobes small, usually bluntly rounded, but sometimes toothlike. Petiole node in profile wedge- shaped, usually with a blunt triangular apex; sometimes the apex somewhat blunt- ly rounded. Subpetiole with a narrow rim. Postpetiole in profile slightly higher than the petiole node, with a convex dorsum. In dorsal view, petiole node and postpeti- ole broader than long. Cephalic dorsum between the frontal carinae highly pol- ished but usually with a few foveolate punctures bordering frontal carinae. A few foveolate punctures present on the genae and many on the ventral surface of the head. Dorsum of alitrunk, petiole, and postpetiole smooth and shining. Gaster unsculptured. Dorsal surfaces of head, al- itrunk, petiole, and postpetiole with sparse erect to suberect hairs. A few hairs present on the first gastral tergite. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some suberect hairs. Color varying from light yellow-brown to blackish- brown, gaster usually darker than alitrunk; sometimes alitrunk lighter than head and gaster so that the specimens show a bicol- ored form. Queen. TL 3.23-3.52, HL 0.80-0.81, HW 0.76—0.85, CI 95-106, SL 0.66-—0.74, SI 87-92, EL 0.18—0.20, PW 0.64—0.72, AL 0.86—0.94, PPW 0.26—0.30, PPL 0.18— O24 PPI 125-144 (m= 3): General shape as in Figures 211-212; except for normal caste differences, other characters similar to worker. 497 Male. Unknown. Comments. This species can be easily recognized because it lacks a pair of pro- podeal teeth or spines in the workers, which is unique within Pristomyrmex. In addition, in the levigatus group, P. inermis has a characteristic shape of the petiole node in the workers and queens: wedge- like in profile view with an apex. This char- acter has originated independently at least twice in the genus because it is also seen in P. fossulatus and P. punctatus of the punctatus group. Holotype Worker. Indonesia: Irian Jaya, PT. Freeport Concession, Siewa Camp., 03.04°S 136.38°E, 200 ft, lowland second- ary rainforest, #98-71 (stray foragers in leaf litter), 12.iv.1998 (R. R. Snelling). Paratypes. One queen with same data as holotype; one queen with same data as ho- lotype but date 7.iv.1998, and number #98-48 (stray on log); two workers with same data as holotype but date 7.iv.1998 and number #98-44 (ex sifted leaf litter); three workers, Papua New Guinea: Bulolo, 2.300 ft, rainforest, 19.xii. 1967 (B. B. Low- ery); three workers, Papua New Guinea: Bulolo, 2,300 ft, rainforest, 27.xii.1967 (B. By Lowery); three workers, Papua New Guinea: Bulolo, 2,600 ft, rainforest, 25.xii.1970 (B. B. Lowery); one worker, Papua New Guinea: Bulolo, 2,800 ft, rain- forest, Ji197l, (Be By Lowery); :thnee workers, Papua New Guinea: Bulolo, 3,000 ft, rainforest, 5.i1.1971 (B. B. Low- ery); one queen, New Guinea: Bulolo (Morobe Dist), 8.i.1971 (B. B. Lowery); two workers, New Guinea: near Vanimo, rainforest, ca. 50 m, ex rotting log, 10— 11.vii.1972 (R.W.T.). Ecological Information. This species oc- curs in rainforest and has been collected in litter samples and on logs. Pristomyrmex largus sp. n. Figures 213-214 Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four ea, of which the third foot counting from the apex, smallest: 495 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 : | Figures 213-214. Pristomyrmex largus sp. n. 213: Worker head, full-face view; 214: Worker, lateral view. HW and HL = 0.90; petiole node in pro- file with the anterodorsal angle higher than the posterodorsal. Holotype Worker (ANIC). TL 3.38, HL 0.90, HW 0.96, CI 107, SL 0.78, SI 81, EL 0.14, PW 0.61, AL 0.92, PPW 0.24, PPL 0.20, PPI 120. Paratypes, 34 workers and one queen (MCZC, R. Clouse’s per- sonal collection). Worker. TL. 3.40, HL 0.90, HW 0.90, CI 100, SL 0.81, SI 90, EL 0.14, PW 0.60, AL 0.90, PPW 0.25, PPL 0.20, PPI 125 (n = jl). Mandibles smooth and shining. A broad-based triangular short tooth present about midway on the basal margin of the mandible. Clypeus flat, unsculptured, smooth, and shining; its anterior margin with three denticles: a weak median tooth and one prominent lateral tooth on each side, sometimes the median tooth indis- tinct. Ventral surface of clypeus unsculp- tured and smooth. Palp formula 1,3. Fron- tal carinae just extending to the level of the posterior margins of eyes. Antennal scrobes absent. Frontal lobes weakly ex- panded. Eyes moderately sized. Occipital margin straight or feebly concave in full- face view. Pronotum unarmed. Propo- deum with a pair of long spines that are about 1.5 times the distance between their bases. Metapleural lobes subtriangular. Dorsum of alitrunk in dorsal view rather flat. Petiole node in profile high, with the anterodorsal angle higher than the poster- odorsal, its anterior face sometimes sub- parallel to the posterior one; anterior pe- duncle of the node about as long as the node, and subpetiole with a narrow semi- translucent rim. In dorsal view, petiole node transoval. Postpetiole in profile high- er than long, rounded dorsally, in dorsal view broader than long and somewhat transrectangular. Both dorsal and ventral surfaces of head smooth and shining but with few small, shallow punctures present on the genae. Dorsum of alitrunk smooth and shining. Petiole and _ postpetiole smooth and shining, except for a lateral longitudinal carina on each side that sep- arates the tergite from the sternite. Gaster unsculptured. Dorsal surfaces of head and alitrunk with some erect to suberect short hairs. Petiole node and postpetiole each with two pairs of hairs as illustrated in Fig- ure 214. A few hairs present near the base of the first gastral tergite. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with abundant erect to suberect short hairs. Color yellow-brown to reddish- brown. Queen. TL 3.94, HL 0.94, HW 0.95, CI 101, SL 0.80, SI 84, EL 0.22, PW 0.74, REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang AIG, TOI TA PWWEO BO). IPP O28) Ie ISO) a = iDy Generally similar to worker, except for normal caste differences; in addition, pro- podeum with a pair of short spines, shorter than those in the conspecific worker, first gaster tergite with numerous hairs. Male. Gulonowns Comments. Pristomyrmex largus is the second largest species in the levigatus group after P. lucidus. Pristomyrmex lar- gus must have evolved from the ancestor of P. levigatus. Pristomyrmex largus can be separated from P. levigatus by ie follow- ing characters in the vores: P. largus Larger species with HW = 0.90, and HL = 0.90 Propodeal spines relatively long, longer than the distance between en bases Ventral surface of head smooth, with only a few small hair pits P. levigatus Smaller species with HW < 0.80, and HL < 0.80 Propodeal armaments relatively short, shorter than or about as long as the distance between their bases Ventral surface of head with numerous foveolate punctures Holotype Worker. Ponape I., Mt. Tolen- kiup; vi-ix.50 (P. A. Adams). Paratypes. One worker, Micronesia: Pohnpei I., Hilltop campsite near Mt. Nanalaud, 400 m, on an old ivory nut, 3.v.1995 (Ron Clouse); 33 workers and one queen, Pohnpei I., Nahnal aud cave, around camp (#155—-158, under dead leaves, on ground; #161, under rotting leaves; #167, under dead leaves. gl iie= 180. [eaves, ander rotten), 24.i1ii.2000 (Ron Clouse). Ecological Information. All par atypes of this species have been collected in high- elevation rainforest, under the rain- eoalked leaf litter, and inside a rotten ivory nut on the forest floor (R. Clouse, personal com- munication). Note: I do not illustrate the queen of 499 this species because this caste, together with 33 workers, collected by Mr. Ronald Clouse, reached me after my manuscript was completed. Pristomyrmex levigatus Emery Figures 215-218, 277, 281 Pristomyrmex levigatus Emery, 1897: 583. Lectotype worker, New Guinea: near Friedrich-Wilhelmshaf- en et Berlinhafen (Biré) (MCSN), here designated, [examined]. Pristomyrmex mendanai Mann, 1919: 341. Two syn- type workers, Solomon Is.: Santa Cruz Is., Graciosa Bay, 19.v.-24.xi.1916 (W. M. Mann) (MCZC, USNM) [examined]. Syn. n. Note: The lectotype of P. levigatus shows some fine differences from the two syntypes of P. mendanai. In the former, the metapleural lobes are subtriangular; the dorsum of the alitrunk is rather flat, with two sides somewhat margined; the pro- podeum is armed with a pair of short spines that are not subtriangular but some- what incurved at their apices. In the latter, the metapleural lobes are rounded; the dorsum of the alitrunk is convex, and its sides are not distinctly margined; the pro- podeal armaments are subtriangular, not distinctly incurved at their apices. How- ever, the presence of some intermediate forms suggests that they belong to the same species for the present. Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four teeth, of which the third tooth, counting from the apex, smallest; HW 0.62-0. 78, HL 0.64—0.78; eyes with five to eight ommatidia in the longest row; pronotum unarmed; propodeum with a pair of teeth or short spines; petiole with a lateral longitudinal carina on each side; subpetiole lacking a pinlike long process; dorsal surfaces of head and _ alitrunk smooth and shining. Worker. TL 2 36-2 .96, HL 0.64—0.78, HW 0.62-0.78, CI 96-106, SL 0.50—0.65, SI 78—89, EL 0.12—0.16 (very rarely 0.10), PW 0.41-—0.52, AL 0.54—0.75, PPW 0.20— O22 RR ON4=OMSaeRe i223 1435 ae 30). 500 217 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 ‘ons N Figures 215-218. Pristomyrmex levigatus Emery. 215: Worker head, full-face view; 216: Worker, lateral view; 217: Queen head, full-face view; 218: Queen, lateral view. Mandibles usually smooth and shining but with a few fine longitudinal rugae in some specimens. A broad-based triangular short tooth present about midway on the basal margin of the mandible. Frontal area concave, usually unsculptured, but some- times with a weak median carina. Clypeus flat, unsculptured, smooth, and shining; its anterior margin with a median denticle and two lateral teeth, but sometimes the median tooth is weak or absent. Ventral surface of clypeus lacking any rugae or toothlike prominences. Palp formula 1,3. Frontal carinae distinct, extending to the level of the posterior margins of eyes. Scrobal impressions shallow, present lat- eral to the frontal carinae. Frontal lobes weak so that the antennal articulations are almost entirely exposed. Antennal scapes, when lying on the head, close to the oc- cipital margin. Eyes moderately sized, with five to eight ommatidia in the longest row. Dorsum of alitrunk in dorsal view flat in the lectotype but convex in some speci- mens. Pronotum unarmed or at most with blunt tubercles, lacking teeth or spines. Propodeum with a pair of triangular teeth or short spines that are slender, slightly in- curved at their apices in the lectotype. Me- tapleural lobes subtriangular or rounded. Petiole node in profile nodiform with the anterodorsal angle higher than the poster- odorsal; its anterior peduncle about as long as the node. Subpetiole with a narrow long flange. In dorsal view, the dorsum of pet- iole node transoval and broader than long, Postpetiole in profile higher than long, with a rounded dorsum, in dorsal view REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang somewhat transverse-rectangular and broader than long. Dorsum of head smooth and shining but sometimes with a few foveolate punctures bordering the frontal carinae; sometimes a few foveolate punctures present on the genae and around the eyes. Dorsum of alitrunk un- sculptured, smooth, and shining. Petiole and postpetiole smooth and shining, each with a longitudinal carina on each side that separates the tergite from the sternite. Gaster unsculptured. Dorsum of head with numerous erect to suberect hairs. Dorsum of alitrunk with some erect or suberect hairs. Dorsal surfaces of petiole node and postpetiole usually with one to two pairs and one to three pairs of hairs, respectively. A few hairs present at the base of the first gastral tergite. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with erect to suberect short hairs. Color golden-brown or reddish-brown. Queen. TL 2.92-3.34, HL 0.74—0.75, HW 0.76—0.81, CI 101—108, SL 0.60—0.65, SI 79-83, EL 0.18—0.19, PW 0.54—0.66, AL 0.80—0.92, PPW 0.23-0.26, PPL 0.18— 0.20) PPI'128=133 @ = 5). Generally similar to worker, except for normal caste differences. Propodeal ar- maments tuberculate or denticulate, slightly shorter than those in the conspe- cific worker. Male (Figs. 277, 281). Two male speci- mens, together with 13 workers and two queens, collected in New Guinea (Nadzab, #1083) by E. O. Wilson, constitute a se- ries; each of the two males was originally mounted together with a worker or with a worker and a queen on the same pin: TL 2.48, 2.54; HL 0.50, 0.51; HW 0.53, 0.53; 1 105;,° 106: SL 0:16, 0.18: SI 30, 34: HWE 0.63, 0.64; EL 0.23, 0.25; PW 0.60, 0.63; AL 0.82, 0.84: PPW 0.18, 0.19; PPL 0.15, 0.16; PPI 119, 120 (n = 2). Head, including the eyes, broader than long. Clypeus lacking a median longitudi- nal carina; anterior clypeal margin rather straight. Frontal carinae reaching the level of the posterior margins of antennal inser- 501 tions. Mesoscutum lacking distinct notauli and parapsidal furrows. Scuto-scutellar sulcus with nine narrow longitudinal ridg- es. Propodeum weakly tuberculate, lacking teeth or spines. Metapleural lobes with a blunt or somewhat rounded apex. Middle and hind tibiae without any spurs. Petiole node in profile low, nodiform, having an anterior face and a long anterior peduncle. Postpetiole in profile low and rounded dorsally, in dorsal view subrectangular and broader than long. Dorsum of head smooth and shining, but frontal area with a median longitudinal carina. Dorsal ali- trunk generally smooth and shining, except for those marked sutures. Dorsal surfaces of petiole and postpetiole smooth and shining. Gaster unsculptured. All dorsal surfaces with abundant rather long hairs. Legs and scapes with numerous erect or suberect short hairs. Color blackish- brown. Wings infuscated. Comments and Discussion. Pristomyr- mex levigatus occurs in New Guinea, Nama Is., Solomon Is., New Georgia, and New Britain Is. It appears to be a basal species within the levigatus group. Many species, such as P. acerosus, P. boltoni, P. inermis, P. largus, P. lucidus, P. mandibu- laris, P. minusculus, P. obesus, and P. sim- plex, may have evolved from a P. levigatus— like ancestor. The workers of these species are separable from those of P. levigatus as follows: P. levigatus differs from both P. simplex and P. obesus by lacking foveolate punctures on the dorsal surfaces of the al- itrunk and the head between the frontal carinae. In P. levigatus, the eyes are larger, usually containing five to eight ommatidia in the longest row but two to three in P. boltoni. Pristomyrmex levigatus is smaller (HW < 0.80, HL < 0.80) than P. largus and P. lucidus (HW > 0.90, HL > 0. 90). Pristomyrmex minusculus bears a pair of pronotal teeth that are not seen in P. lev- igatus. A pair of propodeal teeth or short spines are present in P. levigatus, but ab- sent in P. inermis. Pristomyrmex levigatus possesses a longitudinal ruga on each side of the petiole node that is not seen in P. 502 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 219-220. Pristomyrmex longus sp. n. 219: Worker head, full-face view; 220A: Showing the dorsum of the petiole node of the worker, in dorsal view, is long-oval and longer than broad; 220B: Worker, lateral view. mandibularis. Finally, the subpetiole of P. levigatus does not have a pinlike long pro- cess that is distinct in P. acerosus. Material Examined (ANIC, BMNH, LAMN, MCZC). New Guinea: Nadzab, #1083, dry evergreen forest (E. O. Wil- son); Gogol Val. ca. 24 km W. Madang, ca. 50 m, rainforest, rotten wood, ex small fragment (R. W. Taylor); Bulolo, rainforest, 2,300 ft (B. B. Lowery); Yawasora near Wewak, ca. 50 m, rainforest, berlesate (R. W. Taylor); Gulf Prov., Ivimka Camp, Lak- ekamu Basin, 7.73°S, 146.76°E, 110 m, #96-345 (R. R. Snelling); Port Moresby, Brown River (J. Baloph); N.D. Papua, Sangara (G. Baker); (P. M. Room). Nama Is. near Truk (R. W. L. Potts). Solomon Is.: New Georgia (E. S. Brown). New Britain Is. (L. Weatherill). Ecological Information. This species oc- curs in rainforest and has been collected in a litter sample. Pristomyrmex longus sp. n. Figures 219-220 Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four teeth, of which the third tooth, counting from the apex, smallest; eyes small, with only two to three omma- tidia in the longest row; dorsum of petiole node in dorsal view long-oval and longer than broad; first gastral tergite with nu- merous hairs. Holotype Worker (MCZC). TL 2.51, HL 0.67, HW 0.66, CI 99, SL 0.55, SI 83, EL 0.07, PW 0.44, AL 0.66. Paratypes, 14 workers (MCZC, ANIC, BMNH). Worker. TL 2.22-2.68, HL 0.63—0.70; HW 0.62-0.70, CI 95—100, SL 0.52—0.56, SI 80-87, EL 0.05—0.10, PW 0.40-0.46 AL 0.60—0.70, PPW 0.17—0.18, PPL 0.16— O16, PP 041 35G sla)» Mandibles usually smooth and shining. A broad-based short tooth present about midway on the basal margin of the man- dible. Clypeus flat, its anterior margin with a median tooth and two lateral teeth; sometimes the median tooth is weak. Frontal area concave, with a median carina that usually extends to the clypeus. Ventral surface of clypeus smooth, lacking any ru- gae or prominences. Palp formula 1,3. Frontal carinae distinct, extending to the level of the posterior margins of eyes. Scrobal impressions shallow, smooth, pre- sent lateral to the frontal carinae. Frontal lobes weakly expanded. Antennal scapes, when lying on the head, close to the oc- cipital margin. Eyes very small, consisting of 4 to 10 ommatidia, with only two to three ommatidia in the longest row. Oc- cipital margin straight or feebly concave in full-face view. Profile of alitrunk and ped- REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang icel segments as in Figure tum unarmed. Propodeum with a pair of triangular teeth. Metapleural lobes round- ed. Betiole node in profile high with the anterodorsal angle higher than the poster- odorsal, its anterior fice subparallel to the posterior one; anterior peduncle of the node about as long as or slightly shorter than the node; subpetiole with a narrow semitranslucent lamella. In dorsal view, dorsum of petiole node long-oval and dis- tinctly longer than broad. ‘Postpetiole in rofile rounded dorsally, in dorsal view slightly broader than long or about as long as broad. Dorsum of oad between the frontal carinae mostly smooth but usually with some sparse, small, and shallow punc- tures. Some foveolate punctures present on the genae and around the occipital cor- ners of Mcadk Dorsum of alitrunk, petiole, postpetiole, and gaster usually unsculp- tured, smooth, anal shining. Dorsal surfac- es of head, alitrunk, and gaster with nu- merous erect to suberect haus. ‘Two pairs of hairs usually present on the dorsal sur- faces of petiole node and postpetiole, re- spectively, as illustrated in Figure 220B. A few pairs of forward-projecting hairs pres- ent near the anterior clypeal margin. Scapes and tibiae with numerous erect to suberect short hairs. Color yellow-brown to reddish-brown. Queen and Male. Unknown. Comments. The workers of P. longus are quickly recognizable by the following: In the levigatus group, only three species, P. Polioni.: P. coggii, and P. longus, possess small eyes. Only two species, P. longus and P. obesus, have numerous hairs evenly dis- tributed on the entire first gastral tergite. But in only one species (P. onan) is ahve dorsum of the petiole node in dorsal view long-oval and longer than broad; in the other members of the levigatus group, the dorsum of petiole node is subrounded or transverse-oval (i.e., about as long as broad or broader than long). Holotype Worker. New Guinea: Huon Pen., Lower Busu R., lowl. rainfor., 0.V.1955, #957 (E. O. Wilson). 220B. Prono- 503 Paratypes. Eight workers with same data as holotype; one worker, New Guinea: Huon Pen., Lower Busu R., lowland rain- forest, 6.v.1955, #978 (E. O. Wilson); five workers, New Guinea: 13 km NW Lae, Bubia, lowland rainforest, 26.iii.1955, #688 (E. O. Wilson). Ecological Information. This species has been collected in lowland rainforest. Pristomyrmex lucidus Emery Figures 221-224 Pristomyrmex lucidus Emery, 1897: 584. Holotype worker, New Guinea: Berlinhafen (Bird) (MCSN) [examined]. Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four teeth, of which the third tooth, counting from the apex, smallest; HW 0.98-1. 26. HL 0.92-1.16; postpetiole in dorsal view longer than broad or as long as broad, in profile with an arched anterior face and a steeply sloping posterior face; petiole node in profile with a single evenly rounded blunt apex. Worker. TL 3:71-4.84, HE 0:92=1.16,; HW 0.98-1.26, CI 104—111, SL 0.90—1.16, SI 89-98, EL 0.16—0.20, PW 0.62-0.79, AL 0.90—1.28, PPW 0.24—0.28, PPL 0.26-— OSS RPLS4=100NGr — o>): Mandible smooth and shining but sometimes with a few longitudinal rugae superficial or distinct. A broade based short tooth present about midway on the basal margin of the mandible. Clypeus with a short median longitudinal carina that usu- ally does not reach to the anterior clypeal margin; sometimes this carina indistinct; soniceimnes a few additional short rugae present. Anterior clypeal margin wach a median tooth and two lateral oe some- times the median tooth is smaller than the others. Ventral surface of clypeus lacking any rugae or teeth. Palp formula 1,3. Fron- tal carinae extending to the level of the posterior margins of eyes. Antennal scro- bes absent. F Frontal lobes : slightly expanded basally. Eyes containing 7 to 10 ommatidia in the longest row. reais of alitrunk and pedicel segments as in Figure 222. Pron- 504 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 221-224. Pristomyrmex lucidus Emery. 221: Worker head, full-face view; 222: Worker, lateral view; 223: Queen head, full-face view; 224: Queen, lateral view. otum at most with a pair of blunt tuber- cles, lacking teeth or spines. Propodeum with a pair of armaments, varying from broadly based minute teeth to moderately long acute spines. Metapleural lobes tri- angular or each with a_blunt-rounded apex. Petiole node in profile high, with a single evenly blunt-rounded apex and a long anterior peduncle. Postpetiole in pro- file high (slightly higher than petiole), with an arched anterior face and a steeply slop- ing posterior face, its apex pointing pos- terior-upwardly. In dorsal view, postpetiole broadening from front to back, mostly lon- ger than broad, rarely about as long as broad. Cephalic dorsum between the fron- tal carinae highly polished but usually with a few foveolate punctures bordering the frontal carinae. Sometimes a few foveolate punctures present on the genae and many on the ventral surface of the head. Dor- sum of alitrunk, petiole, and postpetiole unsculptured and highly polished. Gaster smooth and shining. Dorsal surfaces of head and alitrunk with sparse erect to sub- erect moderately long hairs. A pair of hairs present on the dorsum of petiole node and one to two pairs on the dorsum of post- petiole. First gastral tergite with a few hairs. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with numerous REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang suberect to erect short hairs. Body uni- form yellow-brown or blackish-brown or bicolored (i.e., alitrunk and pedicel seg- ments lighter than head and gaster). Oueen WWE oma a2. Sle A: RIV 22 1 27 EL lOss Ile SE 106.413: SI 87, 89; EL 0.26, 0.28; PW 0.96, 1.00; Ee) 1240.42: -PPW 0:31) 0:36 PPE 0:32, 0.38; PPI 95, 97 (n = 2). Generally similar to worker, except for normal caste differences. In addition, pro- podeal armaments usually shorter than those in the conspecific worker; first gas- tral tergite sometimes with numerous erect or suberect hairs. Male. Unknown. Comments. The workers of P. lucidus can be easily recognized by the following characters: (1) The postpetiole, in profile view, shows an arched anterior face and a steeply sloping posterior face, with the apex pointing posterior-upwardly. This shape is unique in the levigatus group. Furthermore, the postpetiole in dorsal view is longer than or as long as broad. In the other 11 species of the group, the post- petiole, in dorsal view, is distinctly broader than long. (2) The petiole node in profile view bears a single evenly blunt-rounded apex that is not seen in the other 11 spe- cies of the levigatus group. (3) P. lucidus has the largest head width (HW) in the levigatus group: In the 55 specimens mea- sured, HW is 1.00 to 1.26 but 0.98 in only one individual. In the other species of the levigatus group, HW is less than 0.90, ex- cept in one species, P. largus, in which, HW falls into the range 0.90 to 0.96. Material Examined (ANIC, MCZC, LAMN, USNM, CASC, BMNH#). Papua New Guinea: Tapini, 1,000 to 1,200 m, rainforest, acc. 2249 (rtw. wood fragment), acc. 2252 and 2262 (nest in soil under log) (R. W. Taylor); near Vanimo, rainforest, ca. 50 m (ex rotting log) (R. W. Taylor); Ya- wasora near Wewak, <50 m, rainforest, ex rotting wood piece (R. W. Taylor); 6 km NE of Wau, Bulolo R. Valley, rainforest 1,100 m, under bark of rotten log (R. W. Taylor); 8 km S of Kokoda, rainforest, 800 505 m, vial 4-36 (ground strays) and vial 37- 191 (ex rotting wood piece) (R. W. Taylor); Wau, Bishop Museum Station, 1,200 m, ex soil under rotten branch (R. W. Taylor); 9 mi on Lae, side of Mumeng, 3,500 ft, rain- forest (B. B. Lowery); Bulolo, 4,000 ft, rainforest (B. B. Lowery); Bulolo Gorge, 2,800 ft, rainforest (B. B. Lowery); along Kokoda Rd., 400 to 1,000 ft, rainforest (B. B. Lowery); 2 mi N Kokoda, ca. 1,000 ft, rainforest (B. B. Lowery); Wau Gorge, 3,000 ft, rainforest (B. B. Lowery); Wau, 4,000 ft, rainforest (B. B. Lowery); ca. 12 km SE Vanimo, 150 m Virgin hill rainfo- rest (W. L. Brown); Wau N on Bulolo Rd. B-278, 650 m (S. Peck); Bewani Rd., near Vanimo km 2 quarry, 40 m, lowland rain- forest (W. L. Brown); Maffin Bay (E. S. Ross). Indonesia: Irian Jaya, PT. Freeport Concession, Wapoga Camp., 03.14°S, 136.57°E, 3,450 ft, #98-230 (Montane pri- mary rainforest, ex rotten stick in litter) (R. R. Snelling). Ecological Information. This species oc- curs in rainforest and has been collected in soil under a log, under the bark of a rotten log, in litter samples, and on the ground. Pristomyrmex mandibularis Mann Figures 225-228 Pristomyrmex mandibularis Mann, 1921: 444. Syn- type workers, Fiji Is.: Viti Levu, Nadarivatu, 1915— 1916 (W. M. Mann) (AMNH, LACM, MCZC, USNH) [examined]. Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four teeth, of which the third tooth, counting from the apex, smallest; a strongly prominent tooth present about midway on the basal margin of mandible; petiole lacking a lateral longitudinal carina on each side; dorsum of head between frontal carinae usually smooth and shining. Worker. TL 2.68—-3.49, HL 0.74—0.84, HW 0.76-0.88, CI 95-107, SL 0.61—0.74, SI 80-93, EL 0.11—0.14, PW 0.48—0.60, AL 0.68—0.90, PPW 0.20-0.25, PPL 0.16— O19] RPMs — se)! Mandibles usually smooth and shining. 506 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 225-228. Pristomyrmex mandibularis Mann. 225: Worker head, full-face view; 226: Worker, lateral view; 227: Ergatoid queen head, full-face view; 228A: Ergatoid queen, lateral view; 228B: Dorsum of the ergatoid queen alitrunk, dorsal view. A prominent tooth present about midway on the basal margin of mandible. Clypeus flat and unsculptured. Anterior clypeal margin with a median tooth and two lat- eral teeth; the median tooth, in size, sim- ilar to or smaller than the others. Ventral surface of clypeus usually with a weak transverse ruga. Palp formula 1,3. Frontal carinae extending to the level of the pos- terior margins of the eyes. Antennal scro- bes absent. Frontal lobes slightly expand- ed. Eyes moderately sized. Occipital mar- gin straight or feebly concave in full-face view. Pronotum unarmed. Propodeum with a pair of short to moderately long spines. Metapleural lobes triangular. Peti- ole node in profile high, higher than long, with the anterodorsal angle being an apex and the dorsum posteriorly rounding into the posterior surface, its anterior peduncle slightly longer than or about as long as the node. Postpetiole in profile high, about two times as high as long, rounded dor- sally. In dorsal view, postpetiole transrec- tangular. Dorsum of head between the frontal carinae usually smooth and shining REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang but sometimes with a few small and shal- low punctures. Sometimes a few foveolate punctures bordering the frontal carinae, present on the genae and around the eyes. Dorsum of alitrunk smooth and shining. Petiole and postpetiole smooth and shin- ing, each lacking a lateral longitudinal ca- rina on each side. Gaster unsculptured. Dorsal surface of head with numerous erect to suberect long hairs. Some of sim- ilar hairs present on the dorsum of ali- trunk, a pair on the petiole, one to two pairs on the postpetiole, and a few on the first gastral tergite. A few pairs of forward- projecting hairs present near the anterior clypeal margin. Scapes and tibiae with nu- merous erect to suberect hairs. Color red- dish-brown to blackish-brown. Ergatoid Queen. TL 3.32, 3.40; HL 0.84, 0.84; HW 0.82, 0.82; CI 98, 98; SL 0.66, 0.73; SI 80, 89; EL 0.14, 0.15; PW 0.54, 0.56; AL 0.78, 0.82; PPW 0.23, 0.24; PIPIE, OS). OfVile IPP TS a Ge, =) Generally similar to worker, color and pilosity as in the worker, but the head with one ocellus, alitrunk in dorsal view with a pro-mesonotal suture, mesonotum more convex, and propodeal armaments shorter than in the conspecific worker. Queen and Male. Unknown. Comments. Pristomyrmex mandibularis is endemic in Fiji and is the only Pristo- myrmex species so far occurring there. It possesses an ergatoid queen caste, which is not seen in the other species of the lev- igatus group; furthermore, its ergatoid queen has only one ocellus. Pristomyrmex mandibularis is closely related to P. levigatus and P. largus, also from the Oriental region. The differences between the workers of P. mandibularis and those of P. levigatus and P. largus are as follows: P. mandibularis Petiole and postpetiole each lacking a lateral longitudinal carina on each side Basal margin of mandible with a strong- ly prominent tooth 507 Anterior clypeal margin usually with a distinct median tooth P. levigatus and P. largus Petiole and postpetiole each with a lon- gitudinal carina on each side that sep- arates the tergite from the sternite Basal margin of mandible with a broad- based short tooth A median tooth, on the anterior clypeal margin, usually lacking or very weak Material Examined (ANIC, MCZC, USNM, BMNH, MHNG). Fiji: Nausori Highlands, #424 (rotting stick in litter) (W. L. and D. E. Brown); Viti, Nadarivatu, rainforest, acc. 83 (berlesate, leaf mold) and acc. 66.51 (forest floor, colony in small crevice, rotting branch fragment) (R. W. Taylor); Nadarivatu (W. M. Mann); Lase- ma (W. M. Mann); Viti, Levu, Nadarivatu Reserve, 17.34'S, 177.57'E, Rainforest 800 m, Q. M. Berlesate No. 775, sieved litter (G. Monteith); Kadavu, 2 km SE Vunisea, 19.04’'S, 178.10'E, rainforest 20 m, Q. M. Berlesate No. 770, sieved litter (G. Mon- teith). Ecological Information. This species oc- curs in rainforest and has been collected in litter berlesates; it nests beneath stones in small colonies (Mann, 1921). Pristomyrmex minusculus sp. n. Figures 229-232 Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and pos- sessing four teeth, of which the third tooth, counting from the apex, smallest; pronotum with a pair of teeth. Holotype Worker (MCZC). TL 3.02, HL 0.76, HW 0.80, CI 105, SL 0.66, SI 83, EL 0.14, PW 0.52, AL 0.78. Paratypes, 16 workers and one queen (MCZC, ANIC, BMNH, NACA). Worker. TL 2.52-3.02, HL 0.66—0.78, HW 0.66-0.80, CI 99-106, SL 0.54—0.66, SI 77-84, EL 0.10-0.14, PW 0.45-0.52, AL 0.59-0.74, PPW 0.20-0.24, PPL 0.14— 0.19, PPI 126-143 (n = 16). Mandibles usually smooth and shining but sometimes with a few small shallow 508 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 229-232. Pristomyrmex minusculus sp. n. 229: Worker head, full-face view; 230: Worker, lateral view; 231: Queen head, full-face view; 232: Queen, lateral view. punctures. A broad-based short tooth pre- sent about midway on the basal margin of the mandible. Clypeus unsculptured. An- terior clypeal margin with a median tooth and two lateral teeth, but the median tooth often smaller than the others, sometimes the median tooth indistinct. Ventral sur- face of clypeus with a weak transverse ruga. Palp formula 1,3. Frontal carinae ex- tending to the level of the posterior mar- gins of the eyes. Scrobal areas shallow, short, present lateral to the frontal carinae in full-face view. Frontal lobes weakly ex- panded. Eyes moderately sized. Occipital margin feebly concave. Alitrunk in dorsal view more or less flat. Pronotum with a pair of acute small teeth; in some small specimens, this pair of teeth are very weak but visible, and in dorsal view they become a pair of sharp points on the two sides of the pronotum. Propodeum armed with a pair of spines, varying in length and shape, straight to slightly upcurved along their length. Metapleural lobes subtriangular, but rarely with a rounded apex. Petiole node in profile high with the anterodorsal angle higher than the posterodorsal, its an- terior surface usually subparallel to the posterior one, its anterior peduncle about as long as the node. Subpetiole with a nar- row rim. Postpetiole in profile high, rounded dorsally. Petiole node and post- petiole in dorsal view broader than long. Dorsum of head between the frontal ca- REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang rinae smooth and shining but sometimes with a few foveolate punctures bordering the frontal carinae, present on the genae and around the eyes. Dorsum of alitrunk smooth and shining. Petiole and postpeti- ole each with a lateral longitudinal ruga on each side that separates the tergite from the sternite. Gaster unsculptured. Dorsal surfaces of head and alitrunk with sparse erect to suberect hairs. Dorsal surfaces of petiole node and postpetiole each usually with two pairs of hairs and first gastral ter- gite with a few hairs, as in Figure 230. A few pairs of forward-projecting hairs pre- sent near the anterior clypeal margin. Scapes and tibiae with some erect to sub- erect short hairs. Color yellow-brown to reddish-brown; appendages lighter. Queen. TL 3.08, HL 0.72, HW 0.74, CI 103, SL 0.60, SI 81, EL 0.17, PW 0.60, AL 0.82, PPW 0.24, PPL 0.18, PPI 133 (n = 1) Generally similar to worker, except for normal caste differences. In addition, pro- notal armaments absent; propodeum with a pair of teeth or short spines that are shorter than those in the conspecific work- er. Male. Unknown. Comments. This species has a wide dis- tribution in the Oriental region. It is also dispersed to North Queensland, Australia. Pristomyrmex minusculus must be derived from a P. levigatus—like ancestor. It is in- distinguishable, in the queen, from P. lev- igatus at present. The workers of P. min- usculus have almost same appearance as those of P. levigatus, except for a derived character—the pronotum with a pair of small teeth. In the levigatus group, this critical character is possessed only by the workers of P. minusculus, thus making them easily recognizable. Holotype Worker. Palau Is: Peleliu L., east coast, 26.1.1948 (H. S. Dybas). Paratypes. Nine workers and one queen with same data as holotype; one worker, Wallis Is.: NukuTapu I., 28.iii.1965 to l.iv.1965 (G. Hunt). Two workers, Indo- nesia: Seram, above Haruru, near Masohi, 509 50 to 150 m, one worker, Indonesia: Irian Jaya, 12 km S of Sorong, forest fragment, 1.v.1981 (W. L. Brown); two workers, Tonga Is.: Fale- hau, Niuatoputapu, moss + lichen, from coconut tree trunks, 1.ix.1971 (W. and G. Rogers); two workers, YapGroup, vii— viii.50. (R. J. Goss). The following additional (non-type) specimens have a pair of very weak pro- notal prominences. They have the follow- ing measurements: Worker: HW _ 0.60— 0.72, HL 0.62—0.72, CI 97-103, SL 0.52— 0.60, SI 77-87, EL 0.10-0.12, PW 0.42-— 0.48, AL 0.57—0.73, PPW 0.19—-0.21, PPL 0.14—0:116, PPI 125-150 (n = 12). Oueen: IED Ow Ol Moe Ini; One, OLwes (Cll IC) LOOSE GO!625 0622 SIeS2aro2 ken OokS: 0.18; PW 0.62, 0.62; AL 0.82, 0.88; PPW ORAS 0:24 SRP OM ORI eA Ss Aa (a= 2). Collecting Data for These Non-Type Specimens (ANIC. USNM, BMNH). Papua New Guinea: Kiunga (J. Balogh); Bisian- umu near Sogeri, rainforest, 500 m (E. O. Wilson); Maffin Bay (E. S. Ross). Micro- nesia: Pohnpei, Agric. and Trade School, in leaf litter (in Ylang- Ylang grove) and in rotting coconut tree (Ron Clouse). Austra- lia: N. Queensland, Cape York, Lockerbie (G. B. Monteith); N.Q., Iron Ra. rainfo- rest, berlesate (R. W. Taylor and J. Fee- han). Ecological Information. This species oc- curs in rainforest and has been collected in litter samples, and in a rotting tree. 18.iii.1981 (W. L. Brown); Pristomyrmex obesus Mann Figures 233-236, 267, 278 Pristomyrmex obesus Mann, 1919; 339. Syntype workers, queen and male, Solomon Is.: Ysabel, Fu- lakora; Malaita, Auki; Three Sisters, Malapaina; 19.v.-24.xi.1916 (W. M. Mann) (MCZC, USNM, AMNH) [examined]. Pristomyrmex pegasus Mann, 1919: 338. Holotype worker, Solomon Is.: Santa Cruze: Graciosa Bay, 19.v.-24.xi1.1916 (W. M. Mann) (USNM) [exam- ined]. Syn. n. Pristomyrmex obesus subsp. melanoticus Mann, 1919: 340. Syntype workers, Solomon Is.: San Cristoval, Pamua: Wai-ai; 19.v.—4.x1.1916 (W. M. Mann) (MCZC, USNM) [examined]. Syn. n. 510 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 233-236. Pristomyrmex obesus Mann. 233: Worker head, full-face view; 234: Worker, lateral view; 235: Queen head, full-face view; 236: Queen, lateral view. Note: Pristomyrmex pegasus and P. obe- sus were described as two new species in Mann’s (1919) paper, on pages 338 and 339, respectively (i.e., P. pegasus appeared before P. obesus; see also the previous ci- tation). However, when proposing that these two names are synonymic, I choose P. obesus instead of P. pegasus as a valid specific name for the following two rea- sons: (1) Mann (1919) mentioned the pres- ence of the elevated sides of the mesotho- rax, and the absence of a median tooth on the anterior clypeal margin are character- istic of P. pegasus. However, “the elevated sides of the mesothorax” are not shown in the unique holotype of P. pegasus, and “the absence of the median tooth on an- terior border of clypeus” is actually an in- dividual variation. (2) P. obesus possesses about a dozen syntypes, including a female and a male, but P. pegasus has only a single type specimen (i.e., holotype). If the ho- lotype of P. pegasus is lost or destroyed in the future, it would be very inconvenient to those people who want to see it. Thus, P. obesus is selected as a valid name here. Diagnosis (Worker). Masticatory margin of mandible lacking a diastema and _ pos- REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang sessing four teeth, of which the third tooth, counting from the apex, smallest; eyes containing = fou to seven ommatidia in the longest row; dorsum of head, except for the scrobes, usually with dense foveo- late punctures; entire first gastral tergite evenly covered with numerous erect or suberect hairs. Worker. TL 2.24-3.22, HL 0.63-0.86, HW 0.61-0.86, CI 94-104, SL 0.52—0.75, Sl 79-88, EL 0.09-0.14, PW 0.41—0.55, AL 0.58—0.88, PPW 0.18—0.24, PPL 0.16— 020) PRL 105-125 Ga = 90). Mandibles usually smooth and shining but sometimes with a few superficial short rugae or a few hair pits. A broad-based tri- angular short tooth present about midway on ahe basal margin of the mandible. Clyp- eus flat, smooth, | aa the median carina of frontal area usually extending to the clyp- eus. Anterior clypeal margin ith a median denticle and two lateral teeth but the me- dian denticle often smaller than the oth- ers; sometimes the median denticle lack- ing; thus, only two teeth are present there. Wentr al surface of clypeus lacking toothlike prominences or rugae. Palp formal RSE Frontal carinae extending to the level of the posterior margins of eyes. Scrobal im- pressions broad, Sellen: present lateral to the frontal carinae. Frontal lobes slightly expanded. Eyes moderately sized, usually containing five to seven (rarely four) om- matidia in the longest row. Profile of ali- trunk and pedicel segments as in Figure 234. Pronotum unarmed. Propodeum with a pair of triangular short spines. Meta- pleur: al lobes rounded. Dorsum of alitrunk in dorsal view usually with a longitudinal impression or furrow at middle, bat some- times this longitudinal impression indis- tinct. Petiole mode in profile high with the anterodorsal angle higher than nile poster- odorsal, its Ankonor “peduncle nearly as long as the node. Subpetiole with a narrow semitranslucent lamella. Postpetiole in profile rounded dorsally. In dorsal view, dorsum of petiole node subrounded, about as long as broad; postpetiole somewhat transversally rectangular and _ slightly 511 broader than long. Dorsum of head, ex- cept for the scrobes, usually with dense fo- veolate punctures. Dorsum of alitrunk with scattered foveolate punctures. Peti- ole, postpetiole, and gaster smooth and shining. Dorsal surfaces of head and ali- trunk with numerous erect to suberect long hairs. Two pairs of hairs usually pre- sent on the dorsum of petiole node and two to three pairs usually on the dorsum of postpetiole. Entire first gastral tergite covered with numerous, evenly distr cece erect or suberect hairs. A few pairs of for- ward-projecting hairs present near the an- terior clypeal margin. Scapes and _ tibiae with numerous erect to suberect short hairs. Color reddish-brown. Queen. TL 2.69-3.40, HL 0.69-0.82, HW 0.67-0.85, CI 97-104, SL 0.54—0.68, SIP 7S—Sose IE Onlds=0 199 RWaO54=0164 AL 0.76—0.94, PPW 0.20-0.26, PPL 0.18— 022) Peis 25 =s1'5) Generally similar to worker, except for caste differences. In addition, foveolate punctures shallow on the mesonotum; pro- podeal armaments slightly shorter than those in the conspecific workers: dorsum of alitrunk lacking a longitudinal furrow. Male (Figs. 267, 278). A syntype male, together with Alyontt a dozen workers and a queen, collected in Fulakora, Solomen Th. by W. M. Mann on 19.v.—24.xi.1916, constitutes a series. A second male speci- men, together with five workers, collected in Guadalcanal, Solomen I., by P. Green- slade, constitutes another series. Each of the two males was originally mounted with two or three workers, respectively, on the same pin. TL 2.92; HL 0.52, 0.53; HW 03533) 0'54- CI 0250102 3S O22 0802235 38, 41: HWE 0.66, 0.68; EL 0.25, 0.26: PW 0.58, 0.60; AL 0.90, 0.92; PPW 0.18; PRE OS SPR LOOnGs—s2): Head, including the eyes, broader than long. Clypeus convex, without a median longitudinal carina. Anterior clypeal mar- gin transverse. Frontal carinae weak or in- distinct. On the mesoscutum, notauli dis- tinct, forming a Y shape; parapsidal fur- rows absent. Scuto-scutellar sulcus rather 512 broad, with five narrow longitudinal ridg- es. Propodeum weakly faberculate: lacking teeth and spines. Metapleural lobes some- what rounded. Petiole node in profile low, rounded dorsally. Postpetiole in profile low, rounded dorsally, in dorsal view sub- quadrate. Dorsum of head unsculptured and shining, except for a median longitu- dinal carina present on the frontal area. Dorsal surface of alitrunk smooth and shining but with well-marked sutures. Pet- iole, postpetiole, and gaster smooth and shining. Dorsal surfaces of head, alitrunk and gaster with abundant long hairs. Legs ane numerous hairs. Body lacigehe brown; wings infuscated. Gonmmients: Pristomyrmex obesus occurs on Solomon Is. It is similar in the workers to P. simplex of New Guinea but can be separated from the latter by possessing nu- merous erect or suberect hairs evenly dis- tributed on the entire first gaster tergite. In the levigatus group, this character is present only in two species, P. obesus and P. longus; in the other species, the first gas- tral tergite has a few or no hairs. In addi- tion, foveolate punctures on the dorsal head are denser in the workers of P. obesus than in P. simplex. Material Examined (ANIC, MCZC, BMHH, USNM, BMNH#). Solomon Is.: Ysabel, Fulakora (W. M. Mann); Guadal- canal, Mt. Austen (P. Greenslade); Guad- aleanal, Kukum (P. Greenslade); Guadal- canal, Gold Ridge, 2,000 ft (P. Green- slade); Guadaleanal, Mt. Jonapau, 2,600 to 3,500 ft (P. Greenslade); Guadalcanal, Na- limbiu R. (P. Greenslade); Guadalcanal, Balesuna R. (P. Greenslade); Guadalcanal, Visale (P. Greenslade); Guadalcanal, Uma- sani R., 1,000-ft ridge, leaf litter (P. N. Lawtence); San Cristoval, Warahito R., 275 ft (P. Greenslade); San Cristoval, for- est, 250 ft (P. Greenslade); Malaita, Small Malaita (P. Greenslade); Malaita, Dala (P. Greenslade); New Georgia, Kolomban- gara, Hunda (P. Greenslade): New Geor- gia, Kolombangara, S. Kusi (P. Green- slade); New Georgia, Kolombangara, N of Kuzi, 500 ft, forest litter (P.N.L.); New Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Georgia, Vella Lavella, Bara koma (P| Greenslade): New Georgia, Vangunu I. (P. | Greenslade); Choiseul, Wagina I. (P Greenslade); Russell Is., Luavic (P. Green- slade); Nggela (P. Green lade). Isabel, Bu- ala (P. Greenslade); Vanikord (P. Green- | slade); Bougainville I., Panguna, 600 to. 800 m (J. i Gressitt). | Ecological Information. This species has _ been eollemnadl | in forest litter. Mann (1919) | found a colony composed of less than a dozen workers, a dealated queen, and one | male under a stone. Pristomyrmex simplex sp. n. Figures 237-240 Diagnosis (Worker). Masticatory margin | of mandible lacking a diastema and pos- sessing four ere of which the third tooth, counting from the apex, smallest; dorsal sinthioest of alitrunk and head bes tween the frontal carinae with scattered foveolate punctures; eyes usually contain- ing five (rarely four) ommatidia in the lon- gest row; a few hairs present on the first gastral tergite. Holotype Worker (ANIC). TL 2.48, HE 0.66, HW 0.69, CI 105, SL 0.52, SI 7& EL 0.12, PW 0.46, AL 0.58. Paratypes, six workers and three queens (MCZC, ANIC, BMNH). Worker. TL 2.42-2.64, HL 0.65—0.683 HW 0.66-—0.70, CI 100—105, SL 0.52—0.54, SI 74-79, EL 0.09-0.12, PW 0.44—0.46; AL 0.58—0.66, PPW 0.18—0.20, PPL 0.14— 0.16, PPI 119-129 (n = 6). Mandibles usually smooth and shining but sometimes with a few small shallow hair pits and a few superficial short rugae. A broad-based triangular short tooth pre- sent about midway on the basal margin of the mandible. Clypeus flat, emoothe and shining, usually unsculptured but some- times fie median carina of the frontal area extending to the clypeus. Anterior clypeal margin wath a median denticle and two lat- eral teeth; the median denticle is often smaller than the others; sometimes the median denticle is absent so that the only two teeth are present there. Ventral sur- REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 513 Figures 237-240. Pristomyrmex simplex sp. n. 237: Worker head, full-face view; 238: Worker, lateral view; 239: Queen head, full-face view; 240: Queen, lateral view. face of clypeus lacking rugae or toothlike prominences. Palp formula 1,3. Frontal ca- rinae distinct, extending to the level of the posterior margins of eyes. Scrobal areas slightly concave, present lateral to the frontal carinae. Frontal lobes weakly ex- panded. Eyes moderately sized, about 0.14 to 0.17 X HW, usually containing five (sometimes four) ommatidia in the longest row. Occipital margin feebly concave in full-face view. Profile of alitrunk and ped- icel segments as in Figure 238. Pronotum unarmed. Propodeum with a pair of tri- angular teeth. Metapleural lobes rounded. Petiole node in profile high with the an- terodorsal angle higher than the postero- dorsal, its anterior peduncle about as long as the node. Subpetiole with a narrow rim. Postpetiole in profile rounded dorsally. In dorsal view, dorsum of petiole node tran- soval and dorsum of postpetiole somewhat transversely rectangular. Dorsum of head between the frontal carinae with scattered foveolate punctures, varying from a few feeble punctures to numerous distinct ones; spaces between foveolae often 514 smooth. Dorsum of alitrunk with a few to some scattered foveolate punctures. Peti- ole, postpetiole, and gaster smooth and shining. Dorsal surfaces of head and ali- trunk with numerous erect to suberect hairs. Two pairs of hairs usually present on the dorsal surfaces of petiole node and postpetiole, respectively, and a few near the base of the first gastral tergite. A few pairs of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect to suberect short hairs. Color reddish-brown. Queen. TL 2.51—2.84, HL 0.66—0.70, HW 0.66—0.71, CI 1OO—101, SL 0.51—0.58, SI 76-82, EL 0.14—0.16, PW 0.50-0.55, AlevO166-0:S81, BRW.0N9=0015 PRM ON5= ONS) PPI 114=1335@7. = 3): Generally similar to worker, except for caste differences. In addition, mesoscutum rather smooth and shining, but mesoscu- tellum with a few foveolate punctures; propodeal armaments slightly shorter than those in the conspecific worker. Male. Unknown. Comments. This species occurs in New Guinea. It is closely related to P. levigatus and P. obesus. It is separable from P. levi- gatus by possessing some foveolate punc- tures on the dorsum of the alitrunk in the workers and on the dorsal head between the frontal carinae in the workers and queens. It differs from P. obesus because the workers of P. simplex have only a few hairs on the first gastral tergite. Pristomyr- mex sinylex differs from P. coggii by the following characters: The eyes usually con- tain five (sometimes four) ommatidia in the longest row in the workers of P. sim- plex but two to three ommatidia in P. cog- gii; the dorsum of the head has foveolate- reticulate sculpture in the workers and queens of P. coggii but scattered foveolate punctures in P. simplex. Holotype. Papua: 8 km S of Kokoda, S00 m, rainforest, ANIC Berleasate, No. 382, L.vi.1972 (R. W. Taylor). Paratypes. One worker with same data as holotype; two workers, Papua: Karema, Brown R., lowl. rainfor., 8—11.iii.1955 (E. Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 O. Wilson); three workers and one queen, Papua New Guinea: Port Moresby, Brown | River, 2.x.1969 (J. Balogh); one queen; Papua: N. Dist., 27.xii.1971 (P. M. Room); one queen, Papua: N. Dist., Debelou, 23.vi.1973 (P. M. Room). Non-Type Material Examined. A worker (NHMV), collected in the Philippines (Lu-_ zon, Lagunas, Mt. Banahaw above Kina- buhayan, 600-700 m) by J. Kodada and B. Rigova, has the following measurements: TL 2.98, HL 0.84, HW 0.84, CI 100, SE 0.70, SI 83, EL 0.10, PW 0.54, AL 0.748 A second worker (LAMN), collected in New Guinea (Gulf Prov., Ivimka Camp, Lakekamu Basin, 7.73°S, 146.76°E, 120 m, #96-291, lowland wet forest, ex sifted leaf litter) by R. R. Snelling, bears a few small, feeble, shallow punctures on the dorsal al- itrunk; this specimen possesses TL 2.48, HL 0.68, HW 0.69, CI 101, SL 0.58, Sil 84, EL 0.09, PW 0.45, AL 0.62. Ecological Information. This species oc- curs in rainforest and has been collected in litter samples. THE PROFUNDUS GROUP Worker. Small sized, with the following combination of characters. (1) Masticatory margin of mandible lacking a diastema and possessing four teeth (i.e., the strongest apical + the sec- ond strongest preapical + the smallest third + an acute basal tooth); basal margin of mandible with a strongly prominent tooth that is adjacent to the basal tooth of the masticatory margin; as a result, five teeth are set close together. (2) Lateral portions of clypeus in front of antennal fossae reduced to margins. (3) Antennal scrobes broad, deep, and extending close to the occipital corners. (4) Base of antennal scape lacking a cir- cling lamella. (5) Mesonotum much higher than pro- podeal dorsum so that the dorsum of ali- trunk in profile is not continuously arched. (6) Metapleural lobes vestigial and in- distinct. The profundus group is closely related to the levigatus group. The form of den- tition of the masticatory margin of the mandible is a critical character shared by the workers and queens of the two groups. Possessing so many autapomorphie char- acters, including that a tooth on the basal margin of the mandible is adjacent to the basal tooth of the masticatory margin, as well as the previously mentioned charac- ters 3 to 6, the profundus group is easily separable from the levigatus group and all other Pristomyrmex species groups. This group contains only a single spe- cies, P. profundus of Malaysia. Pristomyrmex profundus sp. n. Figures 241-244 Diagnosis (Worker). See characters 1 and 5 melee the profundus group. Holotype Worker (BMNH). TL 2.86, HL 0.64, HW 0.74, CI 119, SL 0.44, SI 59, EL 0.13, PW 0.52, AL 0.68, PPW 0.24, PPL 0.16, PPI 150. Paratypes, 19 workers and three queens (BMNH, MCZC, MHNG, ANIC, LAMN). Worker. TL 1.74—2.86, HL 0.46-0.66, HW 0.46-0.74, CI 96-119, SL 0.36-0.44, Slo 97, 9 Ee OL09=0545 RW. 0!34=0:522 mls O.44=0.68: PEW O115-0.24 PPL 0.11— lo. PRL 123-645 m= 20): Mandibles usually smooth and shining but sometimes with a few weak, short, bas- al rugae. Clypeus not depressed, usually with a median longitudinal carina and two transverse carinae. Anterior clypeal margin lacking any distinct denticles. Ventral sur- face of clypeus lacking any toothlike prom- inences but usually with a weak transverse ruga. Palp formula undissected. Frontal carinae divergent, extending beyond the level of the posterior margins of eyes and close to the occipital margin. Frontal lobes completely absent. Antennal scrobes deep, wide, present between the frontal carinae and the eyes for the reception of the scapes and funiculi of antennae. Antennal scapes, when lying in the scrobes, close to the occipital margin of head. Eyes contain- ing five to eight ommatidia in the longest row. In full-face view, head widest near the REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 515 occipital corners; occipital margin rather straight. Dorsum of alitrunk in profile not continuously arched, with the mesonotum much higher than the propodeal dorsum (i.e., a vertical cliff present between the mesonotum and the propodeal dorsum). Pro-mesonotum forming a single convex. Sometimes the mesonotum higher than the pronotum; thus, in dorsal view, a pro- mesonotal impression is seen. Pronotum unarmed. Propodeum with a pair of mod- erately long spines. Metapleural lobes ves- tigial and indistinct. In profile view, petiole node high, ca. 0.21 to 0.29, much higher than long and also higher than postpetiole, with a single evenly blunt-rounded apex and a long anterior peduncle. Subpetiole with a narrow lamella. Postpetiole in pro- file high, ca. 0.15 to 0.23, distinctly higher than long, with a rounded dorsum. In dor- sal view, postpetiole transoval and much broader than long. Dorsum of head, ex- Cope tOmathe atennl scrobes, fully cov- ered with rugoreticulum. Galen sculpture present on tie pro-mesonotum. Propodeal dorsum with a few longitudinal rugae. Pet- iole smooth and shining, usually with a longitudinal carina on Sch side. Postpe- Bole and gaster smooth and shining. Dor- sal sirtaces of head, alitrunk, aide gaster with numerous erect, thick, long eae sometimes some hairs somewhat clavate A pair of hairs present on the dorsum of pet- iole node and usually two pairs on the dor- sum of postpetiole. A few pairs of forward- projecting hairs present near the anterior clypeal margin. Scapes and tibiae with some erect or suberect moderately long hairs. Color light yellow to yellow brown. Queen. PIL, 3 46—5592 ee Euley OL62=0:635 HW 0.76—-0.77, CI 123-124, SL 0.44—-0.46, Si o7— Ole EL Onl5=0s638 RW 710'66=0570) AL 0.92-0.98, PPW 0.28—0.29; PPL 0.18— (0) A0), IAP TAO (= S&S). General shape as in Figures 243-244, with normal caste differences from the conspecific worker; mesonotum unsculp- tured, smooth, and shining; anterior end of mesoscutum medially slightly concave. Other characters similar to worker. 516 P FREES =~ Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figures 241-244. Pristomyrmex profundus sp. n. 241: Worker head, full-face view; 242: Worker, lateral view; 243: Queen head, | full-face view; 244: Queen, lateral view. Male. Unknown. Comments. This is a distinct species be- cause many characters possessed by its workers and queens are unique in the ge- nus. In the entire Pristomyrmex fauna, three species (P. profundus, P. divisus, and P. pulcher) do not possess any denticles on the anterior clypeal margin, but the latter two species belong to the punctatus group. Holotype Worker. Malaysia: Sabah, Por- ing Hot Springs, 500 m, 7.v.1987 (Burck- hardt and Lobl). Paratypes. Seventeen workers and three queens with same data as holotype; two workers with same data as holotype but date 6.v.1987. Ecological Information. Unknown. THE UMBRIPENNIS GROUP Worker. Medium to large sized (HL | 1.04-1.68, HW 1.02-1.74, TL 4.10-7.06) with the following combination of charac- ters. (1) Masticatory margin of mandibles _ with four teeth arranged as the strongest. apical + the second strongest preapical + two broad-based short teeth of similar size; diastema lacking or indistinct. (2) Basal margin of mandible with a) central, broad-based, prominent lobe. (3) Lateral portions of clypeus in front | of antennal fossae reduced to a margin. (4) Anterior margin of the median por- tion of clypeus usually with five to seven REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 517 246 Figures 245-246. Pristomyrmex fuscipennis (F. Smith). 245A: Queen head, full-face view; 245B: Showing a transverse carina on the ventral clypeus; 246: Queen, lateral view. denticles; lateral portions of anterior clyp- eal margin in front of antennal fossae with a few weak blunt denticles. (5) Ventral surface of clypeus with a transverse ridge. (6) Frontal lobes present, partly cover- ing the condylar bulbs of holding antennal scapes. (7) Frontal carinae extending to the lev- el of the posterior margins of eyes. (S) Lamella that encircles the base of antennal scape usually with a broad and deep notch on the center of the dorsal sur- face. (9) Palp formula 1,3. (10) Eyes small; EL is about 0.040 to 0.064 X HW in P. picteti and P. pollux and 0.069 to 0.108 k HW in P. umbripennis. (11) Alitrunk in profile, not including propodeal spines, with a regularly arched dorsum, in dorsal view without any su- tures. (12) Pronotum unarmed. (13) Metapleural lobes bluntly round- ed. (14) Petiole node in profile longer than high, with a long anterior peduncle; its an- cenadlonsall angle is on approximately the same level as the posterodorsal. (15) Foveolate punctures or foveolate- reticulate sculpture present on the dorsal surfaces of the head and the alitrunk. This is a monophyletic group, contain- ing five valid species. It is easily recogniz- able by possessing characters 1, 2,5, 8, and 14. This group is endemic in the Oriental region and restricted to the Philippines, Malaya, Singapore, Brunei, Sabah, Bor- neo, Indonesia, and Papua New Guinea, The males of the wumbripennis group are easily distinguished from the oer known Pristomyrmex males by possessing the fol- lowing characters: (1) medium to large size; (2) palp formula 1,3; (3) propodeum with a pair of broad-based, robust spines (Figs. 256, 279, 280); and (4) the sides of petiole with some longitudinal or reticu- late rugae (Figs. 256, 279, 280). Pristomyrmex fuscipennis (F. Smith) Figures 245-246 Myrmica fuscipennis F. Smith, 1861: 46. Holotype a Indonesia: Celebes, Tondano (A. R. Wal- lace) (OXUM) [examined]. ? Pristomyrmex fuscipennis (F. Smith) Emery, 1901: 567. Pristomyrmex fuscipennis (F. Smith) 1932: 468. Donisthorpe, Queen. TL 6.92, HL 1.62, HW 1.64, Cl OM SE MEAG Sl -SOn Hl 032 ee ENVIR S 2D: AL 1.98 (n = 1). Comments and Discussion. This species, described from a single queen, obviously belongs to the wmbripennis group by pos- sessing the following characters: (1) mas- ticatory margin of Sanandible with four 518 teeth (an apical + a preapical + two broad-based short teeth of similar size), lacking a distinct diastema; (2) basal mar- gin of mandible with a central, broad- based, prominent lobe; (3) frontal lobes partially covering the condylar bulbs of holding antennal scapes; (4) lamella that encircles the base of antennal scape with a broad and deep notch on the center of the dorsal surface; (5) a coarse transverse carina present on the ventral surface of clypeus; and (6) anterodorsal angle of pet- iole node in profile on approximately the same level as the posterodorsal. This queen differs from the queens of P. picteti, P. pollux, and P. umbripennis as follows: In P. fuscipennis, the dorsum of the head possesses foveolate-reticulate sculpture; many foveolate punctures be- tween the frontal carinae are almost aligned so that it seems that several lon- gitudinal rugae are present there. These longitudinal rugae are indistinct or absent in P picteti, P. pollux, and P. umbripennis. In P. picteti and P. umbripennis, the dor- sum of the head possesses scattered fove- olate punctures. In addition, in P. fusci- pennis, the antennal scapes lack longitu- dinal carinae along their dorsal margins, a median longitudinal carina runs through the entire clypeus and frontal area, dad only five teeth are present on the anterior margin of the median portion of the clyp- eus, Sich are different from those in P. pollux. Pristomyrmex fuscipennis may be a sib- ling species of P. picteti because the queen of P. fuscipennis is very similar to that of P. picteti, except for possessing foveolate- reticulate sculpture and longitudinal rugae on the dorsal head. In Battin: I have ex- amined several workers that may belong to P. fuscipennis. The cephalic sculpture of these workers is similar to that of the queen of P. fuscipennis, but the other characters of the workers are identical with those of the workers of P. picteti. Finally, I feel that P. fuscipennis may represent an incipient species. Further Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 collecting and ecological investigations are needed. Ecological Information. Unknown. Pristomyrmex picteti Emery Figures 247-250, 268, 279 Pristomyrmex picteti Emery, 1893: 190. Lectotype worker, Sumatra: Deli (Bedot) (MCSN), here des- ignated, [examined]. Pristomyrmex picteti var. tingiana Stitz, 1925: 120. Syntype workers, Philippines: N. Palawan, Binal- uan, xi-xii 1913 (G. Boettcher) (MNHU, MCZC) [examined]. Syn. n. Diagnosis (Worker). Masticatory margin of mandible with four teeth (an apical + a preapical + two broad-based short teeth of similar size), lacking a distinct diastema; eyes small, with three to four ommatidia in the longest row; propodeal spines fairly long, about 1.3 to 1.5 x the distance be- tween their bases, not strongly upcurved at their apices. Worker ls 4110=5:84-) lge L04= 1836" HW 1.02—1.40, CI 96—108, SL 0.92-1.26; SI 85-97, EL 0.05—0.09, PW 0.68—0.88, AL 1.12—1.58, PPW 0.30-0.41, PPL 0.274 0:36, PRE 100=120°@-=40)) Mandibles rather smooth or sometimes with a few longitudinal rugae. Basal mar- gin of Tanchible with a br eagle based, sub- triangular, short prominence or a some- eohate curved lobe. Clypeus smooth and shining, except for a median longitudinal carina hat usually crosses the entire clyp- eus but sometimes does not reach the pos- terior clypeal margin. Anterior margin of the median portion of clypeus (not ind ing the margins in front of the antennal fossae) with a median denticle and two (sometimes three) lateral denticles on each side. Ventral surface of clypeus with a coarse, transverse, long carina. Palp for- mula 1,3. Frontal carinae usually beyond the eyes. Antennal scrobes indistinct. Frontal lobes present, partially covering the condylar bulbs of holding antennal scapes. Antennal scapes, when lying on the dorsal head, just reaching or slightly be- yond the occipital margin. Lamella, encir cling the base of antennal scape, usually REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 519 Figures 247-250. Pristomyrmex picteti Emery. 247A: Worker head, full-face view; 247B: Showing a transverse carina on the ventral clypeus; 248: Worker, lateral view; 249A: Queen head, full-face view; 249B: Showing a long transverse carina on the ventral clypeus; 250: Queen, lateral view. with a broad and deep notch on the center of the dorsal surface, but in several spec- imens (from Palawan I., Philippines), this lamella is entire, without a notch. Eyes very small, usually with three, at most four, ommatidia in the longest row. Occipital margin straight or very feebly concave in full-face view. Alitrunk in profile with a convex dorsum. Pronotum unarmed. Pro- podeum with a pair of fairly long spines, ca. 1.3 to 1.5 X the distance between their bases; sometimes the spines are weakly upcurved at their apices. Metapleural lobes rounded or somewhat truncated. Petiole node in profile slightly longer than high, with a fairly long anterior peduncle; its anterodorsal angle is usually on approx- imately the same level as the posterodor- sal, but sometimes the former is slightly higher than the latter. Subpetiole with a narrow, semitranslucent lamella. Postpeti- ole in profile higher than long, rounded dorsally, in dorsal view broadening from front to back. Dorsal and ventral surfaces of head, dorsum of alitrunk, as well as the sides of pronotum with numerous foveo- late punctures; space between foveolae of- ten smooth. Antennal scapes smooth, or with one to two longitudinal rugae along their margins. Petiole smooth and shining, except for a longitudinal ruga on each side. Postpetiole and gaster smooth and shining. 520 Dorsal surfaces of head and alitrunk with numerous erect or suberect short hairs. Two or more pairs of hairs present on the dorsal surfaces of petiole node and post- petiole, respectively. A few of forward-pro- jecting long hairs present near the anterior clypeal margin. Scapes and tibiae with nu- merous erect or suberect short hairs. First gastral tergite lacking erect or suberect Raine Color reddish- Broun Queen. TL 6220=72025) Ei e264" HW 1.42=1.69) CI 103-1138, SL 1.21—1.56, SI 85-92, EL 0.29-0.34, PW 1.14-1.42, Alene (4=2 1A PRWa041= 0525 PRI 036= 0.42, PPI 110-125 (n = 13). General shape as in Figures 249-250, with normal caste differences from the conspecific worker; eyes much larger, usu- ally containing more than 17 onuneuchan in the longest row; other characters similar to worker. Male (Figs. 268, 279) la AG=> 2 Se Ee 0.68—0.78, HW 0.69—0.80, CI 95-108, SL 0.30—0.42, SI 40-56, HWE 0.85—0.94, EL 0.35-0.39, PW 0.92-1.14; AL 1.44—-1.78, PPW 0.30-0.38, PPL 0.28—0.34, PPI 103— 114 (n = 5). Head in full-face view, including the eyes, broader than long. Clypeus convex, lacking a median longitudinal carina. Palp formula 1,3. ronal carinae distinct, ex- tending to the level of the posterior mar- gins of antennal insertions. Maximum di- ameter of the median ocellus 0.10 to 0.12. Scapes longer than the other antennal seg- ments, except for the apical ones. On the mesoscutum, notauli rather wide, forming a V shape, separated into several cells by narrow transverse ridges; parapsidal fur- rows absent. Scuto-scutellar sulcus wide, usually with five to six narrow longitudinal ridges. Propodeum armed with a pair of robust, triangular short spines. Metapleur- al lobes subtriangular. Middle and hind tibiae without any spurs. Petiole node in profile low, slightly longer than high, with a fairly long anterior peduncle. Postpetiole in profile rounded dorsally, in dorsal view somewhat transversely rectangular and slightly broader than long. Dorsum of Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 head with some small punctures, varying from scattered and shallow to dense and somewhat coarsely incised. Pronotum with — dense foveolate punctures. Mesonotum | with punctures, varying from a few scat- tered to numerous and from feeble, small to rather large. Propodeum rugulose. Sides of petiole with a few longitudinal rugae as well as some foveolate punctures between them. Sides of postpetiole with some shal- low foveolate punctures. Gaster unsculp- tured. All dorsal surfaces with numerous blackish-brown long hairs; sometimes hairs _ somewhat stiff. Color blackish-brown; wings dusky. Note: The previously described male is assigned to the species P. picteti for the following three reasons: (1) It belongs to the umbripennis group because it is very similar to the males of P. pollux, P. reti- culatus, and P. umbripennis but rather dif- ferent from the other known males of Pris- tomyrmex in the structure and shape of propodeal spines, petiole, notauli, and scu- to-scutellar sulcus and in body size, sculp- ture, and hairs. (2) It differs from the males of the other species of the wmbri- pennis group as follows: In the male of P. picteti, CI is 95-108, EL is 0.35—0.39, the propodeal spines are shorter, the postpe- tiole in dorsal view is slightly broader than long (PPI 103-114), and the notauli form a V shape; but in P. pollux, CI is 80, EL is 0.44, the propodeal spines are longer, the postpetiole is distinctly longer than broad (PPI 80-90), and the notauli form a Y shape. The male of P. picteti (HW 0.69— 0.80, HL 0.68—0.78, and EL 0.35-—0.39) is smaller than that of P. umbripennis (HW 0.98, HL 0.94, and EL 0.47). A distinct rugoreticulum is seen on the dorsal head and also on the mesonotum in P. reticu- latus but not so in the male of P. picteti. (3) Two male specimens were collected in Dumaguete, Philippines, by J. W. Chap- man on May 30, 1950, and each of them was originally mounted together with one worker or one queen of P. picteti, respec- tively, on the same pin. Comments. Pristomyrmex picteti occurs REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang in Papua New Guinea, Borneo, Sabah, Brunei, Malaya, Singapore, and the Phil- ippines. It is closely related to the other several species of the wmbripennis group. A discussion regarding the relationship be- tween P. picteti and P. fuscipennis is pro- vided under P. fuscipennis. Characters separating P. picteti from P. umbripennis are listed under P. umbripennis. The work- ers of P. picteti differ from those of P. pol- lux as follows: P. picteti Propodeal spines relatively short, not strongly upcurved at their apices Smaller species with HL 1.04—1.36, HW 1O2= "40" andi Sie 0:92=1-26 Dorsum of head with scattered foveo- late punctures; space between foveo- lae often smooth Anterior margin of the median portion of clypeus usually with fewer than seven denticles P. pollux Propodeal spines relatively long, strong- ly upcurved at their apices Larger species with HL 1.42-1.54, HW Zea sey emo! Si, 40S ia Dorsum of head with foveolate-reticu- late sculpture Anterior margin of the median portion of clypeus usually with seven denti- cles Material Examined (ANIC, BMHH, BMNH, LAMN, MCZC, NHMV). Papua New Guinea: Gulf Prov., Ivimka Camp, Lakekamu Basin, 7.7°S, 146.8°E, 120 m el- -evation, lowland wet forest, #96-395 (nest in wet rotten log), #96-404 (in rotted log debris) (R. R. Snelling); NETH. Genjam, 40 km W of Hollandia, 100 to 200 m (T. C. Maa); 24 km N Madang, 80 m, 5°01'S, 145°46'E, rainforest, ex rotten log (P. S. Ward); Yawasora near Wewak, <50 m, rainforest, ex rotting log (R. W. Taylor); along Kokoda Rd., rainforest, 400 to 1,000 ft (B. B. Lowery); Lae, Didiman CK., low- land rainforest (E. O. Wilson); Lae, Busu R., lowland rainforest, in rotten wood on ground (B. B. Lowery); Lae, Markham R. 521 Bridge, lowland rainforest, in log (B. B. Lowery); Lae 300 ft, above Bumbu Crk., edge of rainforest, wet gully, in rotten wood (B. B. Lowery); Tage L. Kutubu 2700 tte forest maxreiny ext leavesn (pl: Barrett); Papua, Brown Riv., lowland rain- forest, ex rotten wood fragment (R. W. Taylor). Indonesia: Irian Jaya, PT. Free- port Concession, Siewa camp., 03.04°S, 136.38°E, 200 ft, #98-86, lowland second- ary rainforest, under loose bark of dead log (R. R. Snelling); Borneo, West Kaliman- tan, Gunung Palung Nat. Pk. Cabang Panti Ress Sta 00 sto 400 mmnee el S4SaellOa ke primary forest (Datling, Rosichon, Sutris- no): Borneo, Sandakan (Baker). N. Bor- neo: Tawau, Quoin Hill, Forest Camp 1, 3 to 5 km WSW of Cocoa Res. Sta. (Y. Hir- ashima). Brunei: Belait District, Manilas, Ulu Belait (D. E. Hardy). Malaysia: Ma- laya, Gombak (B. Bolton); Pahang, Kuala Tahan 200 m (J. L. Gressitt); Sg. Patani (G. H. Lowe). Singapore (Baker). Philippines: Mt. Montalban, Rizal Wa-wa Dam, 150 to 200 m (H. M. Torrevillas); Mt. Apo 5 to 6,000 m (C. F. Clagg); N. Palawan, Bin- aluan (G. Boettcher); Palawan, 14 km S Puerto Princesa (9.44°N, 118.44°E), rain- forest 0 to 250 m (B. B. Lowery); Palawan, Iwahig Penal Col., ca. Puerto Pricesa (9.44°N, 118.44°E), coffee plantation, 60 m, edge of rainforest, foraging on ground (B. B. Lowery); Luzon, Los Banos, Mt. Makiling 600 m (14.1°N, 121.11°E), rain- forest, in very wet log (B. B. Lowery); Los Banos (F. X. Williams); Mt. Makiling (Bak- er; A. T. Cencho); Laguna, Mt. Makiling, 150 to 500 m (H. Zettel); Luzon I., Bauqui (R.C.Mcg.); Luzon, Mt. Banahao (?); Du- maguete, 4,500 ft (J. W. Chapman); Horns of Negros, 3,600 ft (J. W. Chapman); Camp (J. W. Chapman). Ecological Information. This species oc- curs in rainforest, nesting in rotten logs. It has been collected on the ground, on rot- ten wood, and under loose bark of a log. Pristomyrmex pollux Donisthorpe Figures 251-254, 269, 280, 282 Pristomyrmex pollux Donisthorpe, 1944: 83. Holo- type male, Malaysia: Penang, 6.xi.1913 (G. E. Bry- ant) (BMNH) [examined]. Figures 251-254. Pristomyrmex pollux Donisthorpe. 251A: Worker head, full-face view; 251B: Showing a transverse carina on the ventral clypeus; 252: Worker, lateral view; 253A: Queen head, full-face view; 253B: Showing a transverse ruga on the ventral clypeus; 254: Queen, lateral view. Note: This species was described only from a single male, which created the dif- ficulty of associating female castes with the male. I place the following workers under P. pollux for three reasons: (1) In MCZC, two males of P. pollux appear to belong to the same series as the following examined 28 workers bearing labels with the same locality and collector and with the absence of collecting date. (2) This male (i.e., the holotype of P. pollux) belongs to the um- bripennis group because it is very similar to the males of P. picteti, P. reticulatus, and P. umbripennis but rather different from the other known males of Pristomyrmex in the structure and shape of propodeal spines, petiole, notauli, and scuto-scutellar sulcus and in body size, sculpture, and hairs. (3) After all available males of Pris- tomyrmex are examined, it seems that, in Pristomyrmex, the propodeal armaments of the male are usually shorter than those of the conspecific worker. The male of P. pollux has a pair of well-developed pro- podeal spines (which are actually the strongest and longest among all known Pristomyrmex males). The following work- ers also possess a pair of long propodeal spines, which matches with those in the male of P. pollux closely. Diagnosis (Worker). Masticatory margin of mandible with four teeth (an apical + a preapical + two broad-based short teeth of similar size), lacking a distinct diastema; eyes small, with three to four ommatidia in the longest row; propodeal spines long, strongly upcurved at their apices. Worker. TL 6.26—-6.80, HL 1.42—1.54, REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang HW 1.42-1.58, CI 95-104, SL 1.40-1.52, SI 96-101, EL 0.06—0.08, PW 0.90—0.96, AL 1.58—1.90, PPW 0.39-0.43, PPL 0.39— 0:43, PPI 100105 (7 = 28). Mandibles with a few longitudinal ru- gae. Basal margin of ToEtRCHS with a roel short, soe wuhiat truncated, promi- nent lobe. Clypeus with a median longi- tudinal carina that does not reach the pos- terior clypeal margin. Frontal area un- sculptured. Anterior margin of the median portion of clypeus with a median denticle and three (sometimes two) others on each side. Ventral surface of clypeus with a coarse transverse carina. Palp formula 1,3. Frontal carinae strong, extending to the level of the posterior margins of eyes. Scrobal areas smooth, present lateral to the frontal carinae. Frontal lobes present, partially covering the condylar bulbs of holding Antennal scapes. Antennal scapes, when lying on the dorsal head, just beyond the occipital margin. Lamella, encircling the base of antennal scape, with a broad and deep notch on the center of the dorsal surface. Eyes small, usually with three, at most four, ommatidia in the longest row. Occipital margin straight or elke: concave in full-face view. Bronouiin unarmed. Pro- podeum with a pair of long spines that are strongly upcurved at their apices (i.e., hook Eee) and laterally compressed. Meta- pleural lobes prominent and somewhat rounded. Petiole node in profile distinctly longer than high, with a long anterior pe- Aamele: its Al terosloreal angle is on ap- proximately the same level as the poster- odorsal. In dorsal view, petiole node longer than broad. Subpetiole with a narrow, semitranslucent lamella. Postpetiole in pr ofile higher than long, rounded dorsally, in dorsal view broadening from front to back. Dorsum of head, except for scrobal areas and frontal area, with foveolate-re- ticulate sculpture. Antennal scapes with longitudinal rugae along their dorsal mar- gins. Dorsum ae alntiannk as well as the sides of pronotum with numerous foveo- late punctures that are often close to each other. Sides of the rest of alitrunk with 523 some scattered punctures. Petiole smooth and shining, but with a longitudinal ruga on each side. Postpetiole and gaster smooth and shining. Dorsal surfaces of head, alitrunk, petiole, and postpetiole with numerous erect or suberect hairs. A row of forward-projecting long hairs pre- sent near the anterior clypeal margin. Scapes and tibiae with some erect or sub- erect short hairs. First gastral tergite lack- ing erect or suberect hairs. Color yellow- brown to reddish-brown. Queen. A single queen (BMNH) was ex- amined. It was collected in Penang, Ma- laysia, by G. E. Bryant in October 1913 (ie., the collecting locality and the collec- tor name for this specimen are the same as the holotype male of P. pollux): TL ca. (oO Ely 164 AW arco s Close Siw a6: SI 92, EL 0.29, PW 1.42, AL 2.14. This queen possesses the following characters: (1) dentition of the masticatory margin and basal margin of mandible as in the previously Hcntoncd worker; (2) den- tition of the anterior clypeal margin as in the previously mentioned worker; (3 ) clyp- eus with a median carina that does not reach the posterior clypeal margin; (4) ventral surface of clypeus with a transverse ruga; (5) frontal lobes and frontal carinae as in the previously mentioned worker; (6) lamella encircling the base of antennal scape as in the previously mentioned worker; (7) antennal scapes with longitu- dinal rugae along their dorsal margins; (8) propodeum with a pair of robust, rather long spines that are longer than the dis- tance between their bases. (9) metapleural cee as in the previously mentioned work- ; (10) petiole and postpetiole as in the Saree mentioned worker; and (11) dorsum of head, except for the scrobal ar- eas, with foveolate-reticulate sculpture. In other words, except for normal caste dif- ferences and the propodeal spines that are neither laterally compressed nor upcurved at their apices, other characters are gen- erally similar to those in the previously mentioned worker. Male (Figs. 269, 280, 282), Th 5.60= 524 5.92, HL 0.85-0.88, HW 0.68—-0.70, CI 80-80, SL 0.38-0.41, SI 54-60, HWE 0.86—-0.88, EL 0.44—0.44, PW 1.26—1.36, AL 2.00—2.20, PPW 0.35—0.38, PPL 0.42— 0.44, PPI 80-90 (@ = 3). Head in profile high and thick, in full- face view, excluding eyes, much longer than broad, and including the eyes about as long as_ broad. Clypeus convex and ar chee in middle without a median longi- tudinal carina. Palp formula 1,3; maxillary palp long. Maximum diameter of the me- dian ocellus 0.12. Scapes longer than the other antennal segments, except for the apical ones. On the mesonotum, notauli rather wide and deep, showing a Y shape, separated into small cells by narrow trans- verse ridges; parapsidal furrows absent. Scuto- soutellar sulcus wide, separated into small cells by seven to eight narrow ridges. Propodeum ‘with a pair of robust, broad- based, rather long spines. Metapleural lobes prominent and somewhat rounded. Petiole node in profile distinctly longer than high with a long anterior peduncle. Postpetiole in profile “slightly longer than high with a convex dorsum, in don sal view rectangular and distinctly longer than broad. ‘Subpostpetiole with a blunt, tooth- like prominence. Dorsum of head cath nu- merous small foveolate punctures that sometimes are coarse and dense. Prono- tum with small, coarse, and dense foveo- late punctures. Mesoscutum with some scattered, small, shallow foveolate punc- tures. Mesoscutellum with dense, coarse foveolate punctures. Propodeum with some irregular coarse rugae. Middle and hind tibiae without any spurs. Each side of petiole with a coarse longitudinal ruga, a few irregular short rugae, and some foc olate punctures. Side ae postpetiole with some small, weak foveolate punctures. All dorsal surfaces with abundant erect or suberect stiff long hairs. Color blackish- brown; hairs blackish-brown; wings rather smoky. (Note: In the holotype, the right antenna is abnormal, with 11 segments, while the left one has 12.) Comments. Pristomyrmex pollux is Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 known from Malaya and Sabah. It must have evolved from a P. picteti—like ances- tor. The workers of this species can be im- mediately recognized by possessing a pair of distinct propodeal spines that are long, strongly upcurved at their apices (i.e., hooklike), and laterally compressed. This character is unique in the genus Pristo- myrmex. Material Examined (MCZC, USNM, ANIC, BMNH, NHMV). N. Borneo: Mt. Dubit, 3,000 ft (E. Mjoberg). Biological Information. Unknown. Pristomyrmex reticulatus Donisthorpe Figures 255-256 Pristomyrmex reticulatus Donisthorpe, 1949: 750. Holotype male, New Guinea: Finschhafen, 27.iv.1944 (E. S. Ross) (CASC) [examined]. Male. TL 4.86, HL 0.82, HW 0.83, CI HORS SIMOlS 47S Ale BOS Shee Wao 4s AL 1.64 (n = 1). Head, including the eyes, distinctly broader than long. Clypeus convex in mid- dle; its anterior margin transverse. On the mesoscutum, notauli forming a Y shape, with several transverse rugae. Scuto-scu- tellar sulcus wide, separated into small cells by several longitudinal ridges. Pro- podeum armed ante a pair of fairly long spines. Metapleural lobes somewhat rounded. Petiole node in profile longer than high with a long anterior peduncle. Postpetiole in profile Fouled dorsally, in dorsal view subquadrate. Dorsum of head with a developed rugoreticulum, except for a narrow, long median strip that is smooth and unsculptured. Clypeus with some longitudinal rugae. Promesonotum with strongly developed rugoreticulum. Sides of petiole with rugoreticulum. Sides of postpetiole with a few longitudinal ru- gae as well as a few superficial foveolate punctures between them. Gaster unsculp- tured, smooth, and shining. All dorsal sur- faces weft numerous long hairs; hairs stiff on the head and alitrunk. Body and. hairs blackish-brown; wings dusky. Comments and Discussion. This species, described from a single male, belongs to REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang es 5295 BBS Ax LoS ASTI 48=.06) HE 1k32=1k63: Ele s0S1h 74 G19 8109S 1202152: SIRS 92. En OM =OMSS PW OrS6—1e 6s AL 1.46=1.88, PPW 0.33-0.47, PPL 0.32-— O44 ERE 10321139 G@r = 52): Mandibles with a few longitudinal rugae that often reach to the masticatory margin. Basal margin of mandible with a central, broadly curved prominence. Clypeus smooth and shining with a strong median longitudinal carina. Anterior clypeal mar- gin usually with seven denticles: a median denticle and three others on each side; sometimes one to two lateral denticles are indistinct; sometimes, two lateral denticles are fused into a broad lobe. Ventral surface of clypeus with a coarse transverse ridge. Palp formula 1,3. Frontal carinae extend- ing to the level of the posterior margins of eyes. Antennal scrobes absent. F rontal lobes present, partially covering the con- dylar bulbs of holding antennal scapes. One third to one half of the antennal scapes usually laterally compressed near the bases. Antennal scapes, when lying on the dorsal head, just beyond the occipital margin. Lamella encircling the base of the antennal scape with a broad and deep notch on the center of the dorsal surface. Eyes generally containing more than 20 (rarely 15) ommatidia, with six to seven (rarely five) in the longest row. Occipital margin straight or feebly concave in full- face view. Alitrunk, in profile, with a con- vex dorsum, in dorsal view without any su- REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 527 tures. Pronotum unarmed. Propodeum with a pair of slightly elongate triangular teeth. Metapleural lobes prominent and rounded. Petiole node in profile longer than high with a long anterior peduncle; its anterodorsal angle is on approximately the same level as the posterodorsal. Sub- petiole with a narrow, semitranslucent la- mella. Postpetiole in profile higher than long, rounded dorsally, in dorsal view slightly broadening from front to back. Dorsal surfaces of head and alitrunk and the sides of pronotum with scattered fo- veolate punctures, varying from shallow, small, and few to distinct, rather large and many; space between foveolae usually smooth. Petiole, postpetiole, and gaster smooth and shining. Dorsal surfaces of head and alitrunk with some erect or sub- erect short hairs. Usually two to three pairs of hairs present on the dorsal surfaces of petiole node and postpetiole, respectively. A row of forward-projecting hairs present near the anterior clypeal margin. Scapes and tibiae with numerous suberect short hairs. First gastral tergite lacking erect or suberect hairs. Color reddish-brown, but the masticatory margin of mandible black- brown. Queen. TL §.00=8.25, AL 1.80—2.02, BIW 21002=222 55 C99 SS Sw o2 16a) Si 7S Ws), 1BIL, O4O O42, JEW. IL XO EGO), AL 2.26—2.40, PPW 0.52—0.52, PPL 0.48— 0.50, PPI 104-108 (n = 4). General shape as in Figures 259-260, with normal caste differences from the conspecific worker; propodeal armaments toothlike: other characters similar to work- er. Male. A single male specimen (BMNH), collected in Papua (Kokoda, 1,200 ft) by L. E. Cheeman in August 1933, has the following measurements: TL ca. 6.04, HL 0.94, HW 0.98, CI 104, SL 0.35, SI 36, EL 0.47, PW 1.28, AL 1.98. Head, including the eyes, distinctly broader than long. Clypeus narrow, trans- verse, convex in the middle, its anterior margin almost straight. Frontal carinae short. Eyes large and prominent. On the mesoscutum, notauli rather wide, forming a Y shape with several coarse rugae. Scuto- scutellar sulcus wide, separated into small cells by longitudinal ridges. Propodeum armed with a pair of strong triangular teeth. Metapleural lobes rounded. Petiole node in profile low, distinctly longer than high, with a long anterior peduncle. Post- petiole in profile slightly longer than high and rounded dorsally. Dorsum of head be- hind the level of the posterior margins of eyes with some foveolate-reticulate sculp- ture, but the centrical disc of the dorsal head, under the median ocellus, smooth and shining. Each side of the dorsal head, between the eye and the frontal carina, with several transverse rugae; spaces be- tween the rugae smooth and shining. Pro- mesonotum with somewhat coarse foveo- late-reticulate sculpture. Sides of petiole node with foveolate-reticulate sculpture. Postpetiole rather smooth, only with few superficial short rugae on each side. Gas- ter unsculptured, smooth, and_ shining. Body blackish-brown, but gaster and scapes reddish-brown. Note: This male is assigned to the spe- cies P. umbripennis for the following rea- sons: (1) It belongs to the wmbripennis group because it is similar to the males of P. picteti, P. pollux, and P. reticulatus but different from the other known males of Pristomyrmex in the structure and shape of propodeal spines, petiole, notauli, and scuto-scutellar sulcus and in body size, sculpture, and hairs. (2) It cannot be placed in the other species of the umbri- pennis group. It differs from the male of P. pollux because the former has a wider head (HW 0.98, CI 104) than the latter (HW 0.68—0.70, CI 80). This male is larger (HW 0.98, HL 0.94, EL 0.47) than the males of P. picteti and P. reticulatus (HW < 0.85, HL < 0.85, EL < 0.40). In fact, it is the largest male specimen so far found in Pristomyrmex. This matches with the workers and queens of P. umbripennis, which are the largest in the genus. (3) The collecting locality and the collector name 528 for this male are the same as the other two queens of P. umbripennis. Comments. Pristomyrmex umbripennis occurs in New Guinea and some islands of Indonesia. It is closely related to P. pollux and P. picteti, but their workers and queens can be separated by the following characters: P. umbripennis Eyes larger, generally consisting of 20 or more ommatidia and containing six to seven (rarely five) ommatidia in the longest row (worker) Propodeum with a pair of triangular toothlike armaments that are shorter than the distance between their bases (worker and queen) One-third to one-half of the antennal scape usually laterally compressed near the base (worker and queen) P. pollux and P. picteti Eyes smaller, generally consisting of 10 or less ommatidia, and containing two to three (rarely four) ommatidia in the longest row (worker) Propodeum with a pair of fairly long or long spines that are longer (or much longer) than the distance between their bases (worker) or with a pair of short spines (queen) Antennal scape not laterally compressed near the base (worker and queen) Material Examined (ANIC, MCZC, LAMN, BMNH, BMHB). Papua New Guinea: Gulf Prov., Ivimka Camp, Lake- kamu Basin, 7.7°S, 146.8°E, 120 m, #96- 266 (R. R. Snelling); NETH. Santani, 90+ m (T. C. Maa): NE Eloa River 488 to 518 m (S. Cutleck); N. Dist. of Papua, Kokoda (P. M. Room); N. Dist. of Papua, Saiho (P. M. Room); N. Dist. of Papua, Lejo Rd (P. Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 M. Room); Papua, Brown Riv., lowland rainfor., ground, rotten tree stump (R. W. Taylor); Maffin Bay (E. S. Sepik Province, ca. 2 to 3 km S of Wirui, S of Wewak 50 tO ICO) tm, ORBES, WaS.S7le CR. J. Ko- hout); Lae, Bupu River, wet rotten log (B. B. Lowery); Bulolo, 3,000 ft, rainforest, in rotten log (B. B. Lowery); near Popondet- ta, <50 m, rainforest, ex trunks and low foliage (R. W. Taylor); 4 mi S of Popon- detta, rainforest (B. B. Lowery); Sangara N.D. (G. Baker). Indonesia: Amboina (Bird); Irian Jaya, 50 km S Manokwari, Ar- fak Mtns. Nature Reserve 25 m, second rainforest, ex log (G. D. Alpert); Irian Jaya, PT. Freeport Concession, Siewa camp., 03.04°S 136.38°E, 200 ft, lowland second- ary rainforest, #98-62 and #98-86, under loose bark of log (R. R. Snelling); Seram I., Solea (M. Brendell). Ecological Information. This species oc- curs in rainforest and has been collected in and on rotten logs and under loose bark of a log. NOMEN NUDUM Pristomyrmex parvispina Emery Emery (1900) mentioned the name Pristomyrmex parvispina but provided neither an indication such as “n.sp.” (or its equivalent, ele, Spe nov. 9)snor a descrip- tion and designated no types. Pristomyr- mex parvispind 1s thus a nomen nudum. Emery (1922: 233) cited P parvispina Emery as a synonym of P. brevispinosus Emery, but this was incorrect because P. parvispina was not a valid name. In addi- tion, Emery (1922: 233) cited the original publication date incorrectly as 1901 [see Bolton’s (1995) catalog for the dating of the original paper]. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 5 Figure 261. Head of the male of Pristomyrmex brevispinosus Emery, full-face view. Figure 262. Head of the male of Pristomyrmex ?flatus (see the text), full-face view; mandible indicated by an arrow. 530 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 a al we : Figure 263. Head of the male of Pristomyrmex longispinus sp. n., full-face view. Figure 264. Head of the male of Pristomyrmex orbiceps (Santschi), full-face view. 266 0.1 mm Figure 265. Head of the male of Pristomyrmex quadridentatus (André), full-face view. Figure 266. Head of the male of Pristomyrmex sulcatus Emery, full-face view. OX >) 1 532 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figure 267. Head of the male of Pristomyrmex obesus Mann, full-face view. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 533 Figure 268. Head of the male of Pristomyrmex picteti Emery, full-face view. Figure 269. Head of the male of Pristomyrmex pollux Donisthorpe, full-face view (antennae missing). 53: Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figure 270. Male of Pristomyrmex brevispinosus Emery, lateral view. Figure 271. Male of Pristomyrmex ?flatus (see the text), lateral view. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 535 1 mm Figure 272. Male of Pristomyrmex longispinus sp. n., lateral view. Figure 273. Male of Pristomyrmex orbiceps (Santschi), lateral view. 536 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 Figure 274. Male of Pristomyrmex quadridens Emery, lateral view. Figure 275. Male of Pristomyrmex quadridentatus (André), lateral view. Figure 276. Male of Pristomyrmex sulcatus Emery, lateral view. REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 537