PELEGRINA FRANGANILLO AND OTHER JUMPING FORMERLY PLACED IN THE (ARANEAE: SALTICIDAE)

WAYNE P. MADDISON'

CONTENTS

Abstract 216 2. proxima (G. & E. Peckham, Introduction 217 1901) 265 Acknowledgments 218 3. Pelegrina dithalea new species 268 Materials and Methods 219 4. Pelegrina edrilana new species 269 Explanation of Morphological and Behavioral 5. (Walckenaer, 1837) __ 270 Terms 222 6. Pelegrina peckhamorum (Kaston, The Subfamily Dendryphantinae 226 1973) 272 Relationships within the Dendryphantinae 231 7. Pelegrina neoleonis new species 273 The Group of Genera 232 8. Pelegrina tristis new species 274 Two Genera That Have Exchanged Species 9. Pelegrina sabinema new species 275 with Metaphidippus: 10. Pelegrina pervaga (G. & E. Packham, and Beata 235 1909) 276 The Proper Placements of Metaphidippus 11. Pelegrina kastoni new species 277 Species 237 12. Pelegrina flavipedes (G. & E. Peckham, Phanias F. P.-Cambridge, 1901 238 1888) 278 Terralonus new genus 239 13. Pelegrina flaviceps (Kaston, 1973) 279 Ghelna new genus 239 14. Pelegrina exigua (Banks, 1892) 281 The Genus Pelegrina Franganillo, 1930 240 15. Pelegrina montana (Emerton, 1891) 283 Phylogeny within Pelegrina 245 16. Pelegrina insignis (Banks, 1892) 284 Identifying Species of Pelegrina and the 17. Pelegrina chaimona new species 286 Metaphidippus mannii Group 247 18. Pelegrina clemata (Levi & Levi, 1951) .. 287 Key to the Males of All Species of Pelegrina 19. Pelegrina aeneola (Curtis, 1892) 288 and Those Metaphidippus mannii 20. Pelegrina halia new species 290 Group Species Occurring in the United 21. Pelegrina chalceola new species 291 States 248 22. Pelegrina furcata (F. P.-Cambridge, Key to the Female Pelegrina of the Eastern 1901) 292 United States and (East of the 23. Pelegrina volcana new species 294 Mississippi River and Manitoba) 257 24. Pelegrina bicuspidata (F. P.-Cam- Key to the Female Pelegrina of the Great bridge, 1901) 295 Plains (between the Rocky Mountains 25. Pelegrina ochracea (F. P.-Cambridge, and the Mississippi River) 258 1901) 295 Key to the Female Pelegrina of the Pacific 26. Pelegrina morelos new species 296 Coast of the United States and Western 27. Pelegrina huachuca new species 296 Canada 258 28. Pelegrina arizonensis (G. & E. Peck- Key to the Female Pelegrina and mannii ham, 1901) 297 Group of Arizona 259 29. Pelegrina helenae (Banks, 1921) 298 Key to the Pelegrina and females 30. Pelegrina verecunda (Chamberlin & of and 260 Gertsch, 1930) 299 Descriptions of the Species of Pelegrina 262 31. Pelegrina clavator new species 300 1. (Walckenaer, 1837) __ 263 32. Pelegrina pallidata (F. P.-Cambridge, 1901) 301 33. Pelegrina variegata (F. P.-Cambridge, ' Biol- 1 901 302 Department of Ecology and Evolutionary ) ogy, University of Arizona, Tucson, Arizona 85721. 34. Pelegrina yucatecana new species 303

Bull. Mus. Comp. Zool., 154(4): 215-368, January, 1996 215 216 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

35. Pelegrina sandaracina new species 304 cussed, and the harfordii group is transferred to the 36. Pelegrina tillandsiae (Kaston, 1973) 305 genus Phanias F. P. -Cambridge, which appears to be 37. Pelegrina bunites new species 306 relatively distantly related to most other dendry- 38. Pelegrina orestes new species 307 phantines. The following new combinations are es- The Genus Nagaina G. & E. Peckham, 1896 ... 308 tablished: Phanias albeolus (Chamberlin & Ivie), 39. Nagaina incunda G. & E. Peckham, Phanias concoloratus (Chamberlin & Gertsch), 1896 308 Phanias dominatus (Chamberlin & Ivie), Phanias Species of the mannii Group of the United furcifer (Gertsch), Phanias furcillatus (F. P. -Cam- States and Canada 310 bridge), Phanias harfordii (G. & E. Peckham), Phan- 40. Metaphidippus mannii (G. & E. Peck- ias monticola (Banks), Phanias neomexicanus (Banks), ham, 1888) 310 and Phanias watonus (Chamberlin & Ivie). Also re- 41. Metaphidippus diplacis (ChamberUn, moved from Metaphidippus are the mylothrus and 1924) 312 castaneus groups, for which the new genera Terra- 42. Metaphidippus tricolor ChamberUn & lonus and Ghelna are described, thus yielding the Ivie, 1941 314 new combinations Terralonus californicus (G. & E. 43. Metaphidippus chera (ChamberHn, Peckham), Terralonus mylothrus (Chamberlin), Ter- 1924) 315 ralonus unicus (Chamberlin & Gertsch), Terralonus 44. Metaphidippus carmenensis (Chamber- shaferi (Gertsch & Riechert), Terralonus versicolor Hn, 1924) 316 (G. & E. Peckham), Terralonus vittatus (Banks), Ter-

45. Metaphidippus emmiltus new species .... 317 ralonus fraternus (Banks), Ghelna castanea (Hentz), Literature Cited 318 Ghelna barrotvsi (Kaston), Ghelna sexmaculata Index 322 (Banks), and Ghelna canadensis (Banks). The new combination Sassacus paiutus (Gertsch) is estab- lished. The vitis group is retained in Metaphidippus. Abstract. The genus Pelegrina Franganillo con- The limits of the genera Dendryphantes C. L. Koch also reconsidered. tains 38 species of dendryphantine jumping spiders and Beata G. & E. Peckham are from North and Central America that were formerly The combination Dendryphantes nigromaculatus placed in the genus Metaphidippus F. O. Pickard- Keyserling is revived and the following combinations Cambridge. The close relatives of the Dendryphan- established: Beata hispida (G. & E. Peckham), Beata tinae may include the Europhryinae and several inconcinna (G. & E. Peckham), Beata maccunii (G. smaller groups, for they share an embolus that is & E. Peckham), and Beata rustica (G. & E. Peck- coiled counterclockwise (left palp) and separated from ham). Dryphias (G. & E. Peckham) is synonymized the tegulum by a fully expandable hematodocha. The with Beata. subfamily Dendryphantinae itself is delimited by the The largest group removed from Metaphidippus derived conditions of a carina on the underside of is placed in the genus Pelegrina Franganillo, 1930, the male chelicera, the coil of the embolus folded whose species are, with some exceptions, distin- back so as to be hidden behind the base of the em- guished from other dendryphantines by the presence bolus, and S-shaped epigynal openings. of two terminal rami retrolateral to the embolus open- Within the subfamily, generic relationships are ing, an embolic hematodocha that bulges distally, poorly understood, but it is clear that the genus Me- wrinkles on the anterior margin of the male cheliceral taphidippus is polyphyletic. The genus should in- fang, a distinct band of pale scales on the side of the clude at most a few species closely related to the face, and male courtship with the first legs held low neotropical genera G. & E. Peckham, Bag- and forward. The following species are moved into heera G. & E. Peckham, and Gastromicans Mello- Pelegrina: P. aeneola (Curtis), P. arizonensis (G. & Leitao. Gastromicans is removed from synonymy with E. Peckham), P. bicuspidata (F. P. -Cambridge), P. Beata G. & E. Peckham. The following new com- clemata (Levi & Levi), P. exigua (Banks), P.flaviceps binations are established for species in this group: (Kaston), P.flavipedes (G. & E. Peckham), P.furcata Bagheera prosper (G. & E. Peckham), Messua cen- (F. P. -Cambridge), P. galathea (Walckenaer), P. he- tralis (G. & E. Peckham), Messua dentiger (F. P.- lenae (Banks), P. insignis (Banks), P. montana Cambridge), Messua donalda (Kraus), Messua lata (Emerton), P. ochracea (F. P.-Cambridge), P. palli- (Chickering), Messua laxa (Chickering), Messua lim- data (F. P.-Cambridge), P. peckhamorum (Kaston), bata (Banks), Messua moma (F. P. -Cambridge), Mes- P. pervaga (G. & E. Peckham), P. proterva (Wal- sua octonotata (F. P. -Cambridge), Messua pura ckenaer), P. proxima (G. & E. Peckham), P. tilland- P. (Bryant), Messua trident ata (F. P. -Cambridge), Gas- siae (Kaston), P. variegata (F. P.-Cambridge), and tromicans albopilosa (G. & E. Peckham), Gastromi- verecunda (Chamberlin & Gertsch). Pelegrina prox- cans hondurensis (G. & E. Peckham), Gastromicans ima is shown to be a senior synonym of Pelegrina levispina (F. P. -Cambridge), Gastromicans noxiosa geniculata Franganillo, the latter being the types spe- (Simon), and Gastromicans vigens (G. & E. Peck- cies of Pelegrina. A neotype is designated for Attus ham). The combination Messua desidiosa G. & E. galathea Walckenaer. Seventeen species are de- Peckham is revived. The proper placement of various scribed as new: P. balia, P. bunites, P. chaimona, P. groups currently assigned to Metaphidippus is dis- chalceola, P. clavator, P. dithalea, P. edrilana, P. Pelegrina Jumping Spiders • Maddison 217

luiaclnica, P. kastoni, P. morelos, P. neoleonis, P. about half of the nearly 300 species of sal- orestes, P. sabinctua, P. sandaracina, P. tristis, P. ticids north of Mexico volcano, and P. ijucaiecana. Eitophnjs leucophaea C. occurring (accord- to the count of L. Koch, Icitis crassiventer Keyserling, and Meta- ing Richman and Cutler, pJiidippiis digitatus F. P. -Cambridge are newly syn- 1978). In the last three decades, increased on\mized with P. galathea; Dendrijphantes uteaniis interest in the family has resulted in re- Chamberhn & Gertsch with P. aeneola; and Dendry- visions of Habronattus (Griswold, 1987), phantes mimiis Chamberhn with P. furcata. Eu- in and ophrys concolor Banks is removed from synonymy (Edwards, preparation), other w ith P. proterva and considered a senior synonym of genera (Proszynski, 1968, 1971a, Sittacus cursor Barrows, yielding the new combina- 1973a, 1980; Cutler, 1981a, 1987; Rich- tion Sittictis concolor. Identification are keys pre- man, 1981, 1989). However, except for sented for all Pelegrina males and for females from works by Kaston (1973) on some eastern restricted geographical regions. All species are de- scribed and illustrated. Male/female associations were species and by Cutler and Jennings (1985) achieved for all species north of Mexico. Courtship on the arizonensis group, Metaphidippus behavior is described for 22 species of Pelegrina, has remained unre vised, perhaps because for 10 and habitat information karyotypes species, its poorly defined limits have made the for most species. scope of any revision trouble- The genus Pelegrina may be closely related to the potentially some. I Metaphidippus mannii group, Nagaina and/or Eris. When first began to revise Meta- Nagaina incunda G. & E. Peckham is described and phidippus, I knew that I would have to G. & E. Peck- illustrated; Dendryphantes vegetns restrict the revision to only some of the ham, Metaphidippus flavoUneatus F. P. -Cambridge, disparate groups placed there. The largest and Metaphidippus expallidatus F. P.-Cambridge are in synonymized with N. incunda. The species of the group placed Metaphidippus, including mannii group (temporarily retained in Metaphidip- the species most commonly collected in pus) that occur in the United States are also described northern and eastern , was and illustrated; two new combinations, Metaphidip- chosen for revision and is here moved to pus chera (Chamberhn) and Metaphidippus car- the genus Pelegrina Franganillo. nienensis (Chamberhn), are established; one species, The in Metaphidippus emmiltus, is described as new; Den- jumping spiders placed Pelegri- dryphantes versicolor G. & E. Peckham is synony- na are medium-sized dendryphantines mized with Metaphidippus mannii (G. & E. Peck- distributed throughout North America, ham), and Metaphidippus franciscanus Schenkel with with some species extending as far south Metaphidippus dtplacis (Chamberhn). as . The 38 species include the well- known P. galathea, P. proterva, P. fla- and P. aeneola. Males of INTRODUCTION vipedes, Pele- grina are generally brown with white For about 50 years after the Peckham's stripes (Fig. 1), and most can be distin- (1909) revision of the jumping spe- guished from other dendryphantines by cies north of Mexico, taxonomic work on the wide embolus with two rami retrola- North American representatives of this teral to the opening (Fig. 3). The spotted large family consisted mostly of scattered females (Fig. 2) have large thickened flaps species descriptions by Chamberlin, over the epigynal openings (Fig. 4). Al- Gertsch, Ivie, and others. Some generic re- though the eastern species were well stud- visions consolidating and clarifying the ied by Kaston (1973), most of the species previous work began appearing in the occur in the western United States, Mex- 1950s (Gertsch and Ivie, 1955; Barnes, ico, and Central America, and they re- 1955, 1958), but most genera remained ceived their last comprehensive treatments untouched, including the three largest by G. & E. Peckham (1909) and F. O. genera, Habronattus* Phidippiis, and Pickard-Cambridge (1901). Many of the Metaphidippus, which together include western species have been inadequately described and illustrated, often from only one sex, making identification almost im- Authors of scientific names are given in the index. possible by anyone other than an araneol- 218 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

ogist familiar with the group. Many spe- encouragement that allowed me to grow cies in the southwest were undescribed, on my own. My other friends and col- and for most species there is little pub- leagues at Harvard and in our spider lab- lished information on natural history. The oratory —Leticia Aviles, Anna Weitzman, present revision has as its main goal to Mark Moffett, Jonathan Coddington, Mark make the species known, by describing and Stowe, Jackie Palmer, Cecile Villars, John illustrating them, their courtship displays, Hunter, Caty Sibble, Ardis Johnston, Laura and their habitats. Although much prog- Leibensperger, Dee Woessner, and Ann ress has been made in distinguishing spe- Blum—made a very supportive commu- cies and matching males to females, many nity in which to work. Leticia Aviles and problems of geographical variation and Christopher Maddison gave me much aid uncertain male-female matching remain and inspiration during the most difficult for future work, especially among Mexi- stages of the project, can Pelegrina. H. W. Levi, L. Aviles, D. Maddison, G. In addition to the Pelegrina species, Na- B. Edwards, P. F. Stevens, and E. O. Wil- gaina incunda and the U.S. species of the son read and commented on this paper. Metaphidippus mannii group are de- Two anonymous reviewers gave useful scribed because they could very well be suggestions. For discussion of jumping spi- confused with species of Pelegrina and be- der issues, I thank B. Cutler, G. B. Ed- cause their is in need of revision, wards, D. B. Richman, J. Proszynski, F. This work addresses the phylogeny of Wanless, and P. Wijesinghe. B. Cutler, G. Pelegrina and the subfamily Dendry- B. Edwards, A. Moldenke, and D. B. Rich- phantinae, but it has no pretensions of be- man have sent live specimens to me for ing a comprehensive or modern phyloge- courtship observations. Brent Opell helped netic treatment. My phylogenetic goals are with trypsin clearing. H. D. Cameron gave to propose some characters that might pro- me helpful advice on the formation of vide an outline of dendryphantine rela- names, though he is not to be held re- tionships, focusing on the question of the sponsiblefor any ill formed, especially since monophyly of Pelegrina and a few other I did not always follow his no doubt proper groups formerly placed in Metaphidippus. advice. Curators at various institutions lent I hope that this and the basic exploratory, me specimens; a list can be seen under species-level taxonomic work will provide Materials and Methods. I thank these cu- the groundwork for future phylogenetic rators for their help. I would especially like treatments. to thank Luis F. de Armas of the Instituto de ACKNOWLEDGMENTS Ecologia y Sistematica, Havana, , for sending the Franganillo collection of This paper formed part of a Ph.D. thesis salticids. for the Department of Organismic and This work was supported in part by a Evolutionary Biology, Harvard Universi- NSERC (Canada) Postgraduate Fellow- ty. The work described in it has roots many ship. For allowing me to participate in a years deep, and during these years I re- major collecting trip to Mexico, I thank S. ceived the help of many friends and col- B. Peck and R. S. Anderson. The trip was leagues. As always, David Maddison has funded by NSERC and had assistance from been foremost among my collaborators in the Universidad Nacional Autonoma de formulating ideas, on collecting expedi- Mexico and Harvard University. The De- tions, on behavioral observations, and in partment of Organismic and Evolutionary many other activities. My thesis advisor. Biology of Harvard supported trips to Cal- Herbert W. Levi, provided a well-orga- ifornia, Arizona, and Spain. Publication nized laboratory in which to work, a model costs of this study were covered in part by of dedication to a the Colles fund. , and helpful Wetmore Pelegrina Jumping Spiders • Maddison 219

MATERIALS AND METHODS white silicone rubber (bathtub caulking). Collections Examined. The taxonomic The silicone rubber is superior to paraffin revision is based on specimens in the fol- for most purposes, for it can hold even lowing collections. The abbreviation for minuten pins firmly and later heal, and it the collection is followed by the name of offers the advantage over sand of allowing the collection and the curator and others appendages to be pinned open. Palpi were on responsible for aiding in loaning the ma- mounted Vaseline in an alcohol-filled terial, to whom many thanks are due: depression slide with a coverslip and drawn at 100 X and 200 x under an Olympus AMNH American Museum of Natural BH-2 compound microscope using inci- History, New York (N. Platnick, dent fiber-optics illumination and a cam- L. Sorkin) era lucida. Not only did the use of a com- BMNH The Natural History Museum, pound microscope allow higher resolution, London (P. Hillyard) but also the axial light path prevented the CAS California Academy of Sci- drawing difficulties caused by the side-to- ences, San Francisco (W. Pu- side shifting of the image that occurs when lawski, D. Ubick) focusing on a stereo dissecting microscope. DU Darrel Ubick personal collec- For the external (ventral) view, epigyna tion were examined using the same technique, lESC Institute de Ecologia y Siste- without clearing. Epigyna were dissected matica, Havana (Luis F. de Ar- off of the specimen to allow for the small mas) working distance of the compound micro- MCZ Museum of Comparative Zool- scope. The Vaseline on which they were ogy, Cambridge (H. Levi) mounted was made opaque by mixing with MSUW Midwestern State University, chalk dust, in order to simulate the cream- Wichita Falls, Texas (N. Hor- colored muscles and glands that would un- ner) derly the epigynum on an intact specimen. TXAM Texas A&M University, College After examination of the specimen. Vas- Station, Texas (A. Dean) eline was removed by a xylene rinse. The UCB University of California, Berke- oblique drawings of the male carapace and ley (E. SchUnger, C. Griswold) were made mostly under the UWBM Burke Museum, University of compound microscope, at 40 x. Most Washington, Seattle (R. Craw- drawings of the female abdomen were ford) made under a Zeiss stereo dissecting mi- WPM W. Maddison personal collec- croscope with a camera lucida. The draw- tion ings of the male face and female abdomen ZMB Zoologishes Museum Berlin (M. show the appearance in alcohol. Most Moritz, S. Fischer) drawings were done on coquille board with ink, a Conte drawing pencil, and white Note that Canadian specimens are, in gen- paint. Small labels with my initials (WPM) in revision eral, underrepresented this be- and the year drawn (e.g., 84) were placed cause two major collections of Canadian in vials of specimens illustrated. Photo- spiders, the Canadian National Collection graphs of living specimens were made with at the Biosystematics Research Centre, Ot- a standard 55-mm lens reversed on exten- tawa, and the Royal Ontario Museum col- sion tubes to yield approximately 2.5 x lection, were not examined due to time magnification on Kodak Technical Pan or limitations. Kodachrome film, using illumination by Routine Examination and Illustra- flash. Measurements of carapace length, tions. Specimens were examined in a glass carapace width, and body length (Galiano, dish with a bottom layer of half black, half 1963; Wanless, 1978) were made from the 220 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

dorsal view using an eyepiece reticle on a fixed, 80% ethanol fixed, and Kahle's fixed Leitz stereo dissecting microscope. Gen- specimens were all used. Specimens not erally only about five specimens of each fresh-killed were first rinsed in water for sex were measured, because little reliance several hours before digestion. The body on these measurements was made in this parts were separated to allow penetration study. They are intended to give only a of the trypsin. The trypsin solution was general idea as to the size and proportions made from about 1 ml purified trypsin of the species. The results are presented as (Fisher #T-.360) in 10 ml water, filtering follows: minimum (median) maximum, after mixing. The trypsin was warmed un- Thus, if five measurements for carapace der a light bulb during digestion. After length for the females of one species are digestion, the parts, especially dark, heavi- 2.0, 2.1, 2.2, 2.3, and 2.3, these would be ly sclerotized palps or epigyna, were = reported as follows: 2.0(2.2)2.3, n 5. bleached. They were first rinsed in water Descriptions. A species description was and then 80% ethanol, then moved to a originally written from a sample of five solution of 1 part ethanol : 1 part 10% Aero- males and five females or more (if avail- solfor a day, and then for bleaching moved able). During subsequent identifications, into a solution of 1 part 30% hydrogen the description was periodically checked peroxide : 1 part ethanol : 2 parts 10% Aero- to ensure that it covered the range of vari- sol. Aerosol (Fisher) was added to the ation within the species. Two characters bleaching solution to inhibit the formation described that were less thoroughly sam- and buildup of bubbles, which otherwise pled are the exact region of contact of the can fill the body part and make it unusable, forehead band with the anterior median After about 1 day of bleaching, the body eyes, which was examined in only four to part was moved to 80% ethanol and then eight males, and the details of the internal transferred to 95-100% ethanol. The body epigynal ducts of the female, which in part was then mounted temporarily in many species was examined in only one to clove oil or permanently in Euparal. For three females. most specimens, the transferral to Euparal Clearing. Clearing was used for detailed was accomplished by placing the body examination of the integument, especially parts in Euparal thinned with either eth- to observe external and internal structures anol or Euparal Essence and letting the of the genitalia and mouthparts. The var- ethanolor essence evaporate off, thus grad- ious body parts were cleared by placing ually taking the specimen through a series them in warm trypsin solution for 1-2 days of stronger Euparal solutions. This was most to digest internal tissues. When the tryp- easily done directly on the microscope slide, sin-clearing procedure is successful, it re- which was then let to dry partially in a veals palp morphology to a level of detail dust-free area. Drying thickened the Eu- not previously published (Figs. 3, 16-27). paral, allowing final positioning of the parts Trypsin was used instead of potassium or before adding the coverslip. After the cov- sodium hydroxide because it damages the erslip is added, the body parts, especially cuticle very little and has shown no ten- the palpus, may move. Thus, the palp was dency to expand the palp except perhaps placed near the edge of the coverslip to when hematodochae are tightly coiled as allow repositioning using a microneedle in Ashtabula and Bagheera. If the tissues slipped under the coverslip. of the specimen are well fixed and firm. Expansion of Palps. Palps were best ex- then digestion will be very slow. Hence, panded permanently by boiling the spec- for best results the specimen should be imen alive and then fixing it in dilute fresh-killed (not fixed) or "fixed" in a poor Kahle's fluid to harden it in an expanded fixative with a low concentration of alco- position followed by gradual dehydraton hoi. Fresh-killed, dilute ethanol-triton to 80% alcohol. This technique is much Pelecrina Jumping Spiders • Maddison 221

like tl.at described = by Sadana (1971). Palps back and forth at tips (n 1), = vigorously so prepared are resistant to contraction and (ca. 5 c/s) (n 1) but at low = amplitude (n can be critical-point-dried successfully 5, 33)." The parenthetical comments in- A few that (Figs. 7, 9). palpi were already dicate the number of observations (n) and preserved in alcohol were expanded in a the number of males in which the feature few minutes b\ placing them in a hot mix- was noted. The number of observations ture of 15% hydrogen peroxide, 40% wa- was counted as follows: a male was ob- and 45% ethanol. ter, served doing a bout of courtship display; Chromosomes. Chromosomes were ex- any features of position or motion were amined using the Feulgen technique as considered to have thus been observed once described by Maddison (1982). The results during this bout. If the male stopped dis- are given under the description of each playing because the female left or rejected species examined. him, and if he began again later (perhaps Courtship. Courtship observations were after I had moved them back together), obtained for Pelegrina species and nu- then the next display was counted as a merous other dendryphantines. Specimens separate observation. While the more ob- were examined within a few days after vious features of the display may have been collecting. A male and a female were observed many times (for instance, that the placed on a cotton beating sheet, usually legs were flickered was observed 12 times not in full sunlight, and manipulated until in six males in the preceding example), the male faced the female and began dis- some of the more subtle details of the dis- play. Behavior of the male was observed play were not noticed in most displays and, by eye, and notes and still photographs thus, would have been observed only a few were taken. Female behavior was not re- times (for instance, that the legs flickered corded. For most species, no filming or "alternately back and forth at tips" was videotaping was done. As salticid behavior observed only once). Where there is vari- can be fast, it is difficult to take notes ac- ation, the description lists each of the al- curately, and there are likely errors in ob- ternatives with an indication of sample servation, especially of subtle differences sizes. For instance, if the legs were usually in timing and positions. Still, consistency flickered at low amplitude, but occasion- of observations and videotape confirma- ally at high amplitude or not at all, the tion of my own previously taken notes for description might read like this: "On each = Pelegrina dithalea, Metaphidippus man- series legs flickered (n 9, 13) noticeably = = nii, Metaphidippus chera, and Phanias (n 1) or with fairly low amplitude (n = watonus all indicate that the descriptions 7, 33) or perhaps not at all (n 3, 13)." should be generally accurate. There are a Sample sizes are in general small. It was number of observations, such as the alter- felt that it is presently more important to a nate palp waving in the Pelegrina fla- obtain broad survey, including many vipedes group and the triangular crouch species, than a deep analysis of a few spe- pose of Pelegrina aeneola, that have been cies. Explanations of terms used to describe repeated a number of times and in which courtship are given in the section Expla- I have strong confidence. In the descrip- nation of Morphological and Behavioral tions of the displays, the sample sizes for Terms and in the description of behavioral the observations are indicated by listing characters (item 7) supporting the mono- the number of observations for each fea- phyly of Pelegrina. ture of the display. For instance, in the Phylogenetic Analysis. Though this description of the courtship of P. galathea, work is primarily concerned with the ge- the following sentence occurs: "First legs nus Pelegrina, an attempt was made to = = flicker (n 12, 63) on each series (n 4, outline the broader structure of the family = 23) up and down (n 4, 25) and alternately and the relationships of dendryphantines, 222 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

partly for their own sake and partly to set vipedes, flaviceps, and exigua were main- the context for the genus Pelegrina. The tained distinct despite apparent hybrid- general discussion of phylogeny within the ization because they differ in numerous family is presented separately (Maddison, features. The two sympatric forms of ex- 1988, unpublished manuscript). My phy- igua were left under one specific name logenetic proposals are presented in a nar- until they can be better studied. rative discussion of groups and characters; Male /Female Matching. Care had to be no numerical phylogenetic analysis was taken in proposing which males and fe- done. Table 2 shows the distribution of males belonged together in the same spe- some of the more important characters, cies, as in other spiders (Levi, 1985). The but it will be noted that some of these do problem was especially bad in Pelegrina not perfectly support the groups proposed. species from the southwestern United Problems with some of the characters are States, Mexico, and Central America, noted in the phylogenetic discussion. Fur- where collecting has been limited. Despite thermore, the presumption of ancestral the strong , similarity state for a given character is usually not in body form and markings could often be accompanied by rigorous outgroup anal- used as evidence. Other criteria used were ysis (e.g., Maddison et al., 1984). We are expected correlations in genitalia (e.g., still making only preliminary sketches of wide, robust embolus with strong epigynal the phylogenetic structure of this large and flaps), co-collecting in same geographical poorly known family, and it would not be region, locality, or microhabitat, and sim- productive to delay phylogenetic hypoth- ilarity of the male and female each to those eses until they can be rigorously defended. of another well-matched species. Com- The suggestions made should have at least ments regarding evidence used to match a glimmer of truth and will, I hope, stim- males and females are given in those spe- ulate future work. cies descriptions where it seems needed. Species Distinctions. Populations were Among the less certain matchings are those considered distinct species if several con- for Pelegrina sandaracina, pallidata, chai- sistent and discrete morphological differ- mona, huachuca, and morelos. ences could be found among them, but when there were few differences, apparent EXPLANATION OF MORPHOLOGICAL intergradation, or little material, the de- AND BEHAVIORAL TERMS cision as to whether one or more species Markings in General. Color patterns are are present was sometimes difficult. In sev- generated by integument coloring and by eral cases, only a single species was rec- covering of setae, though pale setae usually variation, ognized despite geographical overlie and dark overlie because the variation was too pale integument geographical dark. The terms hair and scale are used to at or confusing present (furcata-mimus) , refer to a thin, more or less cylindrical seta because the differences were slight and and a broad, flattened seta, respectively possibly not consistent {aeneola / uteanus, (cf. Hill, 1979). northern and Floridian tillandsiae, north- Markings of the Carapace. Male den- ern and southern carmenensis, mannii/ dryphantines are commonly dark brown versicolor) . In other cases, allopatric pop- with bands of white to yellow scales, usu- ulations that are similar but that differ con- ally including a major longitudinal band sistently in a number of features were rec- on either side of the carapace and abdo- ognized as distinct {sabinema / pervaga, bi- men. The following names are used to re- cuspidata /volcana, neoleonis /tristis, fer to the bands of pale scales on the car- proxima /galathea / dithalea, chera / tricol- 1): in apace (Fig. or / diplacis) , though each of these cases the decision was difficult. Pelegrina fla- side bands: Longitudinal bands on either Pelecrl\a Jumping Spiders • Maddison 223

side of carapace, beginning beside the paired: the first pair just anterior to the anterior lateral eyes (ALEs) and pro- muscle attachment of the second dorso- ceeding just beneath the small eyes and ventral muscle (number 86viii + ix of posterior eyes and beyond, onto the tho- Whitehead and Rempel, 1959), the second rax. pair anterior to the attachments of the third cheek band: Oblique band on the side of dorsoventral muscle, and the third, fourth, the face, starting beneath the ALEs and fifth, and sixth pairs of spots behind this. proceeding down and posteriorly to the The fifth and sixth pairs are very small. carapace margin, in Pelegrina and the Often the fourth through sixth pairs are mannii group. each connected medially and thus form forehead band: A V-shaped marking on small chevrons. Between these pale spots the dorsal cephalic area just behind the may be spots of dark brown, sometimes anterior median eyes (AMEs). very dark (e.g.. Fig. 353). marginal band: On the lower margin of Male Palpus (Figs. 3, 6-9). The descrip- the sides of the carapace, often extended tions generally assume that the left palp is from the cheek band in Pelegrina. Fe- being viewed from the ventral. The ad- males of most species show none of these jectives basal and apical when unqualified bands distinctly; the carapace is instead refer to the appearance in a contracted = often covered more or less uniformly palp (i.e., basal toward the tibia and = with pale scales including a dense white apical toward the tip of the cymbium). covering on the clypeus. In contrast, "anatomically basal" and "api- cal" refer to the cymbium-embolus axis Setae Surrounding Anterior Median of connections of the palp's bulb (ie., an- = Eyes. These are of various colors, from atomically basal toward the basal he- = white to black. In the descriptions of Pe- matodocha and anatomically distal to- legrina males, the colors of setae around ward the tip of the embolus). the circumference of the left anterior me- In the subfamily Dendryphantinae, the dian eye (AME) are indicated using a no- shoe-shaped cymbium holds a bulb con- tation derived from hours on a clock's face. sisting of (from anatomically basal to api- Usually, the colors on only the dorsal part cal) a fully expandable basal hematadocha, of the AMEs are described, for those ven- a small subtegulum, a much reduced me- trally are more variable. Thus, "white dian hematodocha, the large tegulum, a forehead band contacts the AME dorsally fully expandable distal (embolic) hema- 10:30-12:30" means that as one looks from todocha, and the embolus. The basal he- the front at the left AME there are white matodocha expands so as to give a clock- setae from 10:30 o'clock to 12:30 o'clock wise rotation to the subtegulum and te- that are continuous with the forehead band, gulum in ventral view of the left palp, as and dark setae on either side of this. in other salticids. This has been observed Markings of the Legs. The legs are yel- by artificial expansion on many dendry- low to dark brown, but there are often phantines and during copulation of Den- subte- annulate markings (Fig. 1), especially in dryphantes nigromaculatus. The males, so that each leg has pale portions gulum is small and contains little more covered with white scales alternating with than the basal portion of the sperm duct darker portions lacking white. reservoir (Figs. 3, 8). Markings of the Abdomen. The abdo- From the subtegulum, the sperm duct men is usually brown above in males, loops up, down, and around the tegulum in a clockwise direction to ringed by white side bands (Fig. 1). The (Fig. 3) the em- dorsum of the abdomen often shows traces bolus. The distal retrolateral portion of the of the paired pale spots seen in females tegulum, where the sperm duct enters the and from the is (Fig. 2). These pale spots are central tegulum subtegulum, generally 224 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

extended distally and contains a loop of bium), as indicated by artificial expansions the sperm duct pressed against its wall. (Figs. 7, 9). The counterclockwise move- This portion I call the shoulder of the te- ment of the embolus itself probably ex- gulum (Fig. 3). Just proximal to the em- plains why the counterclockwise-coiled bolus and shoulder is a fold extending across embolus can still engage the epigynal the surface of the tegulum, the tegular opening despite the clockwise movement /^(fge (Figs. 3,6, 7), which serves as a pock- of the tegulum (in salticids with the em- et to hold the proximal part of the embolic bolus fixed to the tegulum, the embolus base. Wanless (1984) suggested that the has a clockwise curve, which thus coin- thrust of te- tegular ledge (his M3) is absent from sal- cides with the clockwise the ticids other than spartaeines, in which he gulum). The embolus of dendryphantines described it. However, the fold that I am usually consists of a basal portion, which here calling the tegular ledge may be ho- is transversely directed, and an apical por- mologous with his M3, for it appears as an tion, which is usually thin and erect and extension of the embolic hematodocha cut- has the opening of the sperm duct at its ting across the face of the tegulum and tip (Figs. 40-47; also Figs. 64b, d). The occurs in many groups of salticids. The embolic base consists of a more or less tegulum is hWed with tegular glands, which sclerotized portion of the embolic hema- empty into the sperm duct via a series of todocha that is exposed to the ventral sur- pores in the sperm duct (see Osterloh, 1922; face and that rests betwee the tegular ledge Bhatnagar and Rempel, 1962). As in other and the embolus (Figs. 3, 6, 7). Its wrinkles spiders, these pores are aligned into a band suggest that it should be considered part (Schult, 1980), which, when narrow, ap- of the hematodocha. As noted, the embolus pears as if it were a seam along the length is coiled counterclockwise, a feature much of the sperm duct. Along the narrower part more readily apparent in the euophryines of the sperm duct toward the embolus, the and other subfamilies than it is in the den- glands are connected to the sperm duct via dryphantines. In dendryphantines, the spi- long ducts (Fig. 3; also known from other ral is hidden and best seen in expansions spiders; Osterloh, 1922: figs. 20, 26). These or dissections (e.g.. Fig. 35). The dendry- pores, ducts, and glands have been largely phantine embolus arises prolaterally and ignored in systematics but appear to be a moves across toward the retrolateral side potential source of good systematic char- (the transverse basal portion of the em- acters (for instance, in some salticids the bolus) and then folds back toward the pro- pores are arranged in a broad ribbon, lateral and abruptly rises as the erect apical whereas in others the band is very narrow, portion (Figs. 20-23). In many genera, the and in Sitticus the pores are arranged into erect apical portion is fused against the prominent craters). transverse portion so that there is no open. The embolic hematodocha arises on the freely coiling spiral. A suture on the back back side of the tegulum (Fig. 8), prolater- side of the embolus (the embolic suture), al to the subtegulum. Its wrinkles sweep between the transverse portion and the apically up toward the embolus. The exact erect portion, is often present and indicates arrangement of the folds of the hemato- where the folded-back spiral has not com- docha is probably of systematic value but pletely fused (Figs. 3, 8, 20-23, 31-35). In is very difficult to untangle, for especially cleared palpi, a slight bend in the sperm near the base of the embolus the folds are duct can be seen at this point. The coiling twisted and confusing even in a well- of the embolus and the embolic suture sug- cleared palpus. During expansion, the em- gesting it are clearly visible in Dendrij- bolic hematodocha expands fully to move phantes, Eris, Pelegrina, Phidippus, Tu- other the embolus counterclockwise (i.e., pro- telina, Phanias, and many genera, the is eas- laterally) and back (i.e., toward the cym- In Metaphidippuschera, coihng Pelegrina Jumping Spiders • Maddison 225

ily seen (Fig. 35); in Pelegriria proterva, among the species of Pelegrina (Figs. 236- the only trace is the small suture (Fig. 34); 255). In some, the entire surface behind in Pelegrina flavipedes, the suture is some- the openings is raised into a mound (e.g., times visible and sometimes not. No trace P. clemata. Fig. 246); in others, it is much or of the suture past coiling can be found more concave (e.g., P. furcata. Figs. 249, in Terralouits cf. uiiicus and Poultonella 250). At the posterior margin of the epi- alboimmaculata, but Terralonus myloth- gynum is the notch, or guide (Figs. 4, 5), rus and Tutelina elegans, which are, re- into which fits the male tibial apophysis, spectively, their close relatives, show them although in some dendryphantine groups well. The loss of a trace of coiling may {Poultonella, ) the guide has have evolved several times in the dendry- moved anteriorly as in pellenines (Pel- phantines. In some genera such as Zygo- lenes, Hahronattus) and Bianor. ballus, Hentzia and Mabellina, the coil is The different parts of the copulatory not so compact and, instead, is more open ducts of Pelegrina species are named as (e.g.. Figs. 24-27, 37, 38, 50, 51, 58-63, follows (Fig. 5). From the anterior half of 64c, f). Figure 64 summarizes some of the the openings, the ducts proceed first lat- coiling patterns seen in dendryphantines. erally and posteriorly (the first curve), then The erect portion of the embolus is some- medially (the second curve), and then pos- times a simple spike with the opening ter- teriorly (the third curve), and then twist minal, but in many dendryphantines there a number of times and proceed dorsally to are prolongations that will be referred to the fertilization ducts. The inner surface as rami, and often small denticles occur of the duct is relatively smooth in the first on the surface of the embolus. In Pelegri- and second curves, smooth or rough in the na, in particular, there are two rami re- third curve depending on the species, and trolateral to the opening of the sperm duct, rough with projections in the twisted area the prolateral ramus very near the opening posteriorly. The flower-shaped openings and the retrolateral ramus some distance of the accessory glands occur on the second away (Fig. 3). curve, usually near the junction with the Epigynum. In most dendryphantines, first curve (Fig. 5). The pathway from the the openings are well separated and copulatory opening through the copula- S-shaped (Figs. 65-70), with entry toward tory ducts toward the fetilization ducts is the lateral in the anterior half and toward almost straight, lacking the separate sper- the medial in the posterior half (Fig. 5). mathecal reservoir seen in, for instance, The lateral rim of the opening is often some euophryines. thickened to yield a more or less convex Courtship. Some introduction to the for- teardrop-shaped area, which will be called mat of descriptions of courtship has al- the epigynal flap (Fig. 4). The inner mar- ready been given under the Materials and gins of the left and right flaps may be Methods section. Males, especially early in parallel to each other (e.g.. Figs. 262, 274, display, often walk not straight toward the 322) or be divergent from anterior to pos- female but instead sidle so that they ap- or not at all. terior (e.g.. Figs. 346, 352, 398) or be con- proach obliquely The sidling be first to the left and then to the vergent (e.g., Figs. 280, 297, 332, 363). In may some Pelegrina, the flaps converge to such right, and so on, to form a zigzag dance an extent that their posterior halves are (Jackson, 1978). The male usually walks rotated 90° and become transverse (e.g., P. sporadically, taking a series of steps then kastoni, Fig. 317). Even more extreme ro- pausing, another series of steps then paus- tations are seen in P. arizonensis (180°, ing, and so on. Because the male's pose Fig. 424) and P. helenae (270°, Fig. 430). and motions of the first legs, palp, and The surface of the epigynum between and abdomen often differ depending on behind the openings varies in topography whether he is walking or paused, a dis- 226 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

tinction is made between the walking Plexippus, Hyllus, Evarcha, Thyene, Te- of Bia- phase, which is called the series (of steps), lamonia, Harmochirus, and part and the pauses. While the male is walking, nor, delimited by a modified serrula on the male endite and a on the te- the first legs are generally raised, spread, bump and/or extended forward. The first legs gulum just counterclockwise [left palp] of other familiar may be waved or flickered up and down the embolus), and many or forward and backward, one or repeat- groups such as Pellenes, Salticus, Sitticus, their edly. The distinction between a wave and Phiale, Myrmarachne, Amyous, and a flicker is not precise; in general, "wave" respective relatives. However, the Eu- is used when the motion is of low speed ophryinae, Dendryphantinae, and several or frequency, "flicker" for high-frequency smaller groups are united by an embolus repeated motions. The abdomen may be that is free to move, being separated from twitched (Jackson, 1978) down then up, the tegulum by a fully expandable he- which in many species produces a sound matodocha (Figs. 29-31). In these groups, that may function in courtship (Maddison the embolus is also coiled counterclockwise and Stratton, 1988). Explanations of (left palp). The subfamily Dendryphantin- "crouch" and "raisedspread" stages of ae itself is delimited by the derived con- courtship are given in the discussion of ditions of a carina on the underside of the monophyly (item 7) in the description of male chelicera, by the coil of the embolus the genus Pelegrina. folded back so as to be hidden behind the base of the embolus, and by S-shaped epi- THE SUBFAMILY DENDRYPHANTINAE gy^al openings. Pelegrina and Metaphidippus are den- The Dendryphantinae is a subfamily of dryphantines, which are salticids. Maddi- several hundred species, most of these in son (1988) reviewed the phylogeny of the the New World. Males are usually striped, family Salticidae. The majority of salticid with longitudinal bands of pale scales on species are considered to form a mono- either side of the carapace and abdomen, phyletic group, called the Salticine divi- while females usually have paired spots on sion (Maddison, 1988), which are distin- the abdomen. The chelicerae of males are guished from the remaining groups (Lys- often enlarged or elongate as is the first somaninae, Spartaeinae, and the Cocal- pair of legs. The posterior eyes are smaller odes group) by eye structure (Wanless, than in most euophryines. 1984; Eakin and Brandenburger, 1971; Table 1 lists the generic names (includ- Blest, 1983; Blest and Sigmund, 1984), ab- ing those now considered synonyms) that sence of a tarsal claw on the female palp, I refer at least tentatively to the subfamily, medial displacement of the gnathocoxal Some of these genera are included for the with glands (Figs. 14, 15), asymmetrical tarsal first time. Some genera are included claws, a mound of slit sense organs with hesitation, either because they are obscure an associated seta on the medial edge of and their type species were not examined or the chelicera (Figs. 12, 13), and a small (e.g., Anamosa, Homalattus) because intercheliceral sclerite (Figs. 12, 13), each they exhibited none of the supposed den- of which may be considered derived with- dryphantine synapomorphies listed later in the family. Within the Salticine divi- but have the general body form and mark- sion, there are some prominent subfamilies ings much like those of other dendry- Other that have the embolus fixed immovably to phantines (e.g., Mabellina). — genera the tegulum, including the Heliophaninae are excluded with hesitation for in- (delimited by an apparent stridulatory ap- stance, those genera related to Ballus (Co- paratus and a bump on the tegulum just taxes, , , Pachyballus, and clockwise [left palp] of the embolus; Mad- perhaps Admestina) and to Synageles dison, 1987), the Plexippinae (including (Consingis, Descanso, Peckhamia, and Pelegrina Jumping Spiders • Maddison 227

Table 1. Generic names of jumpinc; spiders tentatively referred to the subfamily Dendryphantinae. Synonymies are not indurated.

Admirala G. & E Peckham, 1901 Nagaina G. & E. Peckham, 1896 Agassa Simon, 1901 Osericta Simon, 1901 Amerotritte Mello-Leitao, 1944 Paradamoetas G. & E. Peckham, 1885 Anamosa G. & E. Peckham, 1895 Parahentzia Bryant Chamberlin, 1925 F. P.-Cambridge, 1901 Anoka G. & E. Peckham, 1893 Parnaenus G. & E. Peckham, 1896 Ashtalnda G. & E. Peckham, 1894 Partona Simon, 1902 Avitus G. & E, Peckham, 1896 Pelegrina Franganillo, 1930 Bagheera G. & E. Peckham, 1896 Phanias F. P.-Cambridge, 1901 Bcata G. & E. Peckham, 1895 Phidippus C. L. Koch, 1846 Bellota G. & E. Peckham, 1892 Poultonella G. & E. Peckham, 1909 Brijantella Chickering, 1946 Raniboia Mello-Leitao, 1944 Cerionesta Simon, 1901 (n. nov. for Cydonia G & Rhene Thorell, 1869 (n. nov for Rhanis C. L. Koch, E Peckham, 1893, preoccupied) 1848, preocc.) Chirothecia Taczanowski, 1878 Rhetenor Simon, 1902 Dendryphantes C. L. Koch, 1837 Rndra G. & E, Peckham, 1885 Donaldius Chickering, 1946 Sassacus G. & E. Peckham, 1895 Dryphias Simon, 1901 Sebastira Simon, 1901 Eris C. L. Koch, 1846 Selimus G. & E. Peckham, 1901 Gastromicans Mello-Leitao, 1917 Semora G. & E. Peckham, 1892 Ghelna new genus Tacuna G. & E. Peckham, 1901 Hentzia Marx, 1883 Terralonus new genus Homalattoides F. P. -Cambridge, 1901 Thammaca Simon, 1902 Homalattus White, 1841 Tulpius G. & E. Peckham, 1896 Lurio Simon, 1901 Tutelina Simon, 1901 Mabellina Chickering, 1946 Uluella Chickering, 1946 Maeviobeata di Caporiacco, 1947 Wala Keyserling, 1884 Megatimus Thorell, 1891 Zeuxippus Thorell, 1891 Messita G. & E. Peckham, 1896 G. & E. Peckham, 1885 Metaphidippus F. P. -Cambridge, 1901

perhaps Cheliferoides and Leptorchestes) Metaphidippus, Paradamoetas, Phidip- may very well be derived dendryphan- pus, Rhetenor, Sassacus, Thiodina, and tines (see Maddison, 1988). Wala, in the subfamily Dendryphantinae; Simon (1901, 1903) placed many of these all except Cheliferoides and Thiodina are dendryphantine genera in his group Den- here considered dendryphantines. dryphanteae or Rheneae, though a num- Specifying distinct apomorphies to jus- ber of them were scattered among other tify my concept of the subfamily is diffi- groups: Beata in the Simaetheae, Bellota cult, for no characters both universal in the Synageleae, Nagaina in the Belli- throughout and unique to the group are eneae, Rudra in the Rudreae, Tutelina in known; homoplasy must therefore be in- the Chrysilleae, and Zygoballus in the Zyg- voked if the subfamily is to be accepted oballeae. G. & E. Peckham (1901b) in- as proposed. Three characters derived cluded in their Phidippus group of genera within the family are proposed as syna- Phidippus, Paraenus, Dendryphantes, Se- pomorphies of the subfamily: limus, and Admirala, here considered dendryphantines, but as well many other 1. Carina on underside of the male che- licerae genera now considered to belong to the (Table 2, character 1). On the Euophryinae and other subfamilies. Pro- ventrolateral edge of the basal segment szynski (1976: 15, 148-150) listed the gen- of the chelicera there is a fold or carina era Cheliferoides, Dendryphantes, Eris, (Fig. 10). This carina occurs in almost 228 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

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all genera considered dendryphantines. ation seen among the dendryphan- It is absent in some groups in which the tines). Despite the first problem, coiling chelicerae are much elongate (Messua, is considered primitive for the subfam- Hentzia, Riidra, Paradamoetas), ily (see earlier discussion under the sec- though this may be related to the elon- tion Explanation of Morphological and gation, as Hentzia mitrata (with short Behavioral Terms). More troublesome chelicerae) does have the carina. More is the second problem: the occurrence troublesome are those dendryphantine of dendryphantine groups such as Par- groups (some Phanias, Tutelina, ^'Eris' adamoetas, the South American "Sas- nidicolens, "Beata" octopunctata) in sacus," Mahellina, Dendryphantes tro- which the chelicerae are not elongate picus, and Bryantella in which the coils and yet the carina is lacking. In at least of the embolus are exposed (Figs. 26, one of these, Phanias, there are some 27, 58-63, 64e, f). However, three lines species with the carina, and its absence of evidence suggest that the exposure in others is presumably a secondary loss. of the coils is not homologous to that of The carina is lacking in all other salti- the euophryines and, instead, is sec- cids I have seen, except for Simaetha ondarily derived from the hidden con- paetula (see Prosynski, 1984: 132) and dition. First, the exposed spiral of Den- Simaetha tenuior, in which a similar dryphantines is always of a different carina is present. These species have a form from that of the euophryines, be- Sitticus-\ike palpus, with a round te- ing placed more retrolaterally and not gulum and immovable embolus, and so small and tightly coiled. Second, oth- appear to be distantly related to den- er characters suggest that these trou- dryphantines. blesome dendryphantines do indeed 2. Coil of embolus compressed, so that the belong with other dendryphantines, embolus folds back sharply on itself and such as the lack of a concave retrolateral superficially appears not to be coiled loop on the sperm duct in the tegulum, as lat- (Figs. 20-25, 31-35, 37-39, 64b-d, g; which may be apomorphic noted Table 2, character 2). Typically, the er with respect to Phanias, the more embolus arises prolaterally and moves basally placed tegular ledge, which may across toward the retrolateral side (the be apomorphic or plesiomorphic, and transverse basal portion of the embolus) the occurrence of the cheliceral carina and then folds back toward the prolat- in at least some species (e.g., Metaphi- eral and abruptly rises as the erect api- dippus vitis). Third, there is an appar- cal portion. A suture on the back side ent morphocline between the hidden of the embolus, between the transverse spiral of Eris aurantia (Fig. 64b), portion and the erect portion, is often through the more marginally open spi- present and indicates where the folded- ral of Eris militaris and a species near back spiral has not completely fused. Zygoballus incertus (Figs. 64d, 53, 57), The coils of the embolus are not folded to the more open spiral of Zygoballus but rather open and exposed (Fig. 19), incertus and Paradamoetas (Figs. 64e, in the other groups, such as the eu- 58), to the open and well-coiled spiral ophryines, that have a counterclock- of Mahellina and Dendryphantes tro- wise-coiled embolus. Two main prob- picus (Figs. 62, 63, 64f). The transition lems with the use of this character are would require merely a retrolateral shift posed by various dendryphantines: in of the erect portion of the embolus and some, the embolus appears not even a loss of sclerotization of the basal (lat- coiled, and in others the embolus ap- erally directed) portion of the embolus pears coiled but the coils are exposed to leave the erect part of the embolus (Fig. 64 summarizes some of the vari- free, at which point it could coil as in Pelecrina Jumping Spiders • Maddison 231

Dendryphantes tropicus and Mabelli- focus on those phylogenetic questions that na prescotti (Figs. 62, 63). are most important to resolve the generic 3. with en- Epigynal openings S-shaped, placement of the galathea group of spe- the lateral in the try toward anterior cies, which is the subject of the species half and toward the medial in the pos- revision that makes up the bulk of this terior half (Figs. 5, 65, 67-70; Table 2, paper. The galathea group has resided in character 3). To my knowledge, this the genus Metaphidippus, but, as ex- sinuate opening is unique among sal- plained next, this genus is polyphyletic. ticids. Though most dendryphantines Here I will begin (but not complete) the have this character, it has similar prob- task of dismantling Metaphidippus. The lems to the preceding one and is cor- galathea group will be moved to the genus related with it. A number of dendry- Pelegrina, the harfordii group to Phanias, phantines (e.g., Eris militaris, Tutelina the mylothrus group to Terralonus, the elegans, Phidippiis clarus, Hentzia, castaneus group to Ghelna, some other Paradamoetas, Anicius, Zygoballus, species to Messua, but some species groups Messua, Bagheera) have C-shaped or (mannii group, vitis group, various neo- simple cavernous openings. In some of tropical species) will remain temporarily these, however, ridges descending into in Metaphidippus for want of a better al- the opening are presumably remnants ternative. Table 3 lists the proposed re- of the teardrop-shaped flaps associated classification of Metaphidippus and some with the sinuate openings (Eris mili- other dendryphantine species. taris, Fig. 66; Tutelina elegans). For Metaphidippus was described in 1901 some of these, there are related species by F. O. Pickard-Cambridge, who gave no that have the sinuate openings {Eris clear justification for its limits. Many of floridana, Tutelina hartii, Phidippus the North American species that had been spp., Anicius sp., Zygoballus tibialis), placed in Dendryphantes were subse- but there remain other genera with no quently transferred to Metaphidippus, in remnant of the flaps or obvious close part because of Bryant's (1941) conclusion relatives with sinuate openings. In gen- that these species did not belong with the eral, the species lacking a compact em- Old World Dendryphantes. In fact, her bolic spiral also lack the sinuate open- cited evidence was mistaken: the Euro- were ac- ings, perhaps because a retrolateral shift pean specimens she compared of the erect portion of the embolus is tually "Eris" nidicolens misidentified as correlated with an expansion of the epi- D. hastatus (Maddison, 1988). Regardless, of North gynal openings and weakening of the the more or less wholesale transfer re- teardrop-shaped flaps. Such a correlat- American species from Dendryphantes ed change seems to mark each of Eris sulted in a Metaphidippus that has been militaris, Pelegrina kastoni, Dendry- desperately polyphyletic, being nothing phantes nigromaculatus, and Phidip- more than a catch-all genus of unremark- pus octopunctatus from its close rela- able North and Central American dendry- tives. phantines spanning much of the diversity of the subfamily. Because the type species RELATIONSHIPS WITHIN THE of Metaphidippus (M. mandibulatus F. O. DENDRYPHANTINAE Pickard-Cambridge) is not closely related in the in- The limits and interrelationships of gen- to most species placed genus, the era within the subfamily Dendryphantin- cluding galathea group, many species at of should be placed else- ae (see Table 1 for a list) are present Metaphidippus discus- where. The relationships of the true Me- poorly understood. The following will first be considered, after sion will make an attempt to resolve only taphidippus for I will which the hmits of Dendryphantes and a small part of the confusion, 232 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Table 3. Summary of placements of various New World dendryphantine species apart from Pelegrina species, discussed in text. ? indicates placement considered possible but unconfirmed. Authors of names are given in index.

Bagheera Phanias Phanias albeolus B. prosper P. concoloratus ?Metaphidippus nigropictus P. dominatus ?M. bicavatus P. flavostriatus P. furcifer Beata P. furcillatus Beata hispida P. harfordii B. inconcinna P. monticola B. longipes P. neomexicanus B. maccunii P. watonus B. magna ?P. salvadorensts B. rustica Terralonus Dendryphantes (partial) Terralonus californicus Dendryphantes nigromaculatus T. niylothrus D. chuldensis T. unicus D. fusconotatus T. shaferi D. hastatus T. versicolor D. rudis T. vittatus Gastromicans T. fraternus

Gastromicans alhopilosa Ghelna G. hondurensis Ghelna castanea G. levispina G. barrowsi G. noxiosa G. canadensis C. vigens G. sexmaculata ?Hasarius lisei Metaphidippus niannii group Messua Metaphidippus mannii Messua desidiosa M. carmenensis M. centralis M. chera M. dentiger M. diplacis M. donalda M. emmilttis M. lata M. tricolor M. laxa M. limbata Metaphidippus vitis group M. niunia Metaphidippus vitis M. octonotata M. texanus M. pura M. mathetes M. tridentata Dendryphantes melanomerus ?Metaphidippus cupreus ?Metaphidippus ovatus ?Metaphidippus iridescens ?Metaphidippus inflatus ?Metaphidippus quadrinotatiis ^Dendryphantes felix

Beata will be discussed. Then, the correct taphidippus appear to fall. Members of placement of various groups placed in Me- this group, which includes such common taphidippus will be considered. species known as "£m" limbata, ''Meta- phidippus" prosper, "Beata" alhopilosa, The of Genera Bagheera Group have a distinctive embolus, which appears, It is within the Bagheera group, com- at first glance, to be coiled or curved clock- mon in the Neotropics, that the true Me- wise in the left palpus, opposite the coun- Pelegrina Jumping Spiders • Maddison 233

terclockwise coiling I have said character- have a twisted embolus, but in each it takes the izes dendryphantines and related sub- a very different form from the twisting in families (Figs. 99-101). This clockwise the Bagheera group of genera. Sebastira, coiling is apparently superimposed upon Thammaca, Lurio, and Parnaenus may the normal coiling by a twisting of the also belong with the Bagheera group of embolus. The main axis of the embolus has genera, though their emboli do not so ob- twisted counterclockwise (as viewed from viously possess the twisting. A brief ac- tip of embolus in left palpus), thus winding count of Bagheera, Messua, Gastromicans the hematodocha and sperm duct around and Metaphidippus is here given. it (Figs. 64h, 99-101). The tip of the em- Bagheera (Figs. 80-85). Bagheera males bolus, though, seems to have been left be- have elongate, horizontal, parallel chelic- hind by the twisting, so that the apical part erae (Fig. 80); the retromarginal teeth are of the embolus takes on a clockwise curl- near the base of the chelicera; in all but ing. In some species, this clockwise coiling one species there is distally, near the fang, is visible in the uncleared palpus (Figs. 81, what appears to be a large retromarginal 84, 87, 93), but in others it is not (Fig. 91), tooth but actually is not (it does not have and the palpus is little modified from the the terminal canal through the cuticle that typical compressed counterclockwise spi- seems to characterize all true teeth), and ral of dendryphantines. The most extreme most species have tubercles bearing setae clockwise coiling is seen in ^'Metaphidip- on the inner margin of the basal cheliceral pus" prosper (Fig. 99) and ^^Beata" al- segment. Included in Bagheera are the type bopilosa (see Fig. 101). species, B. hiplingi G. & E. Peckham, 1901 Species in the group can be sorted pro- (type species by monotypy; holotype ex- visionally into three subgroups, which may amined; Figs. 80-83), and Bagheera pros- or may not be monophyletic, for each of per (G. & E. Peckham) (NEW COMBI- which there exists an available generic NATION; Figs. 84, 85, 99) and at least two name that has been mostly ignored in the undescribed species. Metaphidippus ni- literature: Bagheera G. & E. Peckham, gropictus F. P. -Cambridge and M. bica- Messua G. & E. Peckham, and Gastromi- vatus F. P. -Cambridge may also belong in cans Mello-Leitao. A fourth genus sharing Bagheera. a twisted embolus is Ashtabula G. & E. Messua (Figs. 86-92). Males of this ge- Peckham, though whether or not it belongs nus have elongate divergent chelicerae with the group is unclear. The twisting of (Fig. 86) with a long and sickle-shaped the embolus in Ashtabula (Fig. 102) is hid- retromarginal tooth near the fang; the pro- den beneath the tegulum and is more ex- marginal teeth are near the base, well sep- on treme, though similar to, that in Bagheera, arated from the retromarginal tooth; Messua, and Gastromicans. However, the anterior distal margin of the basal seg- there are several features that cast doubt ment of chelicera near the fang is a flange. on the placement of Ashtabula with these Included in Messua are the type species 1896 genera. Ashtabula has an extra concave M. desidiosa G. & E. Peckham, (type and collec- loop on the sperm duct in the palpus, pos- species by monotypy; holotype of males and females from San sibly placing it near the base of the sub- tions Jose, Mes- family as noted later in connection with Costa Rica examined; Figs. 86-89), Phanias. The body in Ashtabula is not sua centralis (G. & E. Peckham) (lectotype a male from nearly so large and robust as in the Bag- here designated, Chiriqui), P. heera group, instead resembling that of Messua dentiger (F. -Cambridge) (see Hentzia or Anicius. More work is needed Fig. 91), Messua donalda (Kraus), Messua Messua laxa before the place of Ashtabula can be set- lata (Chickering), (Chicker- limbata tled. Species of Ghelna and the arizonensis ing), Messua (Banks) (Figs. 90, P. -Cam- group of Pelegrina (discussed later) both 100, 117), Messua moma (F. No. 4 234 Bulletin Museum of Comparative Zoology, Vol. 154,

P. -Cam- bridge), Messua octonotata (F. bridge (type species by original designa- and Mes- whose known male Rica, bridge), Messua pura (Bryant), tion), single (Costa is unlike sua tridentata (F. P.-Cambridge). All these BMNH, examined), strikingly other that have been except M. desidiosa are NEW COMBI- most jumping spiders NATIONS. Metaphidippus cupreus F. P.- placed in Metaphidippus. Palpus (Figs. Embolus reminiscent of that of Cambridge, M. ovatus F. P.-Cambridge, 97, 98): M. iridescens F. P.-Cambridge, M. infla- Eris species but with the longitudinally tus F. P.-Cambridge, and M. quadrino- directed apical portion not fully erect, in- tatus F. P.-Cambridge may also belong in stead reclined to the prolateral (Fig. 98). Messua. Dendryphantes felix G. & E. The embolic base bears a flange covering Peckham might be considered either a the basal part of the embolus. Chelicerae: horizontal and di- Bagheera or Messua depending on any Long and cylindrical, future lectotype designation: the body (G. verging (Fig. 96), with two promarginal & E. Peckham, 1901b: fig. 6a) in the type teeth near the base and one retromarginal vial and its attached palpus are of a Bag- tooth near the fang. The fang is forked heera species, probably B. prosper, while near its base (Fig. 96). Markings (Fig. 96): side bands ex- the separate palpus in a microvial (G. & Carapace brown, lacking scales on either E. Peckham, 1901b: fig. 6) is of a Messua cept one patch of white white band mar- species. side of fovea. Wide along scales Gastromicans (Figs. 93-95). This genus gin. At least some metallic green-blue with thin white is distinguished from Bagheera and Mes- on cephalic area. Abdomen sua in having short and vertical but very side bands broken basally; just anterior to robust male chelicerae. Included are Gas- each of the main dorsoventral muscle at- tromicans albopilosa (G. & E. Peckham), tachments is a small white patch of scales; Gastromicans hondurensis (G. & E. Peck- in the posterior half of the dorsum are two ham), Gastromicans levispina (F. P.-Cam- pairs of small lateral white bars. The dor- scales. bridge) (Figs. 93-95, 101), Gastromicans sum has some metallic green-blue noxiosa (Simon), and Gastromicans vigens Measurements: Body length 5.4 mm; car- (G. & E. Peckham). All these are NEW apace length 2.4 mm, carapace width/ COMBINATIONS. Hasarius lisei Bauab- length 1.93/2.37. Vianna & Soares probably also belongs in Two features that can be proposed as Gastromicans. Galiano (1980) synony- synapomorphies for the group of Bag- mized Gastromicans Mello-Leitao with heera, Messua, Gastromicans, and Me- Beata G. & E. Peckham because its type taphidippus are the following: species Gastromicans squainulata Mello- Leitao (type species by monotypy) is syn- 1. The tibial apophysis is erect and at its knife onymous with "Beata" albopilosa. But in- base parallel-sided, shaped like a sofar as Beata albopilosa does not belong blade (Figs. 71-74; Table 2, character in the genus Beata, Gastromicans is avail- 4). Almost all other dendryphantines able as a generic name for albopilosa and have an apophysis tapering throughout its relatives. its length (Figs. 75-79), including that Metaphidippus (Figs. 96-98). Though Phanias and other genera appear the placement of the true Metaphidippus to be near the base of the subfamily with these genera of the Bagheera group (see later), and usually the apophysis is to some extent problematical, such a points at least somewhat ventrally. The to placement is the best supported at present. only other dendryphantines known Before discussing the uniting characters, it me with a similar knife-shaped apoph- Poultonella. would be valuable to give the following ysis are Ashtabula and brief description of the type species of Me- Poultonella does not belong with these taphidippus, M. mandibulatus F. P.-Cam- genera; rather, its peculiar chelicerae Pelecrina Jumping Spiders • Maddison 235

assure a relationship with Tutelina. A reasonable given that other species such as few species of Messua, inchiding M. Messua octonotata have little trace of a lata, have a more tapering apophysis. twisted embolus, and the embolus of M. 2. The tegular ledge runs longitudinally mandibulatus shows unusual folds and does (Figs. 87, 90-93, 98; Table 2, character recline to the prolateral as in Messua. Per- 5), instead of obliquely at 0-60° from haps more detailed study of its peculiar the transverse as seen in other dendry- embolus, when more specimens become phantines and other salticids with a teg- available, will allow a more definitive an- ular ledge (Figs. 40-46, 50, 52, 53). swer to the question of twisting in M. man- While this is unusual among dendry- dibulatus. If the genus Metaphidippus is phantines, it is not unique: it also occurs only an offshoot of Messua, then Meta- in Phanias, Anicius, and Zygoballus in- phidippus would fall as a synonym of the certus. older name Messua. However, I am re- luctant to effect such a synonymy at pres- Additional features that suggest a rela- ent given the number of Metaphidippus tionship between Metaphidippus and species that would be left homeless, and Messua in particular are the long, tubular so Metaphidippus will be left standing for divergent chelicerae with a near-terminal the moment. Regardless, the best-support- retromarginal tooth and a distal anterior ed conclusion at present is that the name flange beside the fang base. Other den- ''Metaphidippus" properly applies to a dryphantines have long divergent chelic- small group of neotropical dendryphan- erae, but in all that I have seen except some tines related to Messua, Bagheera, and South American "Sassacus," the tooth ar- Gastromicans. rangement is different than in Metaphi- Two Genera That Have Exchanged dippus and Messua, with the retromar- Species with Metaphidippus: ginal tooth remaining near the base or the Dendryphantes and Beata promarginal teeth near the apex. This dif- ferent tooth placement may indicate that Because most species that have been the elongation occurred in different por- placed in Metaphidippus do not belong to tions of the chelicerae in these other den- the Bagheera group of genera, they cannot dryphantines, and thus the elongation is follow M. mandibulatus and, thus, need not homologous. As well, the general body to be placed elsewhere. The first place we form and occurrence of greenish reflective might look for a possible home for Me- scales are also suggestive of a close rela- taphidippus species are two genera, Den- tionship between Metaphidippus and dryphantes and Beata, which have in the Messua, though these characters are loose- past exchanged many species with Meta- were ly defined and not necessarily unique. The phidippus. Many "Metaphidippus" in and only feature that would exclude M. man- formerly placed Dendryphantes in dibulatus from Messua is the apparent lack several Beata were formerly placed Me- of the reverse twisted embolus in M. man- taphidippus. To discuss the proper place- dihulatus. However, one undescribed spe- ment of species now in Metaphidippus cies from Costa Rica represented by a sin- more clearly, it would be valuable to re- the limits of these two gle male specimen appears very closely consider genera. related to M. mandibulatus in having sim- Dendryphantes (Figs. 65, 103-108, 120). ilar body form and markings and in having The genus Dendryphantes, described last it has a has over the accumulated a slightly forked fang, and yet century, years on the basis of their slightly twisted embolus (Fig. 92). If these many species, mostly unremarkable two species form a monophyletic group, being dendryphantines. were since moved to then the lack of twisting in mandibulatus Many species genera is not un- such as while others re- may be a secondary loss, which Metaphidippus, 236 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

maining in the genus will probably even- The option of returning a number of tually be placed elsewhere (see the com- groups from Metaphidippus to Dendry- ments of Edwards, 1977). Among the New phantes has little merit at present. Moving World species, there is only one species for these back to Dendryphantes would be which there is presently good evidence for useful only if they are likely to stay there, a placement in Dendryphantes: D. nigro- that is, if they are closely related to the maculatus (Keyserling), most recently type species of Dendryphantes. Other- placed in Eris (Kaston, 1973). Several Old wise, we would merely be worsening Den- World species placed in Dendryphantes, dryphantes' status as a catch-all genus and including D. fusconotattis and D. chuld- adding to it the confusion of changing the ensis, appear very closely related to D. generic placement of many common spe- nigromaculatus (see figures of Proszynski, cies. As noted, only D. nigromaculatus 1971b, 1982). Like the Old World D. has- among New World species is a strong can- tatus (the type species) and D. rudis, D. didate to stay in Dendryphantes. nigromaculatus has a slightly elongate Beata (Figs. 77, i09-ii2). The limits of body dully marked. Perhaps the best char- the genus Beata have been greatly over- acter that strictly delimits Dendryphantes estimated (Simon, 1903; Chickering, 1946). is the presence of a fold of embolic he- Because the type species of Beata is fissi- matodocha that lies across the basal part dent (it has a bifid retromarginal cheliceral of the embolic base, covering the wrinkles tooth), it has not only been removed from there (Figs. 103-108). If this character is the dendryphantines (Simon, 1903) but has used to delimit the genus, then it would also been burdened with diverse dendry- be a small genus of mostly Palearctic spe- phantines that happen to have a similarly cies. The placement of nigromaculatus in bifid tooth (Simon, 1903; Chickering, Dendryphantes is further supported by its 1946). Note that the tooth is better consid- sharing with D. rudis, D. fusconotatus, ered a single bifid tooth rather than two and D. chuldensis a much elongated prong fused teeth because the inner boundary of coming off of the base of the embolus and the cuticle does not extend to the tip of curving toward the cymbium (Figs. 106, the second cusp. The second cusp shows 108). The embolus therefore appears to all gradations of development in the den- have two rami, much as in Pelegrina fla- dryphantines, with most lacking it, some uipedes, though not homologous according showing a slightly swollen margin (e.g., to reasoning given below. Species of the Pelegrina proterva. Fig. 10), and others other two major groups of Old World den- having a well-developed cusp. It is there- dryphantines, the "Eris" nidicolens group fore best to place far less emphasis on this (Fig. 61) and the genus Rhene (Fig. 52), character. Beata magna G. & E. Peckham lack the fold across the embolic base seen (Fig. 109), the type species of Beata (by in Dendryphantes. Though Rhene species monotypy), bears few resemblances to most often have a prong arising from the base of the other fissident dendryphantines, in- of the embolus, in Rhene it is not curled stead having many more resemblances with toward the cymbium and, instead, is erect the other robust-bodied dendryphantines as in Beata and the mannii group. Rhene previously placed in Dryphias, Homalat- has been considered a close relative and foic/^s, and Anamosa. The following char- possibly a synonym of Dendryphantes acters, which appear derived within the (Proszynski, 1973b), but a number of other subfamily, delimit this group containing features of Rhene such as the presence of Beata magna: epiandrous fusules and the concave retro- marginal loop of the sperm duct of the 1. Tibial apophysis narrow and bent to- palpus also cast some doubt on this place- ward the ventral, almost paralleling a ment. ridge on the tibia below it (Fig. 77). Pelegrina Jumping Spiders • Maddison 237

This tibial ridge is similar to that in The Proper Placements of Pelegrina (Fig. 78), but it is longer and Metaphidippus Species sharper in Beata. 2. First leg tibia dark and enlarged at least Now that the relationships of the true slightly compared to patella, even in Metaphidippus and the limits of Dendry- females. phantes and Beata have been reconsid- 3. Carapace distinctively wide and high, ered, we are in position to discuss how the higher than in Sassacus, Agassa, and various groups within Metaphidippus Rhene, wider than in Sassacus and might be dispersed. Some of these conclu- Agassa. Unlike Zijgohallus, but like sions are summarized in Table 3. Sassacus and Agassa, the carapace is Four groups are here removed from wide well past the posterior eyes before Metaphidippus, as discussed in subsequent it abruptly drops and narrows. sections. The galathea group is transferred 4. Carapace scales erect (in at least some to the genus Pelegrina (and its species re- but perhaps not all species). vised), the harfordii group to the genus 5. Retromargin of base of embolus with Phanias, the mylothrus group to the new prong rising parallel to apical erect por- genus Terralonus, and the castaneus group tion of embolus (Fig. 110). Such a prong to the new genus Ghelna. is also seen in Rhene (Fig. 52) and the Two species groups occurring in the Metaphidippus mannii group (Fig. United States are retained temporarily in 499). Metaphidippus pending further study: the mannii group and the vitis group. The mannii group is discussed later in connec- The following species are placed in Bea- tion with the revision of its U.S. species. ta and NEW COMBINATIONS therefore The status of the vitis group (Figs. 27, 59), established: Beata hispida (G. & E. Peck- including Metaphidippus vitis (Cocker- ham) (Figs. 77, 110-112), Beata inconcin- ell), M. texanus (Banks), M. mathetes na (G. & E. Peckham), Beata maccunii (Chamberlin), and Dendryphantes me- (G. & E. Peckham), and Beata rustica (G. lanomerus Chamberlin, is not clear. These & E. Peckham). Also included is Beata species have a characteristic hooked em- longipes F. P. -Cambridge, which may be bolus (Figs. 27, 59) and are small, some- the male of B. magna. Dryphias (type spe- what elongate, and brown to black and cies maccuni by original designation) is a shiny. Metaphidippus vitis was placed in NEW SYNONYM of Beata. The genus Sassacus by Hill (1979) on the basis of scale Beata as here delimited excludes B. digi- morphology and courtship, but scale mor- tata (= Pelegrina galathea) and B. var- phology is known in only few dendry- iegata (= Pelegrina variegata), B. alhopi- phantines, and a similar courtship is also losa (= Gastromicans albopilosa), B. fla- seen in many other dendryphantines volineata (= Nagaina incunda), B. ce- (raisedforward, Table 2, character 9). Fur- phalica F. P. -Cambridge, B. jubata (C. L. thermore, the genitalia of M. vitis are very Koch), B. munda Chickering, B. pernix different from those of the true Sassacus (G. & E. Peckham), B. venusta Chicker- and, instead, are similar to those of the ing, B. wickhami (G. & E. Peckham), and neotropical species placed in Sassacus such B. zeteki Chickering. No new placements as S. arcuatus, which may better be placed are suggested for the last-mentioned seven in the genus Ramboia (see Bauab-Vianna species. Other species placed in the genus and Soares, 1982). The vitis group, here but probably not belonging there are B. retained in Metaphidippus, may eventu- cinereonitida Simon, B. germaini Simon, ally find its place in a primarily neotropical B. lineata (Vinson), and B. striata Pe- genus. trunkevitch. Some species placed in Metaphidippus 238 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

belong in Bagheera or Messua, as already primitive as it is also found in Synageles noted. Metaphidippus paiutus Gertsch is and Admestina (Figs. 29, 30). a Sassacus (Sassacus paiutus (Gertsch), 3. Courtship with first legs raised, for- NEW COMBINATION), possibly a syn- ward, and parallel (Fig. 118) and waved onym of S. papenhoei. Other species of asymmetrically so that the leading leg Metaphidippus require further study be- on sidles is waved exclusively or more fore their placement can be settled. Among strongly (seen by me otherwise only in the neotropical species listed under Me- Anicius). taphidippus by Bonnet (1957), M. longi- 4. Small blunt teeth on the embolus (Fig. palpus F. P. -Cambridge, M. nitidus (G. & 20). Many other dendryphantines (e.g., E. Peckham), and perhaps M. taylori (G. Pelegrina montana. Fig. 204) have & E. Peckham) seem to belong to Parna- teeth on the embolus, but their teeth enus, M. pallens F. P. -Cambridge in Eris, are sharp. The blunt teeth are lacking, M. perfectus (G. & E. Peckham) (Fig. 60) however, in some species of Phanias in Selimus, and M. tropicus (G. & E. Peck- (e.g., P. watonus). ham) (Fig. 62) in Bryantella. For the re- 5. Longitudinal bands of white scales, in- maining neotropical species (see Proszyn- stead of passing below and beside the ski, 1990), I have no placement to suggest. posterior eyes as in other dendryphan- tines, pass around and directly poste- Phanias F. 1901 P.-Cambridge, riorly from the posterior eyes. The dis- tribution of this character is not well Type species, Phanias flavostriatus F. P.-Cambridge, 1901, by monotypy. known.

The unusual, elongate dendryphantines The following NEW COMBINATIONS of the harfordii group (Figs. 20, 36, 47, are established: Phanias alheolus (Cham- 70) include several species in the western berlin & Ivie) (Figs. 20, 70), Phanias con- United States but more in the high- many coloratus (Chamberlin & Gertsch), Phan- lands of Mexico. A generic name is avail- ias dominatus (Chamberlin & Ivie), Phan- able for this group, Phanias F. P.-Cam- ias furcifer (Gertsch), Phanias furcillatus bridge, 1901, based on Phanias flavostria- (F. P.-Cambridge), Phanias harfordii (G. tus, described from two females from Om- & E. Peckham) (Fig. 47), Phanias mon- ilteme, Mexico (BMNH, examined), and ticola (Banks), Phanias neoniexicanus previously considered to belong in the (Banks), and Phanias watonus (Chamber- Marpissinae. The species of the Metaphi- lin & Ivie). Phanias marginalis Banks (type dippus harfordii group are therefore specimen examined) is a Menemerus, not transferred to Phanias. The members of a Phanias, while Phanias salvadorensis the Phanias share these genus characters, Kraus may be either a Phanias or an An- which be within the may apomorphies icius. Also included in Phanias are at least subfamily: 15 undescribed species from Mexico and 1. Tegular ledge expanded so as to cover the southwestern United States. Phanias the tegular shoulder (Fig. 47). The teg- may be placed near the base of the sub- ular ledge of other dendryphantines and family, for it has two features that are ar- other salticids is not so expanded. guably ancestral for the subfamily, name- 2. Embolic hematodocha reduced and ly, the presence of epiandrous gland fu- sclerotized prolaterally and basally. In sules (Machado, 1951; Table 2, character at least some species, much of the ex- 6; see also Maddison, 1988), which it shares pansion occurs from out of the tegular with Hentzia, "Beata" wickhami, ^^Eris" ledge instead of from the prolateral nidicolens, Rhene, and groups apparently dorsal surface (back side) of the tegul- related to dendryphantines (euophryines, Phle- um (Fig. 36), but this feature may be synagelines, ballines, Mopsus, Itata, Pelegrina Jumping Spiders • Maddison 239

gra, though not Neon), and a concave certainly they do not belong in the genus sperm duct loop along the retromargin of Metaphidippus, because they lack the the tegulum (Fig. 20), which it shares with characters of the Bagheera group of gen- some Hentzia, "Eris" nidicolens (Fig. 61), era. I have chosen to describe a new genus Rhene (Fig. 52), some Tutelina, Anicius, for the group to remove it from its uneasy euophryines (Fig. 19), and Admestina. placement in Metaphidippus. I do this with some hesitation, given the overabundance Terralonus new genus of obscure genera in salticids, but the my- lothrus group apparently does not reach Type species, Dendrijphantes mylothrus Chamber- the where it have found lin, 1925, here designated. Name treated as mas- Neotropics might culine. an available generic name, and so it is un- likely to be synonymized soon. Describing Description and Diagnosis (Figs. 22, 44, a genus allows easier discussion of this dis- 68). These western North American spe- tinctive group. The three species associ- cies are unusual among dendryphantines ated with the group for the first time (fra- in being ground-dwelling, usually on ternus, vittatus, and versicolor) can there- ground more or less barren of vegetation, fore be moved directly into Terralonus often under rocks. The body shape and without being temporarily sentenced to markings are distinctive and uniform Metaphidippus. throughout the group. They are somewhat elongate and have relatively low-contrast Ghetna new genus mottled markings of coarse brown or gray Type species, Attus castaneus Hentz, 1846, here des- pubescence. The embolus is long and its ignated. Name treated as feminine. base is more longitudinally directed than is usual in dendryphantines (Fig. 22), ex- Description and Diagnosis. These east- cept for T. californicus, which has a more ern North American species, like the spe- typical embolus (Fig. 44). cies of Terralonus, are ground-dwelling, Included Species. The following species though in more mesic habitats. They share are moved to Terralonus: Maevia califor- a dark granulate carapace with fine golden nicus G. & E. Peckham, Dendryphantes scales, posterior lateral spines on the first mylothrus Chamberlin, Dendryphantes tibia displaced anteriorly, reduced spines unicus Chamberlin & Gertsch, Metaphi- on the first femora, first coxae nearly dippus shaferi Gertsch & Riechert, Icius touching, and the female palpus slightly versicolor G. & E. Peckham, Menemerus swollen. The embolus, at least in the first vittatus Banks, and Menemerus fraternus two species mentioned and perhaps in all, Banks (type specimens of last-mentioned is twisted so as to wind the embolic he- three species examined). This establishes matodocha around the embolus much as the NEW COMBINATIONS: Terralonus in the Bagheera group of genera, though californicus (G. & E. Peckham), Terra- the twisting takes a very different form. lonus mylothrus (Chamberlin), Terralon- Included species. The species Attus cas- us unicus (Chamberlin & Gertsch), Ter- taneus Hentz, Metaphidippus barrowsi ralonus shaferi (Gertsch & Riechert), Ter- Kaston, Icius sexmaculatus Banks, and ralonus versicolor (G. & E. Peckham), Ter- Icius canadensis Banks are here moved to ralonus vittatus (Banks), and Terralonus Ghelna to establish the following NEW fraternus (Banks). COMBINATIONS: Ghelna castanea Discussion. By appearance, these are not (Hentz), Ghelna barrowsi Kaston, Ghelna typical dendryphantines—two of the spe- sexmaculata (Banks), and Ghelna cana- cies were described in the genus Mene- densis (Banks). merus and have remained there to this Discussion. As with Terralonus, the rel- day. Their relatives are unclear, but almost atives of Ghelna are unclear but, likewise. No. 4 240 Bulletin Museum of Comparative Zoology, Vol. 154,

and two rami are not near [he genus Metaphidippus. The opening bearing (Figs. 3, is sim- 190-216, 220-224) is a di- justification for a new generic name generally good feature for the but it is ilar to that for Terralonus. agnostic genus, absent in a number of species. Tibial ventral to THE GENUS PELEGRINA apophysis stout; just apophysis '' "'""^^^y ^ ""^^^ ^^'^- ^^^' de^^^^P^^ ^"^^ rnMiNvjrviNii_i_»^,FRANRANILLO 1930c^ ^ second apophysis in some species (Jur- 44. in Pelegrina Franganillo, 1930: Type species by cata group) or a wide flange other spe- and original designation monotypy Pe/egrma gen- j^jgg (arizonensts group). Females gray, iculata (= P. proximo). Franganillo yellow, or brown with mottled markings Notes on Synonymy. The problem of of four prominent pairs of pale spots on the abdominal dorsum the generic name to be given to the gal- (see, e.g., Figs. 2, athea group was not an easy one to solve. 263, 269, 358, 382). Epigynal openings rel- In my thesis (Maddison, 1988), I concluded atively long. Among small dendryphan- of have that a new generic name was needed for tines, the species Pelegrina per- the the group, because (1) it clearly does not haps best-developed epigynal flaps belong with Metaphidippus and (2) the (Figs. 236-255), which are the teardrop- lateral rims of the The group is arguably monophyletic and of re- shaped openings. the spectable size for a genus, and (3) it ap- flaps are usually convex and overlap peared that there was no genus whose type medial rim of opening. All species of Pe- fell within the However, legrina examined have the same chro- species group. = subsequent investigation has indicated that mosome complement, 2n6 26-1- XXO, an obscure Franganillo name, Pelegrina, as is prevalent throughout the family, should be applied to the galathea group. Monophyly. Thirty-eight species are in- Although the revival of obscure names is eluded in Pelegrina. Most of these species often undesirable, in Fe/egrfna 's case there can be easily recognized as belonging to is little harm because no other published the genus by an experienced identifier on name is available for the group. In 1930, the basis of body form, size, and markings, Franganillo described Pelegrina and based but to articulate precisely characters that it on Pelegrina geniculata Franganillo, could serve as evidence for monophyly is which he placed in the section Unidentati, more difficult. The following characters subfamily Heliophaninae. As discussed un- support the monophyly of the genus, der the description of P. proxima, P. gen- though none provides a simple, strict de- iculata is here considered a junior syn- limitation. Some of the characters delimit onym of Dendryphantes proximus G. & a group slightly smaller than the genus, E. Peckham; thus, the name Pelegrina is others a group slightly larger. Thus, each applicable to the galathea group. character provides only indirect and par- Description and Diagnosis. Small to tial evidence for monophyly. medium-sized dendryphantines distribut- ed throughout North and Central Ameri- 1. Embolus with two terminal rami retro- ca. Males (Fig. 1) typically brown with lateral to opening (Figs. 3, 190-216, longitudinal bands of white scales on either 220-224). The sperm duct opening lies side of the carapace and abdomen. The on the prolateral side of the embolus, inverted white V-shaped marking on the often below the tip. Retrolateral to the forehead that contacts the AMEs distin- opening are two rami, one just distal to guishes Pelegrina from most other den- the opening; the other often elongate dryphantines, though it is not present in (see especially P. tristis, Fig. 197) and all Pelegrina. Legs often with annulate forming the retrolateral tip of the em- markings. The relatively wide embolus bolus. While other dendryphantines with the tip expanded retrolateral to its such as Tulpius, Phanias, and Tutelina Pelegrina Jumping Spiders • Maddison 241

have accessory rami emerging from the grina, except P. orestes, it reaches dis- embokis, none have such rami in the tally to the fang opening. This contrasts position or form seen in Pelegrina. This with Dendryphantes, the mannii group, is perhaps the clearest character deUm- and most other dendryphantines, which iting the genus, but some Pelegrina ap- lack such wrinkles. The only other sal- pear to lack it {verecunda, tillandsiae, ticids seen with such a "secondary ser- bunites, orestes, arizonensis, helenae; rula" are (a) Phanias (four species ex- Figs. 217-219, 225-227), while others amined), in which the wrinkles are re- have the rami but in a much modified stricted to a depression that does not form (flavipedes and related species; reach the opening; (b) Selimus sp. and Figs. 201-203). These problems are dis- '^Beata" octopunctata, in which the cussed later. wrinkles are long and regular; and (c) 2. Hematodocha of embolus bulges as far Eris militaris, in which the wrinkles do distally as the base of the erect portion not quite reach the opening. Except for of the embolus (Fig. 3; Table 2, char- E. militaris, the wrinkles are of differ- acter 7). This feature is present ent form, suggesting they are not ho- throughout Pelegrina, including P. or- mologous. estes and P. bunites. In other dendry- 5. Distinct cheek bands on the male face phantines examined, including the (Figs. 1, 258, 264, and so on). Though mannii group and Eris, the hemato- other dendryphantines may have pale docha joins the retrolateral edge of the scales on the side of the face under the embolic base more basally so that the ALEs, in most these pale scales do not hematodocha fails to bulge as far dis- form a distinct band separated from the tally (Figs. 22, 23). side bands by a dark area. Such a sep- 3. Epigynal flaps well developed, long, and aration of the cheek and side bands is wide and not descending into the open- also seen in some species of the mannii ing posteriorly. This character is diffi- group (Figs. 493, 514, 534). It is lacking cult to assess, for there are other den- in other dendryphantines except one dryphantines with well-developed flaps, species from examined, pos- the char- though in these the flaps do not exactly sibly an Admirala sp., though match those of Pelegrina, being either acter has not been surveyed intensively. varie- much shorter {Phidippus, Fig. 67; Bel- Some species of Pelegrina {P. do not have lota), narrower {Beata; Figs. 109, 112), gata. Fig. 447), however, a distinct cheek band. or less convex (e.g., "Pseudicius^^ siti- inverted mark of culosus). The flaps of most Pelegrina 6. An V-shaped pale differ from those of the mannii group scales on the forehead, contacting the and so and Eris, which have weak flaps that AMEs (Figs. 1, 258, 264, on). males show this fore- descend into the opening at posterior Most Pelegrina some lack it end (Figs. 66, 256, 257); the flaps in the head band, though (e.g., the character has two most problematical Pelegrina spe- P. aeneola). While it is lack- cies, P. bunites and P. orestes, are not been surveyed thoroughly, somewhat like those of the mannii ing in other dendryphantines except for Tutelina hart group. a few species (e.g., it). with 4. Wrinkles present on anterior margin of 7. Male courtship prolonged "crouch" In the of male cheliceral fang (Fig. 11; Table 2, display. courtship Eris the character 8). Running parallel to the Pelegrina, militaris, Metaphi- mannii and in- serrate edge of the fang (Hill, 1977b), dippus group, Nagaitia there is a crouch in just anterior to it, is a line of transverse cunda, display, first are held low and wrinkles, which appears like an irreg- which the legs horizontal and below ular secondary serrate edge. In Pele- forward, usually 242 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

the level of the body (Figs. 125-128, from the brief stage II seen in other 134, 140, 162 for Pelegrina; Figs. 124, salticids, in being much more pro- 178, 184 for Eris and mannii group; longed, performed farther from the fe- Peckham and Peckham, 1889; Rich- male, and having a more rigid appear- man, 1982: fig. 3; Table 2, character 9). ance. The crouch display has been ob- The more distal segments may be raised served in all Pelegrina and mannii (e.g., Pelegrina kastoni. Fig. 140), but group species examined, with the fol- at least the femur is low. The male pro- lowing exceptions: in P. furcata a strik- ceeds toward the female in this pose, ingly different display (the semaphore waving the first legs at low amplitude display) is seen, whereas in Pelegrina if at all. The body is held horizontal, in arizonensis, P. chalceola, and Meta- many species close to the substrate phidippus diplacis only a raisedspread (hence the name "crouch"). This pose display was seen. The lack of observed contrasts with that seen in most other crouch display in these latter species dendryphantines (Table 2; Peckham calls attention to the difficulty of using and Peckham, 1889, 1890; Crane, the character. When seen, the crouch 1949a; Richman, 1982; Jackson, 1978; display is distinctive and can be con- Hill, 1977a; and my own unpublished sidered present. However, when not observations on about 50 in species seen, it may still be characteristic of the which the first are raised legs generally species but not observed because the and spread (the raisedspread display, male simply failed to perform it, re- 113-115, 121; Phidippus, Figs. e.g., maining longer than usual in the raised- Paradamoetas, Zygoballus) or raised spread display. Nevertheless, the crouch and held forward (the raisedforward display was scored as absent in these display. Figs. 118-120; e.g., Phanias, Pelegrina species because the males Sassacus papenhoei, Dendryphantes were observed for a reasonable or held horizontal and sample nigromaculatus) , of Another with the spread very wide (the lowspread dis- displays. problem character is the occurrence of a similar play, Figs. 116, 117; e.g., Messua lim- first in Tutelina and Hentzia bata, Tulpius). The distinctions among leg pose these poses are, of course, vague. It (Figs. 122, 123). However, the exact leg should be noted that Pelegrina does poses and motions in these other genera have a raisedspread display, generally differ in a number of respects from those performed with the carapace held high of Pelegrina. and the abdomen low, but this is per- 8. Ridge under tibial apophysis (Figs. 78, formed usually when the male is far 389, 421, 427; Table 2, character 10). from the female. A Pelegrina male thus Under the tibial apophysis is a ridge often begins with a raisedspread display that in extreme cases can make the and then proceeds to a crouch display apophysis appear bifid (e.g., Pelegrina as he approaches the female. Insofar as bicuspidata) . The ridge is present other dendryphantines often reach with throughout Pelegrina (except fla- the legs parallel, forward, and low just vipedes and similar species) but is also before mounting and copulation (stage present in some species of the mannii II courtship. Crane, 1949b), as in the group (mannii, diplacis), Beata (Fig. crouch display, it may be claimed that 77), and in Tulpius hilarus. It is poorly the crouch display is merely stage II developed in Eris militaris (Fig. 79), courtship and, thus, not restricted to Pe- Bellota wheeleri, and Metaphidippus legrina and relatives. Indeed, the crouch vitis. It is lacking in other dendryphan- display may represent a prolonged stage tines including Dendryphantes (Figs. II. If so, however, it is still distinctive 75, 76). Pelegrina Jumping Spiders • Maddison 243

The preceding characters give reasonable structures seen in the M. arizonensis support to the monophyly of the genus group." Perhaps the similarity they no- Pelegrina, though none provides a strict ticed was the looping of the duct just inside dehmitation. Even if the genus as consti- the opening. The epigynum of D. czeka- tuted here is not monophyletic, the char- nowskii is much like that of D. nigroma- acters provide good evidence that it is at culatus and D. fusconotatus. This looping least mostly monophyletic, to the extent in both Dendryphantes and the arizonen- that monophyly could probably be sis group is related to the rotation of the achieved by including or excluding only a epigynal flaps medially at the posterior end. few troublesome species. The following In Dendryphantes, the rotation reaches 90°; discussion regards the Pelegrina species in the arizonensis group, it is much more that fail to show some of the characters extreme, 180-270°. This rotation must not supposedly delimiting the genus and why be considered a critical character; similar I have chosen to include these species in rotations are seen in the Pelegrina pervaga the genus. Whether or not species in the group (P. kastoni), in Phidippus octo- mannii group might be included in Pe- punctatus, and in Agassa (as compared to legrina is discussed in the introduction to Sassacus). Indeed, in other respects the the revision of the species of the mannii epigyna of the arizonensis group and Den- group. dryphantes are rather different, with the Pelegrina arizonensis and P. helenae. flaps being on the surface in the former These species lack the two rami on the (as in other Pelegrina, Fig. 251), whereas embolus. Instead, the embolus is blade- they descend beneath the opening as a sim- shaped, shifted retrolaterally, and concave ple ridge in the latter (Fig. 65). on the exposed (ventral) surface (Figs. 217, Pelegrina flavipedes, P. flaviceps, and 218, 422, 428). However, there are a num- P. exigua. The biramous embolus of this ber of other characters that would other- group (Figs. 201-203) might be interpret- wise place the two species within Pelegri- ed either as arising from an embolus like na. These species have distinct cheek and that of Pelegrina sahinema (Fig. 198) by forehead bands, a ridge just ventral to the more deeply splitting the two terminal rami tibial apophysis, and the male fang with of the embolus or as arising from an em- wrinkles as in other Pelegrina. The pe- bolus like that of Dendryphantes rudis and culiar embolus might be derived from that D. nigromaculatus (Figs. 106, 108) by pro- of a typical Pelegrina embolus by twisting longation of the retrolateral projection on about its longitudinal axis so as to reverse the shoulder of the embolus. The fla- pro- and retrolateral edges and to present vipedes group lacks a ridge under the tibial the embolus's concave surface, normally apophysis, which otherwise seems char- facing inward to the cymbium, outward acteristic of Pelegrina, thus supporting the toward the front. This is indicated by the interpretation that the flavipedes group presence of a ridge cutting across the face does not belong with Pelegrina. However, of the embolus joining the prolateral sur- there is more compelling evidence that the face of the base with the retrolateral edge flavipedes group is derived from within of the embolus, the position of the opening Pelegrina. Like the pervaga group, the^^a- on the retrolateral side, the concave ex- vipedes group members are conifer dwell- posed face of the embolus, and a pro- ers with yellow chelicerae. Like Pelegrina, nounced furrow on the embolis hemato- they have wrinkles anterior to the fang docha as if folded inward. Cutler and Jen- serrula and an embolic hematodocha bulg- nings (1985) noted that "internal epigynal ing distally, cheek bands on the male face, structure of his [Proszynski, 1982:1] and a crouch display in male courtship D[endryphantes]. czekanowskii bears a (Fig. 127). Like Pelegrina and a few other close resemblance to the internal epigynal dendryphantines, the embolic base is well 244 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

sclerotized, with few wrinkles over most hematodocha to the shoulder of the em- of its exposed surface. Finally, if the fla- bolus, and the forehead band contacting vipedes group were derived from those the AMEs. On balance, then, a case can Dendryphantes with long retrolateral pro- be made for tentatively describing bunites longations on the embolus shoulder (rudis, in Pelegrina. The situation with orestes is the should have more difficult. orestes lacks the nigromaculatus) , group Pelegrina the synapomorphy for Dendryphantes, the cheliceral fang wrinkles of Pelegrina, and fold across the embolic base. The fla- one character, the presence of a ridge on vipedes group lacks this fold. the chelicera (Fig. 483), gives positive ev- Pelegrina furcata. This species has two idence to place orestes in the mannii group, terminal rami on the embolus, robust epi- though the ridge is especially weak and its in gynal flaps, and cheek bands, but court- not present all males. However, orestes ship display is unlike that of any others in lacks the bulge just dorsal to the tibial the genus. The first legs are held wide and apophysis characteristic of the mannii high, unlike Pelegrina, Eris militaris, and group and does have the bulging embolic the mannii group, but like most other den- hematodocha characteristic of Pelegrina. dryphantines, and waved in a distinctive Because of this, orestes will be described semaphore-like fashion (Fig. 121). in Pelegrina, though it may eventually Pelegrina verecunda. Arizonan speci- have to be moved. mens lack the two distinct terminal rami Natural History. Species of Pelegrina on the embolus (Fig. 219), but Chihuahuan are found in various habitats from the Arc- specimens identified as this species have tic to the tropical lowlands of Central the rami. America. While most species in Mexico Pelegrina tillandsiae. This species lacks and Central America appear to occur in the two terminal rami on the embolus (Fig. the highlands (cloud forest and oak-pine 225) and is in many respects atypical. It is zones), there are some lowland tropical tentatively included in Pelegrina because species (P. sandaracina and P. yucate- its epigynum shows strong flaps that, as in cana). All species in the genus are pri- Pelegrina, do not descend into the open- marily dwellers on foliage, being only oc- ings (Fig. 254). casionally found on the ground. Most other Pelegrina orestes and P. bunites. These dendryphantines are also foliage dwellers, species present the greatest problems with though some dendryphantines, in partic- inclusion in Pelegrina, and I might have ular those with more elongate and dully treated them as belonging to the mannii marked gray or brown bodies, are ground group or elsewhere. They lack the two ter- or bark dwellers (Terralonus, Ghelna, some minal rami on the embolus characteristic Phanias species). A number of species of of Pelegrina (Figs. 226, 227), though at Pelegrina appear to be most common on least in P. orestes the embolus is obliquely or restricted to particular sorts of plants: truncated distal to the opening and has one the flavipedes group to various confiers, ramus well separated from the opening. the pervaga group and P. balia to junipers, On the other hand, the epigynal flaps of P. clemata and P. helenae to sagebrush bunites are flatter and narrower than in {Artemisia tridentata), and P. tillandsiae other Pelegrina, more as in the mannii to {Tillandsia usneoides). group (Figs. 225, 481). Pelegrina bunites, Other species do not appear so specialized though, lacks the bulge above the tibial to particular plants, yet in my collecting apophysis characteristic of the mannii they do seem to prefer certain habitats: P. group and has three characters that might proterva occurs in forest understory, at least argue for the placement of bunites in Pe- in the south of its range; P. galathea, in legrina: the occurrence of wrinkles on the fields; P. variegata, in desert scrub; P. cheliceral fang, the bulging of the embolic montana, in streamside vegetation; and P. Pelegrina Jumping Spiders • Maddison 245

insignis, on low plants in fields and bogs. 1. Distinct markings on yellowish male A number of southwestern species are most palpus, consisting of prominent patches commonly collected from oaks. One gen- of white scales on the femur, tibia, and eralist species is P. aeneola, which is found cymbium, interrupted by dark hairs on on trees and herbs of various sorts in the the patella. Other Pelegrina have no Pacific Northwest, though not usually in white scales on the tibia, or fewer on the arid regions. The silken retreats and the tibia than on the patella. egg sacs are constructed among the foliage 2. Cheek band extended posteriorly par- on which the adults are collected. allel to the side band, separated from the side band by dense band of dark PHYLOGENY WITHIN PELEGRINA hairs (Figs. 304, 309, 314). This yields the of white-black-white While insufficient evidence was found appearance carapace side bands. Other Pelegrina to indicate the basal divisions of Pelegrina, have dark setae between side and cheek the delimitation of a number of smaller bands, but in none is the cheek band so groups can be made (see cladogram given horizontal and the dark hairs so dense. in Fig. 129). One clearly delineated group One species in the mannii group, Me- is the flavipedes group, whose three mem- taphidippus emmiltus, has a superfi- bers (flavipedes, flaviceps, and exigua) cially similar pattern. share the following characters derived within the genus: These two groups, the flavipedes and pervaga groups, may together form a T Embolus deeply bifid (Figs. 201-203). monophyletic group, as delimited by the All other species of Pelegrina have the following: division between the terminal rami of the embolus not nearly so deep as in 1. Chelicerae yellow in males. Other Pe- the flavipedes group. Other dendry- legrina have dark brown chelicerae ex- phantines have either a simple embolus cept southern males of P. tillandsiae. or one with accessory rami different Brown chelicerae are present in other from those in Pelegrina and the fla- similar dendryphantines such as Eris, vipedes group. Nagaina, Beata, and the mannii group 2. Chelicerae of male yellow with dark except Metaphidippus emmiltus, which spot in medial concavity (Figs. 319, 324, also has yellow chelicerae. 329, 334). The dark concavity is unique 2. Conifer dwelling. All members of the to this group. flavipedes group, and P. pervaga and 3. Asymmetrical circling of palps in male P. kastoni, are known to dwell more or courtship. During the crouch display, less exclusively on conifers. The habitat This con- the palps are waved fairly slowly at high of P. sabinema is unknown. amplitude in circles such that as one is trasts with the habitat of most other inhabit broadleaf rising the other is falling (Fig. 127). Pelegrina, which Though other Pelegrina may wave their trees, shrubs, and herbs. However, P. are palps out of phase, in none examined proterva, P. aeneola, and P. jurcata whereas P. is this asymmetrical waving made so known to frequent conifers, obvious by the large slow waves. No halia appears restricted to conifers. Also, other dendryphantines examined have the polarity of this character is unclear. em- such waving, except some Phidippus. Outside the genus, Metaphidippus miltus is a juniper dweller, whereas Eris well defined is Another group similarly species are often collected from coni- the pervaga group, consisting of pervaga, fers. sabinema, and kastoni, which share the The neoleonis group, including tristis following: 246 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

from Arizona and neoleonis from Mexico, tation; flap rotations in Terralonus and is distinguished by the broad, dark rotated Ghelna, for example, are in the oppo- epigynal flaps. Though their appearance site direction. is much more Hke that of typical brown- 2. Erect portion of embolus displaced to legged Pelegrina, they share the following retrolateral side and tegulum rotated characters with the preceding two groups somewhat clockwise (Figs. 422, 428). of yellow-legged species: The embolus displacement is seen in some other dendryphantines, but the 1. Retrolateral ramus of embolus much tegulum rotation is perhaps unique. elongate and curled to the prolateral 3. Embolus twisted about longitudinal axis The other Pele- (Figs. 196-203). only so as to reverse pro- and retrolateral with such a ramus is the grina long edges and to present the back side for- furcata group, though in that group all ward. but one have it to the species curling 4. Ridge under tibial apophysis uniquely retrolateral. The exception is morelos, developed into flange (Figs. 421, 427). which has a curl prolateral (Fig. 215), 5. Markings of female abdomen some- but because it seems very close to fur- what lineate (Figs. 425, 431). Lineate cata the curl is con- prolateral probably markings are also seen in P. tillandsiae vergent. and the Metaphidippus tnannii group. 2. Embolus very broad. Such a broad em- The of Central bolus is not found in other Pelegrina furcata group America except peckhamoruni. and southwestern North America includes 3. First curve of duct of epigynum broad. the widespread and common P. furcata as well as a number of rare Though unusual, a duct as broad is species (huachu- found in P. huachuca and P. morelos. ca, morelos, bicuspidata, volcana, ochra- cea). Apomorphies supporting the group The relationships of this proposed fla- are as follows: vipedes-pervaga-neoleonis clade are not 1. under tibial altogether clear, though the sickle-shaped Ridge apophysis unusually so as to form a sec- retrolateral ramus of the embolus of pro- strongly developed, ond 389; even terva, galathea, edrilana, and pallidata apophysis (Fig. stronger in the other in the This may be viewed as a preliminary version species group). is found in all it is of the very long ramus of this clade. Most species, although in some of of these species together with dithalea, lacking specimens furcata. 2. Wrinkles on the retrolateral basal proxima, and peckhamoruni have an angle edge of the embolic base either transverse or on the embolus just basal to the tip of the apically toward the retrola- retrolateral ramus (Figs. 259, 265, 271, 277, ascending teral 390-394, 404, 406, 411, 283, 289, 310) which may be a synapo- (Figs. 416). This contrasts with the wrinkles of other morphy for a large clade (Fig. 129), but and similar whether or not the angle is absent from all Pelegrina dendryphan- tines, which have wrinkles descending other Pelegrina is unclear (e.g., see Fig. toward the retrolateral. Pele- 215). basally have wrinkles sim- Another clearly delineated group is the grina insignis may ilar to those in the arizonensis group (Cutler and Jennings, furcata group. 3. convex, unlike oth- 1985), including the two species P. ari- Epigynal flaps very er zonensis and P. helenae. The apomorphies dendryphantines except Pelegrina and supporting the group are as follows: proxima peckhamorum. 4. Epigynum concave behind flaps, unlike 1. Epigynal flaps far rotated, at least 180° the case in Eris, the mannii group, and (Figs. 424, 430). No other dendryphan- Nagaina, though also seen in Pelegrina tines known to me have a similar ro- proxima, peckhamorum, and balia. Pelegrina Jumping Spiders • Maddison 247

Pelegrina furcata itself has a very pe- Females, however, pose many more culiar courtship display. Whether or not problems for identification. Though the this feature is shared by other members of abdominal markings and epigyna of fe- this group awaits their examination. males vary in a number of features, the The montana group includes three spe- differences can be subtle and difficult to cies: montana, insignis, and chaimona. describe. One might think of the abdom- These species share the following: inal markings as falling in two major cat- egories: those in which the paired white 1. Concavity on back of embolus restrict- spots dominate the dorsum (Figs. 263, 275, ed to distal half of erect portion. In 293, 382, 441, 451) and those in which the other Pelegrina species, the concavity dark patches between and beside the white on the back of the embolus (Figs. 7, 34, spots dominate the dorsum 281, 287, 35) extends from the base of the erect (Figs. 347, 358, 364, 377, 387). Epigynal features portion to its tip. Some Pelegrina, how- to note are the its have topography of surface ever, no clear concavity at all (e.g., and the size, convexity, color, and place- P. aeneola, P. chalceola). ment of the teardrop-shaped flaps cover- 2. Small, sharp denticles on front surface ing the openings. Even once experienced of embolus (Figs. 204-206). Other den- with these characters, an identifier can still dryphantines have denticles on the em- have difficulties with some specimens. The bolus, but they are usually on the re- problems are lessened within a trolateral surface or are of different given geo- graphical Because of this, form. This character has not been well region. separate female keys are given for five the surveyed, however. regions: eastern United States and Canada, the Among the remaining species of Pele- Great Plains, Pacific Coast Untied States grina are many that have a narrow em- and western Canada, Arizona and Mexico, bolus with small rami {aeneola, clemata, and Central America. Parts of the south- chalceola, balia, variegata, verecunda, western United States are therefore with- sandaracina, and others). However, this out a key to females of Pelegrina, namely, may be the primitive condition for the Texas and the Rocky Mountain states. For genus. No clear subgroups were found Montana and Wyoming, the Pacific Coast among these species. key can be used (except possibly for prairie For Colorado, Utah, and New IDENTIFYING SPECIES OF species). PELEGRINA Mexico, most identifications can be accom- AND THE METAPHIDIPPUS MANNII GROUP plished using the Pacific Coast and Ari- zona keys, though the Great Plains key will In general, the genitalia provide the best be needed occasionally. For Texas, the Ar- means of identifying species, but facility izona and Great Plains keys will usually in recognizing the distinguishing features suffice, but the eastern United States key may require some experience. Males are will be needed on occasion. In general, much more easily identified than females, mannii group females are not included in as the palpus and face markings provide the keys. Metaphidippus mannii is in- more readily described and interpretable cluded in the Pacific Coast key, but five differences than the differences in epi- other species in the group that occur in gyna. Take note especially of the width of southern California are not included; man- the erect portion of the embolus and the nii and chera are in the Arizona key, but size and orientation of the two terminal carmenensis is excluded. rami. Indeed, it is usually possible to iden- The keys are written for adult speci- tify males simply by referring to the two mens, but the keys for females will be of pages of Figures 190-235. A single key for some aid to identifying immatures based all species is given for males. on markings. A key for immature Pele- 248 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

central Ari- grina from Minnesota has been given by prolongation (Fig. 479); zona south to Oaxaca .. 37. bunites (part) Cutler (1981b). 5(4). Carapace and abdomen shiny dark or 258-538 the most com- Figures provide coppery brown contrasting strongly prehensive set of illustrations of the spe- with dense white cheek band and chel- iceral with small or no white cies, but important aid can be obtained patches, side bands (Figs. 178, 180, 493); pro- from Figures 130-189, which show living longation on retrolateral edge of base 190-235, which sum- specimens. Figures of embolus blunt or small (Fig. 494) marize the male emboli, and Figures 236- where sympatric with diplacis; British 257, which show the surface topography Columbia to California east to Idaho and central Arizona 40. mannii of the epigyna. - Carapace and abdomen with more ex- tensive white side bands (Figs. 182, Key to the Males of All Species of 503); cheek band not distinct from side Pelegrina and Those band (Fig. 503); prolongation of retro- Metaphidippus mannii Group Species lateral edge of base of embolus distinct Occurring in the United States* and long (Figs. 504, 505); along Pacific Coast of southern California and 1. Erect portion of embolus extremely nar- Baja California 41. row or tapers to point, lacking two ter- diplacis (part) 6(3). Erect of embolus minal rami (Figs. 226-235); lateral portion straight (Fig. face dark under margin of chelicera usually with ridge 233); eyes (Fig. 514) 43. chera near base of fang (Figs. 483, 493, 503, Erect 509, 514, 529); at least small patch of portion of embolus curves ven- white or orange scales on chelicerae; trally (Fig. 234); face extensively cov- western United States and southwest- ered with white scales (Fig. 529) ex- ern Canada, south into Central Amer- cept in Baja California Sur ica {Metaphidippus mannii group, in 44. carmenensis Erect of embolus trun- part, and some Pelegrina) 2 7(2). portion obliquely broad at base - Erect portion of embolus in most species cated, (Figs. 227, 484); and abdomen dusted with wide at tip and with two terminal rami carapace to brown scales (Figs. 190-225); chelicera lacking ridge beige light (Figs. 172, 483); Arizona and Mexico near base of fang (e.g.. Figs. 258, 264); scales on chelicerae varied; widely dis- 38. Orestes (part) tributed 10 Erect portion of embolus not so broad or truncated 2(1). Long patch of white scales on chelicerae, (Figs. 505, 510); markings dark brown with white longer than V2 length of chelicerae (Figs. 182, 184, or (Figs. 478, 493, 514, 529) 3 503, 509) mostly yellow (Fig. 176) - 8 White or orange patch small (Figs. 483, 503, 509) 7 8(7). Legs yellow, first legs fringed with white anterior 3(2). Embolus wide at base of erect portion (Fig. 176); median eyes ringed with chelicerae vertical rel- (Figs. 226-231) 4 red; and - weak erect Embolus very narrow (Figs. 516-523, atively (Fig. 534); portion of embolus thin south- 535) '. 6 very (Fig. 535); ern California to 4(3). Forehead white band lacking so that se- New Mexico 45. emmiltus tae above AMEs are dark (Figs. 493, (part) - 503); retrolateral edge of base of em- Legs with dark brown markings (Figs. anterior bolus with prolongation (Figs. 494, 499, 182, 184, 503, 509); median 505) 5 e\ es ringed with dark setae; chelicerae - White forehead band present and con- at least slightly divergent; embolus thick or Pacific of Califor- tacting AMEs above (Fig. 478); retro- thin; Coast lateral edge of base of embolus lacking nia and Baja California 9 9(8). Embolus wide at base of erect portion (Figs. 231, 504, 505); scales on chelic- erae white; body with bronze reflec- * The mannii group species in Mexico and Central tions (Fig. 182); prolongation on re- America are not included; they can be distinguished trolateral edge of base of embolus large from Pelegrina by their narrower embolus tip, which and distinct (Figs. 504, 505); southern lacks the two rami. Nagaina incunda, described later, California and Mexico is brown and yellow striped with the first legs brown _ 41. diplacis (part) - and the posterior legs yellow (Fig. 174). Embolus thin (Figs. 232, 510); scales on Pelegrina Jumping Spiders • Maddison 249

in North and Central America. 2. P. in the . 1 . proxima Maps 1 -7. Distributions of Pelegrina species. Pelegrina galathea 6. P. neoleonis in Mexico. 7. P. tristis in 3. P. d/matea in Arizona. 4. P. edr/Vana in Mexico. 5. P. proterva in North America. Arizona. 250 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Maps 8-15. Distributions of Pelegrina sp>ecies. 8. P. peckhamorum in the eastern United States. 9. P. sabinema in the southwestern United States. 10. P. pervaga in the central United States. 1 1 . P. kastoni in the southwestern United States. 12. P flavipedes in North America. 13. P. flaviceps in eastern North America. 14. P. exigua in the eastern United States. 15. P. clavator in Mexico. Pelegrina Jumping Spiders • Maddison 251

- chelicerae orange; body dark and dull No medial black spot on chelicerae (Figs. (Fig. 184); prolongation on retrolateral 304, 309, 314, 534); .southern United edge of base of embolus small or absent States and northern Mexico 20 (Fig. 510); central and northern Cali- 19(18). Forehead flat (Fig. 329); forehead dark fornia 42. tricolor brown in alcohol; bod\ and legs brown chelicerae 10(1). Clypeus with patch of white or yellow (Fig. 146); yellow laterally scales bet\\ een AMEs (not merely hairs (Fig. 329); southeastern United States overhanging chelicerae or circling an- north to Massachusetts and New York terior median eyes) 11 14. exigua (part) - - Clypeus without white or yellow setae Forehead bulbous (Fig. 324); forehead between AMEs, except perhaps for yellow in alcohol; body and legs pale hairs surrounding anterior median eyes (Fig. 144); chelicerae with dark spot and overhanging chelicerae 17 laterally (Fig. 324); northeastern Unit- ed States and southeastern Canada 11(10). Chelicerae with large white or yellow patch of scales medially, at least for 'A 13. fla viceps length of chelicerae (Figs. 159, 437, 20(18). Erect portion of embolus very thin (Fig. 442, 457); Mexico and Central Amer- 535); AMEs ringed with red; first legs ica 12 fringed with white (Fig. 176) _ Chelicerae lacking pale scales (Figs. 282, 45. emmiltus (part) United States east of 288, 319, 334); - Erect portion of embolus thicker (Figs. 14 Rocky Mountains and Canada 305, 310, 315, 474); AMEs ringed with ramus of embolus much lon- 12(11). Retrolateral white or brown; legs not fringed 21 than (Fig. 221) ger prolateral 21(20). Cymbium yellow; band of dark setae un- 32. pallidaia der carapace side band (Figs. 304, 309, - Retrolateral ramus of embolus small, 324); embolus wide at tip (Figs. 305, to 220, 224) 13 subequal prolateral (Figs. 310, 315); dwelling on conifer; Kansas embo- 13(12). Pale markings usually yellowish; west to Arizona 22 lus small and tapers to tip (Fig. 224); dark Cymbium dark distally; no band of lowland 35. sandaracina setae under carapace side band (Fig. - Pale white; embolus broadly markings 472); embolus tapers to narrow tip (Fig. truncate 220); montane (Fig. 474); dwelling on Spanish moss, Flor- 31. clavator (part) ida and North Carolina west to Texas em- 14(11). Chelicerae yellow (Figs. 319, 334); 36. tillandsiae (part) divided bolus deeply (Figs. 320, 330); retro- 22(21). Clypeus brown (Figs. 304, 309); lacks white scales; dwellers cymbium lateral ramus of embolus long (Figs. on conifers 15 198, 199, 305, 310); embolus broad at Chelicerae brown (Figs. 282, 288); em- base of erect portion 23 bolus not divided 260, deeply (Figs. with white band except cen- of Clypeus 266); cymbium with dorsal patch ramus of trally (Fig. 314); retrolateral white scales; habitat varies 16 embolus short (Fig. 200); embolus 15(14). First tibia yellow or with thin black stripe; re- rectangular, narrow, and displaced retrolateral ramus of embolus thick, kastoni (Fig. 315) H. than trolaterally only slightly thinner prolateral embolus 23(22). Abdomen brown above; (Fig. (Figs. 201, 320) 12. flavipedes 305) wider than in pervaga 9. sabinema First tibia dark; retrolateral ramus much Abdomen with central longitudinal pale thinner than prolateral (Figs. 203, 330) embolus stripe as in females (Fig. 313); 14. exigua (part) sabinema (Fig. 310) narrower than Embolus to tip (Fig. 284), with 16(14). tapers 1 0. pervaga long hooked retrolateral ramus (Fig. under tibial apophysis usually de- 283) 5. proterva 24(17). Ridge into acute second apophysis Embolus broad at 290), veloped very tip (Fig. on embolic base (Fig. 389); wrinkles retrolateral ramus short (Fig. 289) transverse or ascending apically to- 6. peckha moru m ward the retrolateral edge (Figs. 390, 17(10). Chelicerae yellow (Figs. 304, 309, 314, 404, 406, 411, 416); southwestern 324, 329, 534); dwellers on conifer and 1" United States to Panama {jurcata Spanish moss - 2^ 24 group) - Chelicerae brown (e.g., Figs. 258, 264) At most small ridge or broad flange un- 18(17). Medial black spot on chelicerae (Figs. der tibial apophysis (Figs. 78, 421, 427); 324, 329); embolus deeply divided wrinkles on embolic base descending 325, 330); eastern and central (Figs. toward the retrolateral edge United States and Canada _ - 19 basally 252 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 Pelegrina Jumping Spiders • Maddison 253

315. (e.g.. Figs. 438); widely distrib- chelicerae robust; embolus appears to uted 29 in taper ventral view but iti an ()bli(|ue Rami of embolus small and 25(24). subequal view the two small subequal rami are 213, 214) (Figs. 26 easily seen (Fig. 222); abdomen with - Retrolateral ramus of embolus much long, strong white spots (Fig. 166); arid than longer prolateral (Figs. 212, 215 regions of Mexico and Central Amer- 216) 27 ica 33. variegata (part) 26(25). White patches on chelicerae extend to at Side and cheek bands separate (Fig. 452); least 1/2 their length (Fig. 403); embolus chelicerae narrower; embolus weaker more or less straight (Figs. 213, 404); with two rami not easily visible; ab- Panama 23. volcano domen with unusual transverse bands - White on chelicerae small patches (Fig. (Fig. 169); seasonal tropical forests of embolus bent to 405); retrolateral (Figs. Yucatan Peninsula 34. yucatecana 214, 406); and Mexico 33(29). Embolus with long retrolateral ramus

- 24 . bicuspidata (e.g.. Figs. 196, 197, 209); western Retrolateral ramus curls to 27(25). prolateral United States and Mexico _ 34 (Figs. 215, 411) 26. morelos Embolus with short retrolateral ramus - Retrolateral ramus points retrolaterallv (e.g.. Figs. 190, 206, 210), or rami not or distally (Figs. 212, 216, 390-394, distinct (e.g.. Figs. 219, 225); widely 416) 28 distributed 38 Retrolateral ramus 28(27). points more distally, 34(33). Retrolateral ramus curled prolaterally about twice as long as prolateral ramus (Figs. 196, 197); embolus very broad (Figs. 212, 390-394); widely distrib- at base 35 uted _ 22. furcata (part) Retrolateral ramus erect or pointing pro- - Retrolateral ramus points more retrolat- laterally (Figs. 193, 209, 212); embolus erally, more than four times longer narrower at base 36 than prolateral ramus (Figs. 216, 416); 35(34). Prolateral ramus of embolus obtuse (Figs. southern Arizona 27. huachuca 196, 259); retrolateral ramus blunt and 29(24). Large white patches on chelicerae at least with bump (Fig. 295); embolus nar- to V2 their length (Figs. 437, 447, 452, rower at base than in tritis (Fig. 196); 478); embolus twists to tip; Arizona, Mexico 8. neoleonis Mexico, and Central America 30 Prolateral ramus acute (Figs. 197, 300); - Chelicerae with at most a small medial- retrolateral ramus sharp; embolus basal patch of scales (e.g., Figs. 258, broader at base (Fig. 197); Arizona 348); embolus varied; widely distrib- 7. tristis uted geographically 33 36(34). Side and forehead bands on carapace re- 30(29). Embolus broad and truncated (Figs. 220, duced or absent (Figs. 156, 365); west- 438); dorsum of abdomen mostly ern United States and Canada brown between side bands (Fig. 164); 19. aeneola (part) - montane _. _ 31. clavator (part) Side and forehead bands on carapace well - Embolus narrower, tapers to tip; dorsum developed (Figs. 158, 276, 388); south- of abdomen may have large white spots western United States, Mexico, and (Fig. 166); varied habitats 31 Central America 37 31(30). Embolus tapers to narrow tip with ter- 37(36). Retrolateral ramus of embolus longer and minal opening (Fig. 226); abdominal diverging from prolateral (Fig. 212); dorsum brown between white side widely distributed 22. furcata (part) - Retrolateral of vertical bands (Fig. 170); montane habitats ramus embolus _..._ _ _ 37. bunites (part) (Fig. 193); embolus narrows abruptly

- near central .._ _ Embolus tip wider, opening subterminal tip; Mexico 4. (Figs. 223, 224); abdominal dorsum _ edrilana (part) Erect of with mixed pale and dark spots (Fig. 38(33). portion embolus arises on retro- 166) as in female; deserts and tropical lateral side (Figs. 422, 428); flange un-

lowlands .._ 32 der tibial apophysis (Figs. 421, 427; arizonensis 32(31). Side and cheek bands fused (Fig. 447); group) 39

in America. 17. P. in North America. 18. P. Maps 16-20. Distributions of Pelegrina species. 16. P. montana Nortti insignis 20. P. balia and P. chalceola in western North chaimona In Mexico and Arizona. 19. P. clemata in western North Amehca. America. 254 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Map 25 y Map 27 Pelegrina Jumping Spiders • Maddison 255

Erect of embolus arises portion centrally; living on Spanish moss ridge under tibial not de- apophysis 36. tillandsiae (part) into veloped flange (Fig. 78) 40 Abdomen brown or spotted above; em- 39(38). Embolus tip sharp (Fig. 422); tibial, bolus usually broad at tip though var- broad apophysis flange and short (Fig. ies; habitat varied 47 421) 28. arizonensis Erect 47(46). portion of embolus very thin (Fig. Embolus blunt tip (Fig. 428); tibial 535); anterior median eyes ringed witli narrow and apoph>sis flange elongate red; first legs yellow fringed vvitli u hite (Fig. 427) 29. helenae (Fig. 176) 45. emmiltus (part) Chelicerae scales 40(38). lacking pale 41 Erect portion of embolus thicker; eyes Chelicerae with small of white patch or ringed with white or brown; legs not yellow scales 46 fringed 48 Embolus narrows 41(40). abruptly just basal to 48(47). Abdomen with paired black spots on opening (Fig. 190); retrolateral ramus brown dorsum; pale markings yellow- is small hook southern (Fig. 259); On- ish; embolus long and rectangular, tario, eastern United States south to leaning slightly retrolaterally (Figs. Central America 1. galathea 205, 349, 350) truncate at tip and with Embolus parallel-sided or widens near retrolateral ramus apparently absent tip; retrolateral ramus is small angle (Fig. 205); Canada and northeastern or apparently absent; Canada and United States 16. insignis mountainous areas of - United States 42 Abdomen lacking distinct black spots; Retrolateral 42(41). ramus of embolus longer pale markings white or yellowish; em- than prolateral and leaning retrolater- bolus shorter (Fig. 219) or if long, then ally (Figs. 208, 209); forehead band straight and with more prominent rami absent 19. aeneola (part) (e.g.. Figs. 192, 206, 207, 222) 49 - Both rami of embolus small; forehead 49(48). Abdomen with large and distinct paired band present or absent 43 white spots as in female (Fig. 166); side 43(42). Embolus swollen near tip (Figs. 204, 344) and cheek bands fused (Fig. 447); che- 15. montana licerae robust (Fig. 447); erect portion - Embolus with sides parallel or slightly of embolus parallel-sided and with two tapering near tip (Figs. 207, 210, 211) subequal rami (Fig. 222); deserts of

- 44 Mexico and Central America 44(43). Forehead band well developed and con- 33. variegata (part) - tacting AMEs (Figs. 152, 359); tegul- Abdomen with only small (Figs. 130, 132, um with prominent prolateral bump 152) or indistinct (Figs. 162, 172) (Fig. 361) 18. clemata (part) paired pale spots, side and cheek bands - Forehead band absent or if present then separate; chelicerae not so robust; em- not contacting AMEs (Figs. 378, 383); bolus varied 50 tegulum with at most small prolateral 50(49). Side bands of carapace and abdomen bump (Figs. 379, 384) 45 weak or absent and abdomen mottled 45(44). Cheliceral fang with flange (Fig. 378); (Figs. 162, 172); embolus small, lack- carapace side bands broad (Fig. 378), ing two distinct rami (Figs. 219, 227) embolus bends slightly (Figs. 210, 379); 51 California and northern Arizona north - Side bands well developed and abdomen to Washington 20. balia not mottled (Figs. 130, 132, 152) 52 - Pale white or erect Cheliceral fang lacking flange (Fig. 383); 51(50). markings gray; por- tion of embolus widens on side bands narrower (Fig. 383); em- gradually Iside as it contacts basal bolus straight (Figs. 207, 211); south- prolateral por- ern Arizona to southern Illinois tion (Fig. 433); tip of embolus rounded 30. verecunda 2 1 . chalceola (Fig. 219) Pale or erect 46(40). Abdomen with striking lineate markings markings orange tan; por- tion of embolus widens on as in female (Fig. 477); embolus ta- abruptly side as it contacts basal pering to sharp tip in ventral view; prolateral por-

Maps 21-27. Distributions of Pelegrina species. 21. P. aeneola in western North America. 22. P. furcata in Mexico and the southwestem United States. 23. P. furcata group members in Mexico and Central America. 24. P. huachuca in Arizona. 25. P. arizonensis and P. helenae in western North America (see Cutler and Jennings, 1985, for additional records). 26. P. verecunda in western North America. 27. P. tillandsiae in southeastern North America. 256 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 Pelecrina Jumping Spiders • Maddison 257

tion so as to make a distinct corner Posterior margin of epigynal flap rovmded, of embolus (Fig. 484); tip pointed ret- not transverse, or if tr;ins\ersc then flaps rolateral to opening (Fig. 227) flat and flush \vith surlacc behind them

_ 38. orestes (part) (Figs. 238, 239, 241-245, 254); legs not 52(50). Retrolateral ramus longer than prolater- distinctK annulate; abdomen w itli white al (Figs. 190, 193) ._ 53 spots smaller, often thinner ami elongate 54 Rami subequal (Figs. 191, 192, 206, 207) (e.g.. Figs. 287, 323, 353) 2 53(52). Embolus sides parallel below opening 2(1). Abdomen with prominent paired black retrolateral ramus (Fig. 190); narrow spots on orange-brown background (Figs. (Fig. 259); widespread 161, 353); epigynal flaps divergent (Fig. 1. galathea (part) 352); epigynal surface rises dramatically - Embolus inflated below opening (Fig. from low area around flaps to high pos- 193); retrolateral ramus wide (Fig. terior margin (Fig. 245); legs and face 277); central Mexico 4. edrilana (part) yellowish; mostly northern 16. insignis 54(52). Abdomen with two central broken lon- Abdomen lacking prominent paired black gitudinal pale stripes in addition to side spots though may have brown or reddish bands as in female (Fig. 269); Carib- patches; epigynal flaps parallel, conver- bean 2. proxima gent, or divergent; epigynal surface in - Abdomen lacking central longitudinal most species with little relief (e.g.. Figs. bands; western North America 55 238, 241); legs and face varied; locality 55(54). Embolus parallel-sided or tapers slightly varied 3 to tip (Figs. 207, 360, 361); basal to 3(2). Epigynum with ridge just behind each flap opening the prolateral side is straight (Figs. 244, 346); posterior notch often (Fig. 207); sagebrush of western Un- rectangular; body large and dark with tied States and Canada very small paired white spots on dark _ 18. clemata (part) abdominal dorsum (Fig. 347); Canada Embolus widens slightly near tip (Figs. and mountains of United States _ 192, 206); just basal to opening on pro- 15. montana lateral side is angle (Figs. 192, 206); Epigynum lacking ridges behind flaps; pos- Arizona and Mexico 56 terior notch triangular; body smaller; ab- 4 56(55). Rami of embolus well separated (Fig. domen varied _ _ 192); side bands have extensions join- 4(3). Abdomen strongly striped longitudinal!) ing between posterior eyes (Fig. 132); yellow and brown (Fig. 477); epigynal of the no denticles on exposed surface of em- flaps pale; living on Spanish moss States 36. tillandsiae bolus (Fig. 192) 3. dithalea southeastern United and brown Rami close together (Fig. 206); side bands Abdomen not striped yellow without extensions; surface of embolus longitudinally, usually spotted; epigynal - 5 with denticles (Fig. 206) 17. chaimona flaps varied _ 5(4). Epigynal surface and flaps very flat (Figs. 24i-243); flaps not much darker than rest Pelecrina of the Key to the Female of epigynum except for narrow rim (Figs. Eastern United States and Canada 322, 327, 332); carapace often w ith shiny and (East of the Mississippi River scales and pale spot above and between Manitoba) anterior median eyes (e.g.. Fig. 143); co- nifer dwellers group) 6 1. Posterior margin of epigynal flap truncated (flavipedes surface and with more relief so as to be transverse, and standing high Epigynal flaps distinctly over surface behind it (Figs. 236, 262); (Figs. 237, 238); flaps usually dis- darker than rest of epigynum (Figs. 286, epigynal flaps convex, parallel; legs 292); carapace lacking shiny scales 8 tinctly annulate (Fig. 131); abdomen Forehead dark above and between AMEs; marked with four pairs of prominent 6(5). behind head often bulbous; legs pale yellow, usu- white spots with small black spots with thin dark lines on _ 1. ally longitudinal them (Figs. 131, 263) galathea

the mannll species group, and 28. P. Maps 28-37 Distributions of species of Pelegrina, Metaphidippus Nagainaincunda^ in and Central America. America. 29. P. P. yucetecana. and P. sandaracina Mexico variegata in Mexico and Central pallidata, Amenca. 32. M. tricolor n Californ a. 31 . M. mannim western North 30 Pbunites and P. orestes in Arizona and western Mexico. United States. /Vl California. 34. M. carmenensis in Mexico and the southwestern 33' M. diplacis in Califomia and Baja 35^ States. 36. P. emmiltus in California and New Mexico. 37. Nagaina incunda in chera in Mexico and the southwestern United Mexico and Central America. 258 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

transverse femora; epigynum \\ ith second curve of so as to be and standing high spermathecal duct wider than in fla- over surface (Figs. 236, 262) 1. galathea - vipedes but no so wide as in exigua (Figs. Legs not distinctly annulate; abdomen with 340, 341); northeastern United States more prominent dark areas on either side bordering Canada and southeastern Can- of smaller pale spots (though pervaga with ada 13. flaviceps pale spots coalesced into single large spot); Forehead with pale spot above and be- epigynal flaps varied, but posterior mar- tween anterior median eyes; head not gin not truncated 3 into bulbous; legs generally lacking longitu- 3(2). Area behind epigynal flaps raised high dinal lines or if persent then wide and mound (Figs. 245, 246); carapace densely scales mostly on anterior legs; epigynum oth- covered with white or yellow (Figs. 161 4 erwise; distribution generally farther 153, ) north or farther south than flaviceps 7 Area behind epigynal flaps more or less flat 7(6). Epigynal flaps parallel (Figs. 241, 322); sec- (Figs. 238, 240); carapace thinly covered ond curve of spermathecal duct narrow with pale scales (e.g.. Fig. 135) 5 and oblique (Figs. 321, 338, 339); cara- 4(3). Scales on carapace white; legs beige and pace narrow; mostly northern (Canada brown; abdomen with large dark patches and northern United States) though found on either side of central paired spots but occasionally on southern mountains lacking strong black spots (Fig. 364), an- 12. flavipedes teriormost pale spots fused into short lon- - Epigynal flaps convergent (Figs. 243, 332); gitudinal bands; epigynal surface behind second curve of spermathecal duct very flaps raised into broad dark shiny round broad and transverse (Figs. 331, 336, 337) mound (Fig. 246); flaps convergent (Figs. carapace broader; mostly southern Unit- 246, 363); usually collected from sage- ed States north to Massachusetts and New brush 18. clemata York 1 4. exigua Scales on carapace yellowish; legs yellow; 8(5). Abdomen marked with large square brown abdomen with paired black spots (Fig. spots on or between paired pale spots 353); epigynal surface behind flaps raised (Figs. 2, 135, 287); epigynal flaps con- gradually but steeply into high mound vergent and fairly flat, short (Figs. 238, along posterior of epigynum (Fig. 245); 286); surface rises quickly behind flaps flaps divergent (Figs. 245, 352); low herbs to broad mound (Fig. 238) 5. proterva in fields and bogs 16. insignis - Abdomen uniformly light brown with small 5(3). Abdomen marked with large square brown white spots (Figs. 137, 293); epigynal flaps spots between small paired pale bands long and fairly convex (Figs. 239, 292); (Figs. 135, 287); epigynal flaps conver- surface rises gradually behind flaps to gent and fairly flat, short and dark (Figs. mound at posterior (Fig. 239) 238, 286); surface rises quickly behind 6. peckhamorum flaps to broad mound (Fig. 238); cara- pace narrow; widespread (5) proterva - Key to the Female Pelegrina of the Abdomen with large central pale spot (Fig. Great Plains (between the 313); epigynal flaps large and flat, fairly Rocky Mountains and the pale (Figs. 240, 312); carapace very wide; Mississippi River)* Kansas to Texas 10. pervaga

1. Epigynal flaps rotated 180° (Fig. 424); ab- domen with lineate strongly markings Key to the Female Pelegrina of the 425) 28. arizonensis (Fig. Pacific Coast of the United States and rotated at most abdom- Epigynal flaps 45°; Western Canada* inal markings not so clearly lineate 2 2(1). Legs distinctly annulate, and abdomen Epigynal flaps rotated 270° so that flaps are marked with four pairs of prominent transverse (Fig. 430); abdomen with li- white spots with small black spots behind neate markings (Figs. 155, 431); com- 29. helenae them (Figs. 131, 263); epigynal flaps con- monly found on sagebrush vex, parallel, posterior margin truncated Epigynal flaps rotated less than 45°; abdo-

* * Not included are some tree-dwelling species whose Includes California, Nevada, Oregon, Washing- ranges reach into the Great Plains: Pelegrina fla- ton, Idaho, British Columbia, and Alberta. Included and vipedes exigua, which occur on conifers in the is Metaphidippus mannii, as well, but not the other north and P. east; peckhamorum, on oaks in the south- mannii group species, which are restricted to the and east; chalceola, in Texas to extreme southern southern part of the area of the key. Not included is Illinois. P. verecunda (see Arizona key). Pelegrina Jumping Spiders • Maddison 259

men with not lineate markings except side of central paired spots (Fig. 364), in clemata, habitat varied 2 occasinally anteriormost pale spots fused into short surface and 2(1). Epigynal flaps very flat (Figs. longitudinal bands; commonly found on with bronze or 241, 256); body shiny cop- sagebrush 18. clemata scales 143, 3 per (Figs. 179) Area behind flaps only moderately raised - more convex and Epigynal flaps epigynal into broad mound (Fig. 238); carapace surface with more relief 244- (Figs. 238, not densely covered with white; abdo- 248); darker than flaps usually distinctly men marked with large square brown rest of without epigynum; body usually spots between paired pale spots (Figs. 2, habitat metallic sheen; varied 4 135, 287); found on various shrubs and 3(2). Orange scales between and beside AMEs trees 5. proterva just above clypeus; body fairly smooth 8(6). Body dark, with very small pale spots on with shiny coppery scales (Fig. 179); usu- abdomen (Fig. 347); white scales be- ally on oaks, holly, Arctostaphijlos, and tween AMEs; epigynum dark; surface other shrubs and trees with leathery leaves rising immediately behind flap to ridge 40. mannii (Fig. 244); collected from waterside - White or dark scales around eyes; carapace shrubs and trees 15. montana - often with pale spot above and between Body mottled beige and tan, with large AMEs (e.g., Fig. 143); body with rougher yellowish spots on abdomen (Fig. 382), appearance; on conifers 12. flavipedes orange scales between AMEs; epigynum surface behind 4(2). Epigynal flap angled where flap bends down pale; rising gradually flaps 20. halia toward opening (Figs. 247, 374, 376); (Fig. 248); juniper-dwelling surface rises immediately behind flap to Key to the Female Pelegrina and mannii broad plateau covering posterior of epi- GROUP OF Arizona* gynum (Fig. 247); carapace thinly cov- ered with white scales that often form 1. Epigynal flaps thin and rotated 90°, lying an inverted T behind the AMEs (Fig. in cavity (Fig. 317); markings gold and 157); abdomen often with anterior me- beige; junipers of southern mountains

dial paired spots coalesced into one large 1 1 . kastoni white spot (Figs. 157, 377); common on Epigynal flaps rotated less than 60° (or, various plants including conifers if rotated 90°, rarely in tristis, then 19. aeneola flaps very broad); markings varied 2 broad and flat 302, Epigynal flaps not angled; surface of epi- 2(1). Epigynal flaps (Figs. surface more or less flat gynum varied; carapace lacking T-shaped 307); epigynal 3 marking on head; abdomen with anterior narrower; sur- medial paired spots separate (Figs. 347, Epigynal flaps epigynal 4 364, 382); habitat varied 5 face varied 3(2). Anterior end of epigynal opening deep, 5(4). Scales on carapace yellowish; legs yellow; with the surface there pale and de- abdomen with paired black spots (Fig. scending deeply under flap; flaps dark 353); epigynal surface behind flaps raised brown; southern Arizona 7. tristis gradually but steeply into high mound Anterior end of epigynal opening shal- of epigynum (Fig. 245); along posterior shallow herbs low, with the surface not so flaps divergent (Figs. 245, 352); low nor descending so deeply as in tristis; in fields and bogs 16. insignis usually light brown; northern Ar- Scales on carapace white, beige, or tan; epi- flaps izona and New Mexico 9. sahinema gynal surface behind flaps either more flat or behind 4(2). Epigvnal surface convex or less flat (Figs. 244, 248) or raised flaps (Figs. 247, 252, 255-257); flaps quickly behind flaps into mound (Figs. often flat 5 238, 246); habitat varied 6 surface concave behind flaps raised into a Epigvnal 6(5). Area behind epigynal flaps convex 14 con- (Figs. 248-250); flaps broad mound (Figs. 238, 246); flaps narrow and flat (Fig. 257), with whitish scales 7 5(4). Epigynal flaps vergent; carapace often and difficult to see or con- transparent Area behind flaps more nearly flat or cave (Figs. 244, 248); flaps divergent dark or covered convergent; carapace * ^ Not included in the key are northern species that with yellowish scales raised occur in Arizona but have been at most rarely 7(6). Area behind epigynal flaps strongly may collected there: montana, flavipedes, in- into round dark shiny mound (Figs. 246, Pelegrina signis, and clemata. Metaphidippus carmenensis is 363); flaps convergent; carapace densely a similar to chera with one known covered with white scales (Fig. 153); ab- species specimen either from Arizona. It is not included in the key. domen with large dark patches on 260 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

(Figs. 525, 526); flaps divergent; com- darker brown spots (Fig. 387), setae mon in desert vegetation though oc- around AMEs darkest dorsally curs at higher elevations 43. chera 21. chalceola (part) - - Epigynal flaps wider and more pig- Carapace covered with white scales; ab- mented, usually more robust than in domen with pale longitudinal side chera, flaps parallel, convergent or di- bands enclosing brown dorsum with vergent; generally found in oak-coni- paired white spots (Fig. 358), setae

fer habitats above 1,200 m elevation 6 around AMEs all white . ._ _ 17. 6(5). Epigynal flaps narrow, flat and mostly chaimona (part) parallel except for sharp bend inward 14(4). Epigynal flaps dark, long, narrow, and surface near posterior end (Figs. 255, 481); convex (Fig. 418); epigynal body vellow, sometimes with paired with strong relief consisting of raised dark spots on abdomen almost as in bumps just medial to each flap, a con- behind to insignis (Fig. 171) 37. bunites cavity flaps rising posterior - first curve of Epigynal flaps without sharp posterior edge (Fig. 418); epigynal bend; if abdomen yellow then lacking ducts broad and long (Fig. 417) large paired dark spots 7 _ 27. huachuca not so nor is 7(6). Abdomen very pale, yellowish, with Epigynal flaps long, epigynal markings consisting of little more than surface so strongly sculptured; first of small dark speckles (Figs. 436, 487, 502) 8 curve ducts narrower 15 - Abdomen more darkly marked with 15(14). Epigynal flaps strongly convex (Figs. 249, or 250); end rounded and stand- brown grav' (Figs. 275, 358, 377, posterior 387) _ _ 10 ing high abo\e surface (Figs. 249, 250); second curve of duct broad 8(7). Epigynal flaps divergent and narrow (Fig. (Figs. 397, 501) 40. mannii (part) 400) _ 22. furcafa - Epig\nal flaps parallel or convergent Epigynal flaps less convex (Figs. 248, 357, not (Figs. 435, 486) _ 9 381, 386); posterior end standing surface 9(8). Epigynal flaps pale (Fig. 486) 38. orestes high above (Fig. 248); second - of Epigynal flaps small and dark (Fig. 435) curve duct narrower (Figs. 356, 380, _ 30. verecunda 385) _ 16 with round white 10(7). Epigynal flap angled about midway along 16(15). Abdomen marked large its length where flap bends down to- spots (Fig. 382); carapace wide; epi- narrow and ward opening (Fig. 247); surface rises gynal flaps pale (Fig. 381) _ 20. balia immediately behind flap to broad pla- - teau covering posterior of epigynum Abdomen with small white spots if any (Fig. 247); carapace thinly covered (Figs. 358, 387); carapace varied; epi- with white scales that often form an gynal flaps broader and shorter (Figs. 1 7 inverted T behind the AMEs (Fig. 157); 357, 386) abdomen often with anterior medial 17(16). Carapace covered with brown reflective paired spots coalesced into one large scales; abdomen brown with large white spot (Fig. 157) 19. aeneola paired darker brown spots (Fig. 387); - setae anterior Epigynal flaps not angled so abruptly in around median eyes middle; epigynal surface varied; car- darkest dorsally 21. chalceola (part) - apace lacking T-shaped marking on Carapace covered with white scales, ab- head; abdomen with paired spots sep- domen with pale longitudinal side arate 11 bands enclosing brown dorsum with setae 11(10). Epigynal flaps divergent and narrow (Fig. paired white spots (Fig. 358); 501) _ 40. mannii (part) around anterior median eyes all white - Epigynal flaps parallel or convergent, not _ 17. chaimona (part) so narrow (Figs. 274, 357, 386) 12 Key to the Pelegrina and Nacaina 12(11). Abdomen marked much as in galathea, females of mexico and with four pairs of prominent white Central America* spots with small black spots behind them (Figs. 133, 275); epigynal flaps 1. Body and legs mostly yellow (Fig. 163, short, fairly flat, and parallel (Fig. 274) 173, 175, 436, 461, 487, 492), with small 3. dithalea Abdomen dark areas more prominent than paired white spots (Figs. 358, 387); Females of the Metaphidippus mannii groups epigynal flaps varied 13 are not included. These can usually be distinguished 13(12). Carapace covered with reflective scales; from Pelegrina females by their weaker epigynal with abdomen brown large paired flaps, which descend into the openings posteriorly. Pelegrina Jumping Spiders • Maddison 261

dark markings if any; epigvnum and dominal markings not so clearly li- flaps mostly flat (Figs. 252,' 253, 255) 2 neate 10 - Bod> and legs well marked with brown 10(9). Epigynal flaps dark, wide, flat, and and gray (e.g.. Figs. 298, 318, 409, 414, strongly convergent (Fig. 297); mon- 425); epigynal surface flat or more or tane 8. neoleonis - less concave (e.g.. Fig. 250) 9 Epigynal flaps not so dark and wide; hab- 2(1). First femur, patella, and/or tibia with itat varied 1 1 small subterminal dark transverse bar 11(10). Epigynal flaps rotated 90° and in pits (e.g., Feckham and Peckham, 1896: fig. (Fig. 317); body yellowish (Fig. 141); 10); clypeus covered with yellow scales northern Mexico, on junipers

except for barren patch beneath AMEs, 1 1 . kastoni beneath which on chelicera is dark line; Epigynal flaps rotated less than 60°; body epigynal flaps weak (Fig. 491); dis- varied; distribution varied 12 turbed lowland habitats 12(11). Abdomen with peculiar transverse 39. Nagaina incunda markings (Figs. 169, 456); fourth pair - Legs uniform in color or if annulate, with of spots in particular a transverse stripe; dark annulae more extensive; clypeus legs strongly annulate (Fig. 169); face densely covered with pale scales even thinly covered with pale scales; epi- below AMEs; habitat varied 3 gynal flaps pale and convergent (Fig. 3(2). Epigynal surface more or less flat except 455); Yucatan Peninsula 34. yucatecana - for longitudinal ridge between flaps Abdomen without such transverse mark- (Fig. 253); flaps convergent, narrow, ings; fourth pair of spots not a trans-

only slightly convex (Figs. 253, 450) .. verse stripe; legs, face and epigynal 33. variegata (part) flaps varied 13 - Epigynal surface lacking central ridge 13(12). Epigynal flaps with abrupt bend near (Figs. 252, 255); flaps varied 4 posterior end (Figs. 255, 481); epigyn-

4(3). Body and legs uniformly orange-yellow al flaps and surface more or less flat .. except sometimes for discrete small 37. bunites (part) - dark spots on abdomen; epigynum Epigynal flaps without abrupt bend near transparent so that spermathecae eas- posterior end; epigynal surface varied 14 ily visible without dissection (Figs. 460, 14(13). Epigynal surface more or less flat except 463); flaps convergent; southern Mex- for longitudinal ridge between flaps ico and Central America (Fig. 253); flaps convergent, narrow, 35. sandaracina only slightly convex; abdomen marked - Body and legs pale yellowish beige, not with large white spots (Figs. 167, 451) so orange; epigynum varied 5 33. variegata (part) 5(4). Abdomen uniformly yellowish, with Epigynal surface usually rises to poste- small discrete dark speckles only (Figs. rior edge; if flat then lacking longi- 6 tudinal 436, 487 ) ridge; flaps varied; markings - Abdomen mostly yellow but any dark varied 15 markings are larger spots and patches 15(14). Epigynal flaps strongly convex (e.g.. Figs. or con- (e.g.. Figs. 171, 409, 446) 7 236, 249, 250), parallel slightly 6(5). Epigynal flaps pale, transparent (Fig. vergent 16 - 486), convergent _ 38. orestes Epigynal flaps flat or only slightly convex Epigynal flaps dark (Fig. 435), conver- (similar to those in Figs. 247, 252); may gent to divergent 30. verecunda be strongly convergent 20 7(5). Epigynal flaps strongly convex (Fig. 408); 16(15). Epigynal surface concave behind flaps epigynum concave behind flaps (Figs. 249, 250), rising gradually to 25. ochracea posterior margin (Jurcata group) 17 - Epigynal flaps flat (Figs. 255, 482, 445); Epigynal surface rises quickly behind epigynal surface more or less flat 8 flaps to mound covering most of pos- 8(7). Epigynal flaps narrow, with abrupt bend terior (Fig, 236) 19 near posterior end (Figs. 255, 481); Ar- 17(16). Epigynal flaps fairly short, pale (Fig. 408); izona to Oaxaca 37. bunites (part) southern Mexico and Guatemala - Epigynal flaps wider, convergent, but 25. ochracea without abrupt bend (Fig. 445); Chia- Epigynal flaps generally longer, dark pas to 32. pallidata (part) (Figs. 298, 413) 18 9(1). Epigynal flaps rotated 180° (Fig. 424); 18(17). First curve of duct narrow, second curve abdomen with strong lineate markings very broad (Figs. 397, 400); abdominal (Fig. 425) 28. arizonensis markings shiny, pale spots generally Epigynal flaps rotated at most 90°; ab- small (Figs. 396, 398, 402) 22. furcata 262 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

- First curve of duct wide, second curve Even if no additional species are discov- narrow 412); on (Fig. strong pale spots ered in Arizona, there is the danger that dark abdominal dorsum 414) (Fig. males and females of some of the known 26. morelos have been mismatched. In Mexico 19(16). Epigynal flaps long and convergent, not species truncate posteriorly (Figs. 4, 280); lon- and Central America, the situation is worse, dark bands on abdomen gitudinal where there are probably several species 4. edrilana prominent (Fig. 281) for - that will remain undescribed some time Epigynal flaps shorter, parallel, truncate to come. there are known some posteriorly (Figs. 236, 262); abdomen Already marked with prominent white spots, female Pelegrina from southern and cen- without prominent dark bands (Figs. tral Mexico that apparently represent spe- 131, 263) 1. galathea (part) cies not described here. I shall not give 20(15). Epigynal flaps convergent, higher me- names to them here so as to avoid dially and tilted down laterally (Fig. making based difficult to 440); epigynal surface high between more species names on and behind teardrops, lower lateral to determine females and because with ad- this (Fig. 440) 31. clavator we discover that - equate collecting may Epigynal flaps not so tilted; epigynal sur- they are females of already-described face flat or only slightly higher me- males. I do, of some dially than laterally 21 however, give figures 21(20). Epigynal flaps truncate posteriorly (Fig. of them (Figs. 464-471). Figures 464-466 above surface at 236), high posterior show a single female from Neriaco, Mexico end 1. galathea (part) which an - (state unknown), may represent Epigynal flaps not truncate posteriorly extreme southern form of P. chalceola. nor so high above surface 22 22(21). Abdomen marked much as in galathea, Figures 467 and 468 show a form from with four pairs of prominent white Guerrero, Jalisco, and Michoacan that may with small black behind spots spots be a southern form of P. dithalea. Figures them (Figs. 133, 275); epigynal flaps 469-471 show a form occurring in collec- fairly flat and parallel (Fig. 274) 3. dithalea tions from Durango. Abdomen with smaller white spots (Figs. The descriptions follow a more or less 358, 446); epigynal flaps usually con- consistent format except that occasionally vergent 357, 445) 23 (Figs. a feature is noted in a few species that is 23(22). Epigynum very flat (as in verecunda, Fig. not noted in others: for instance, 252); dark band along inner margin of any are noted under P. epigynal opening very wide (Fig. 444) strongly annulate legs 32. pallidata (part) yucatecana, but leg annulation is usually Epigynum with some relief; dark band not even mentioned, and in P. balia the along inner margin of epigynal open- flange on the cheliceral fang is noted but ing narrow (Fig. 356) 17. chaimona* the fang is ignored in most other descrip- DESCRIPTIONS OF THE SPECIES OF tions. In the case of leg annulation and PELEGRINA male abdominal markings, the species should be assumed to be characterized by The Pelegrina species of Canada and the usual condition annu- the northern and eastern United States can Pelegrina (legs late, but and male ab- be considered reasonably well known, but fairly indistinctly, domen brown above, with at most small the same cannot be said for the species of white and white side Arizona, Mexico and Central America. In spots, ringed by bands) unless otherwise mentioned. In the Arizona are many species, some poorly col- case of the other such as the lected such as P. huachuca, P. chaimona, characters, in balia, the distribution of the fea- P. tristis, P. chalceola, and P. dithalea. flange ture in all species is not fully known. Such a character is described to aid in separating the from similar that are * A number of unmatched females from Mexico species species known to lack it this chalceola may be P. chaimona or a species easily confused with (in example, it. lacks the flange). Pelegrina Jumping Spiders • Maddison 263

" Information on the labels of type ma- "Dendryphantes ornatus Bks type, "Ithaca, N. Y." and "Nathan Banks Coll.," examined. terial is cited, and, where possible, the au- Dendryphantes hondurensis: —G. & E. Peckham, thor of handwritten labels is identified. in 1896, part: 48, pi. 4, fig. 4a, 9. Type material in Banks, Chamberlin, Kaston, and Levi types MCZ 22 from Belize labeled "449 Dendryphantes still have with them the author's original hondurensis Peck., Type, British 2 1423, G. W. & E. G. Peckham C:oll." labels, handwritten except those of Kaston, (in Bryant's haiuT- writing) which both belong to the genus Gastromi- whose typewriter was distinctive. F. Pick- cans, and \$ 29 labeled "461 Dendryphantes hon- and some of the Peck- ard-Cambridge's durensis Peck., Guatemala, G. W. & E. G. Peckham hams' types no longer have their original Coll." (in Bryant's handwriting), of which 16 19 labels. F. P. -Cambridge's labels have been are P. galathea and 19 is in the genus Messua, examined. One Gastromicans 9 from Belize is here replaced by labels handwritten in pencil, designated LECTOTYPE of D. hondurensis, and Pocock or (Levi, perhaps by Browning thus D. hondurensis is not properly a synonym of Some of the personal communication). P. galathea. — Peckhams' labels were rewritten by Bry- Metaphidipptis capitatus: F. P. -Cambridge, 1901: 272; Bonnet, 1957: in , but most labels of Pelegrina types are 2810, part. Metaphidippus digitatus F. P. -Cambridge, 1901: 269, apparently original. Some are in George pi. 24, figs. 12, 12a-c, S. Type material in BMNH but most are in a Peckham's handwriting, 15 and fragments of two other S labeled "Dendry- handwriting that is probably that of Eliz- phantes digitatus, sp. n. Type 5, Guatemala (Sarg.)" and 26 labeled n. abeth Peckham, for it occurs in other orig- "Dendryphantes digitatus, sp. S's, Mexico H. S", examined. NEW SYN- inal labels in the Peckham Collection and (Teapa) ONYMY. in some of Peckham's George correspon- Beata digitata:—Simon, 1903: 841. Roewer, 1954: dence to Henshaw. 1007. Bonnet, 1955: 873. Dendryphantes capitatus: —G. & E. Peckham, 1909:

469, pi. 38, fig. 5, possibly also pi. 36, figs. 4, 4a, 9. 1 . Pelegrina galathea Metaphidippus galathea: —Chamberlin and Ivie, (Walckenaer, 1837) 1944: 203. Kaston, 1973: 117, figs. 47-50, <59. combination new Dendryphantes galathea: —Roewer, 1954: 1203. Figures 5, 10, 11, 13, 35, 78, 125, 130, 131, 190, 236, 258-263; Map 1 Notes on Synonymy. (1) I interpret Bosc's ambiguous figure (photograph of Attus galathea Walckenaer, 1805: 23 (cites Bosc's MS plates in MNHN Paris seen) as the species figure, pi. 1, fig. 4, 9) (nomen nudum). we call P. following recent us- Attus galathea Walckenaer, 1837: 456, sp. 100. Type galathea, shows a material lost or destroyed. Walckenaer (1837) cited age. Abbott's figure 405 probably and also Abbott's 405 Bosc's MS pi. 1, fig. 4, fig. 9 proterva. (2) Walckenaer's Attus atten- but as Walckenaer refered to Bosc's (9), (1805) only tus and Attus furtivus might also refer to figure, this is to be taken as figure of type. Insofar this species. (3) The epigynum figured by as A. galathea is such a common and well-known the Peckhams for D. hondurensis was that species, and Bosc's ambiguous figure could be in- this terpreted as another species, a NEOTYPE is here of a P. galathea female, but despite designated, 1<3 in MCZ with label "NORTH CAR- the female Gastromicans from Belize was OLINA: 24-31 1943, Brim- Raleigh, garden, May chosen as lectotype because of the name ley." British Honduras 9. hondurensis (suggesting Attus nuhilis Hentz, 1846: 358, pi. 21, fig. 15, Type intended as the label material lost or destroyed. was type locality), is not Euophrys leucophaea C. L. Koch, 1846: 216, fig. 1261, "Type," and because their figure 5 in ZMB with S. Holotvpe 1(5 from Pennsylvania of P. galathea. Their description appears labels "E. ZMB 1794," "1794," "Hol- Leucophaea to apply to the mixture of species in the otypus," examined. Was dried; now rehydrated. NEW SYNONYMY. vials. (4) Kaston used the name nubilis for 9. identifications of material Icius crassiventer Keyserling, 1884: 503, fig. 11, his numerous Holotype in MCZ 19 with labels "18 Icius crassi- around 1940. venter 2 Massachusetts." and "18.", exam- Keys., Diagnosis. A widespread species, for- ined. NEW SYNONYMY. merly confused with others in eastern Dendryphantes ornatus Banks, 1892: 75, pi. 4, fig. in 12 with labels North America, from which it is distin- 29a, pi. 5, fig. 29, 2. Holotype MCZ 264 Bulletin Museum of Comparative Zoology, Vol. 154, No, 4

guished by the embolus shape of males and not so much as in proterva; second curve speckled abdominal dorsum, annulate legs, proceeds medially. Markings (Figs. 131, and convex epigynal flaps of females. Car- 263): Carapace covered above with white apace wider than in most eastern species. to gray scales. Clypeus relatively thinly Similar especially to the Caribbean prox- covered with white setae. Abdominal ima and southwestern dithalea. Can be markings dominated by central pale spots separated from proterva by (males) the each of which is shadowed by dark behind. narrower embolus with smaller hook, Measurements: Body length 4.0(4.6)5.7 darker face, and broader carapace and by mm; carapace length 1.7(1.8)1.9 mm, = (females) the abdominal markings and width/length 0.78(0.82)0.82; n 5$ from convex epigynal flaps. Georgia, Alabama, and Michigan. = Male. Palpus (Figs. 190, 259, 260): Em- Chromosomes. 2n3 26 acrocentrics -\- bolus rectangular, narrowing abruptly just XXO (13 with full count plus 13 with only basal to opening, with small pointed, XXO observed, Toronto, Ontario). curving hook at retrolateral tip (Figs. 190, Courtship (73 observed from seven lo- 259). Markings (Figs. 130, 258): Forehead cations: Rowan Co., Kentucky; San Jacin- band often well developed, with each to, Gonzales, and Hidalgo Co., Texas; San branch forked and extending back to pos- Luis Potosi: 99°42'W, 22°28'N; near Tux- terior eyes (Fig. 258). Cheek band weak. pan, Veracruz; and north of Ciudad Ca- = Clypeus brown, lacking central white spot margo. Chihuahua). Raisedspread (n 9, - on clypeus, with hairs overhanging chelic- 53). Crouch (Fig. 125; n 14, 63): Body = = erae dark except sometimes a few white low (n 5, 33) and horizontal (n 14, 63). hairs medially. White forehead band con- First legs held forward and horizontal (n = = tacts AMEs dorsally 10:30-12:30. Chelic- 12, 53), or slightly raised (n 1), or raised = erae lacking pale scales except in some about 45° (n 1); bowed and touching or = southern males. Femur of palpus only almost touching at tips (n 9, 43), or = slightly paler than more distal segments, straight forward (n 4, 13), or slightly cymbium dark brown and lacking white spread, though more parallel and lower as = = scales except in some southern males. Fem- 3 gets closer (n 1). First legs flicker (n = ora of second, third, and fourth legs often 12, 63) on each series (n 4, 23) up and = more uniformly dark than in proterva, light down (n 4, 23) and alternately back and = brown base graduating to dark brown apex, forth at tips (n 1), vigorously (ca. 5 c/s) = = though in some S3, especially in south, base (n 1) but at low amplitude (n 5, 33). = abruptly pale. Abdomen shows trace of Palpi held down (n 9, 53), either resting = white spot pattern of 92. Measurements: on first leg femora (n 1), tucked beside = Body length 3.0(3.4-3.8)4.0 mm; carapace chelicerae (n 3, 13) or over chelicerae = = length 1.4(1.6-1.8)2.0 mm, width/length (n 2, 13) and pointing inward (n 4, = = 0.77(0.79)8.1; n 63 from Michigan and 23). Palpi waved (n 10, 43) up and down = = Georgia. (n 3, 13) on each series (n 6, 23) vig- Female. Epigynum (Figs. 261, 262): orously (ca. 5 c/s) but at low amplitude (n = Flaps convex (Fig. 236), inner edges often 1). Repertoires: 13 raisedspread only; 23 parallel and close together, back edge of- crouch only; 43 raisedspread and crouch. ten perpendicular to body axis and stand- Distribution (Map 1). Eastern North ing much higher than surface immediately American north to southern Ontario, west behind it (Fig. 236). Surface rises fairly to the Rockv Mountains, south to Florida quickly behind flaps so that posterior sur- and Costa Rica. face is mostly raised, unlike the more con- Records. Manv in MCZ and cave surface of and specimens, especially proxima peckhamo- AMNH, from; CANADA: ONTARIO: Burlington, rum though not so uniformly high as in Hamilton, Port Credit, Windsor. UNITED STATES proterva. First curve of duct broad, but (county records): NEW HAMPSHIRE: Cheshire, Pelegrina Jumping Spiders • Maddison 265

Strafford; VERMONT: Hillsborough, Windham; (1991) examined segregation of isozyme MASSACHUSETTS: Barnstable, Dukes, Essex, Mid- markers in P. galathea. dlesex, Nantucket, Norfolk, Suffolk; RHODE IS- LAND: Newport; CONNECTICUT: Fairfield, Hart- ford, Litchfield, Middlesex, New Haven, Tolland; 2. Pelegrina proxima NEW YORK: Dutchess, Nassau, Suffolk, Tompkins, & E. Wyoming; NEW JERSEY: Bergen, Cape May, (G. Peckham, 1901) Gloucester, Hunterdon, Middlesex, Morris; PENN- new combination SYL\'ANIA: Adams, Berks, Bucks, Erie, Montgom- Figures 191, 237, 264-269; Map 2 ery; OHIO: Ashtabula, Champaign; DELAWARE: Sussex; MARYLAND: Baltimore, Montgomery, Dendryphantes proxima G. & E. Peckham, 1901b Washington; DISTRICT OF COLUMBIA: Washing- (January; see G. & E. Peckham, 1909: 457): 327, ton; WEST VIRGINIA: Mercer; VIRGINIA: Alle- pi. 28, figs. 3, 3a, S9. Types in MCZ 15 12 2imm. gheny, Botetourt, Fairfax, Suffolk, Surry, Portsmouth, "Dendryphantes proxima Pkm, 1901. Cuba Type. Richmond, Washington; KENTUCKY: Rowan; S 9." and "G. W. Peckham Coll." (label is original; TENNESSEE: Benton, Unicoi; NORTH CAROLI- handwritten, probably by Elizabeth Peckham), ex- NA: Avery, Buncombe, Camden, Craven, Durham, amined. The type vial also contains one palpus of Johnston, Macon, Mecklenburg, Nash, New Hanover, another species, perhaps Metaphidippus mannii, Pender, Transylvania, Wake, Washington, Yancey; which is probably misplaced. SOUTH CAROLINA: Oconee, Orangeburg; GEOR- Dendryphantes prudens G. & E. Peckham, 1901a GIA: Chattahoochee, Clarke, Cobb, Glynn, Thomas, (May): 15, pi. 4, figs. 13, 13a, 13b, S. Types in MCZ Ware; FLORIDA: Alachua, Escambia, Hillsborough, 2S 12 with labels "1131 Dendryphantes prudens Indian River, Jefferson, Leon, Madison, Orange, Palm Peckhams, B.0155, , Kingston 31423, 24123" Beach, Pinellas, Polk, Putnam; ALABAMA: Baldwin, (in George Beckham's handwriting) and "B.0155," Colb, Coosa, Dallas, Mobile, Tallapoosa; MISSISSIP- examined. Roewer, 1954: 1199. PI: Harrison, Rankin; LOUISIANA: Baton Rouge, Dendryphantes (Metaphidippus) proximus: —Pe- Caddo, Jefferson, St. Charles; MICHIGAN: Calhoun, trunkevitch, 1911: 640.

Gratiot, Hillsdale, Jackson, Livingston, Micosta, Mid- Pelegrina geniculata Franganillo, 1930: 45, fig. 17, land, Montcalm, Muskegon, Newaygo, Oakland, 2. Types from Sierra Maestra, Cuba, in lESC, orig- Washtenaw, Wayne; INDIANA: Clay, Howard, Mar- inally labeled only by a numerical code but 12 here ion, Starke; ILLINOIS: Adams, Champaign, Peoria; designated as lectotype with labels "PF 548," "Pe- MISSOURI: Berry, Boone, Jackson, Nevada, St. legrina geniculata Franganillo, Lectotype, desig. Charles, St. Louis, Vernon; ARKANSAS: Carroll, W. Maddison 1990" (see comments regarding the Conway, Hempstead, Lincoln, Washington; KAN- generic name Pelegrina, earlier). 42 here desig- SAS: Bourbon, Cherokee, Jefferson, Riley; OKLA- nated as paralectotypes, 3 deposited in lESC, and HOMA: Cleveland, Kiowa, Payne; TEXAS: Aransas, 1 deposited in MCZ. Franganillo, 1936: 138,fig. 76. Bexar, Brazos, Cameron, Comanche, Dallas, Denton, NEW SYNONYMY. Galveston, Grayson, Harris, Hidalgo, Jim Wells, Metaphidippus proximus: —Bryant, 1940: 501 (= Karnes, Kleberg, Leon, Llano, McLennan, Nueces, prudens). Bonnet, 1957:— 2817. Tavlor, San Jacinto, San Patricio, Wichita; COLO- Metaphidippus prudens: Bryant, 1943: 496, figs. 56, RADO: Boulder, Denver, Sedgwick; NEW MEXICO: 57, 63, .52. Bryant, 1950: 189. Bonnet, 1957: 2817. Dona Anna, Rio Arriba, MEXICO: TAMAULIPAS: Dendryphantes proximus: — Roewer, 1954: 1199. Santa Gracia, Reynosa; SAN LUIS POTOSI: near Ciu- Notes on dad del Maiz; NUEVO LEON: Villa de Santiago; Synonymy. Bryant synony- COAHUILA: Gloria; CHIHUAHUA: 21 km N of mized prudens with proxima in 1940 but Ciudad Camargo, Delicias; VERACRUZ: just S of then, in 1943 and 1950, used the name Tuxpan, Fortin; CHIAPAS: Tuxtla Gutierrez. GUA- prudens without explanation. The synon- TEMALA: Amatitlan, Capetillo. COSTA RICA: Chi- of is based on ral Paraiso, Cartago. : Grasmere. ymy Pelegrina geniculata Franganillo's description and an exami- Natural History. In eastern North nation of all surviving specimens of the America, this species is generally found in Franganillo collection, kindly sent to me sunlit places such as oldfields, in contrast from the lESC by Luis F. de Armas via to P. proterva, which is generally more of Herbert Levi and Charles Dondale. The a forest dweller. In Chihuahua, P. galathea collection consists of 26 numbered vials lives in riparian vegetation. Horner (1972) containing at least 17 species (Table 4). has investigated the bionomics and im- The number and diversity of species rep- portance of P. galathea in biological con- resented is approximately what might be trol in sorghum. Steiner and Greenstone expected from Franganillo's papers; thus. 266 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Table 4. Franganillo's collections of salticids. The identifications are by me (with vial number; E.G., PF 548, IN parentheses).

Agobardus cubensis (Franganillo) sensu Bryant: 1 penultimale 6 29 (PF 546) Agobardus sp.: 19 (PF 551) Corijthalia cf. arcuata sensu Bryant: 12 (PF 539) Corythalia cf. squamata Bryant: 2$ (PF 540) 2$ 1 innm. 26 12 Corythalia sp. (not C. arcuata sensu Bryant): 49, (PF 539), (PF 540) . \$ (PF 543) Hentzia cf. tibialis: IS 19 1 imm. Hentzia sp.: 12 (PF 544), 12 (PF 544-2), (PF 562), (PF 564) 4 imm. Lijssomanes antillanus. 22, 2 imm. (PF 535), 39 (PF 536), 5$ 19 (PF 542), 19, (PF 543) Lyssomanes sp., 1 imm. (PF 532) 12 \$ Menemerus bivittatus. 32, 1 imm. (PF 541), 1 penultimate 9 (PF 567), 1 penultimate S, (PF 568), (PF 569) Metacyrba taeniola. 22 (PF 575) Metacyrba sp., 12 (PF 575) Pelegrina proxima, 52 (PF 548), 1

Nilakantha sp.: 12 (PF 544-2) . 15, 1 penultimate $ (PF 571) Platycryptus sp., U (PF 560) Plexippus paykulli: 26 (PF 534), 32 (PF 535), 12 (PF 566) Synemosyna smithii: 12 (PF 550).

the collection may remain more or less among the species in Franganillo's col- complete. The collection lacks labels in- lection. Table 5 lists species in his col- dicating locality or species (Alayon, 1982); lection and their spination. The spina- thus, it is possible that we will never iden- tion of 3-3 on the tibia and 2-2 on the tify the type specimens of Pelegrina gen- metatarsus described for Pelegrina iculata with complete certainty. However, geniculata narrows down the species to I will argue that Pelegrina geniculata is a a dendryphantine, and the fourth tibia junior synonym of Dendryphantes proxi- spination matches P. proxima exactly. mus and that, in particular, the types are Table 5 also lists a few other Cuban in vial 548. the females PF Franganillo's salticids not in Franganillo's collection. is rather detailed in some re- description Among species known from Cuba but and a of the was spects, figure epigynum not listed in the table, Bryant's (1940) The described of the provided. size, shape descriptions indicate that none have the carapace, nature of the clypeus, chelicer- leg spination described for Pelegrina, ae, sternum, eyes, and legs all proxima. fig except Sidusa turquinensis, Icius wick- The placement of Pelegrina in the Uni- hami, Phidippus spp., and Neon nigri- dentati implies a single simple tooth on the ceps, which can be ruled out as Pele- retromargin of the cheliceral fang furrow, grina geniculata on other grounds. consistent with proxima and inconsistent Though spination can sometimes be un- with some such as Hentzia. The genera reliable (Maddison, 1987), spination dif- of con- description Pelegrina geniculata ferences such as those seen between tains nothing that would rule out proxima, dendryphantines and the other subfam- and several features that in particular point ilies listed in Table 5 are reasonably to this These are as follows: species. reliable. 1. Leg spination: Franganillo's descrip- 2. Epigynum: Franganillo's (1930) epi- tion of the spination of the first and gynal figure shows two dark teardrop- fourth legs can apply only to P. proxima shaped objects and posterior notch. No Pelegrina Jumping Spiders • Maddison 267

Table 5. Leg spination of Cuban salticids. Specimens marked by asterisk (*) are from Franganiro's COLLECTION. ALL SPECIMENS IN FRANGANILLO's C;0LLECTI0N ARE INCLUDED EXCEPT LYSSOMANES AND Synemosyna, which are c;learly not Pelegrina by the description. The spination pattern of Pelegrina genicvlata is taken from Franganillo's (1939) description. => 268 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

several adults, all females of P. proxi- Distribution (Map 2). Known from the ma. The description notes the material larger Caribbean islands. being six females; the vial contains five Records. BAHAMAS: Grand Bahama Island, Free- females. No other vial in the collection port (19, AMNH); Rum Cay, near Port Nelson (25, so females without ac- contains many AMNH). CUBA: Havana (many <39, MCZ); Havana: companying males or immatures. Santiago de las Vegas (66 52, MCZ); Marianao Habana (29, AMNH); Soledad, Cienfuegos (55 49, MCZ, AMNH); Oriente: de Cuba (19, MCZ); Trin- This evidence taken together indicates that Santiago idad Mtns., Hanabanillo Falls {IS, MCZ); Holquin the description of geniculata Pelegrina (.59, MCZ); Banes (19, MCZ); 7 km N of Vinales {26 to P. in to applies proxima and particular 19, AMNH); Vega Alta, Santa Clara (43, AMNH); San the females in vial PF 548. Accordingly, Vicente, Pinar del Rio (43 22, AMNH). JAMAICA: Christiana Claremont one of these females has been designated (19, AMNH); (13, AMNH); Spanish Town (29. MCZ); St. Andrew: Mona (53 19, as a lectotype of Pelegrina geniculata. MCZ); St. Andrew: (19, MCZ); St. Ann, 1.6 The known Caribbean Liquanea Diagnosis. only km E of Moneague (12, MCZ). DOMINICAN RE- Pelegrina, differing from the similar gal- PUBLIC: S. O. de las Matas (13, MCZ); La Vega (19, athea in having more lineate abdominal MCZ); Ciudad Trujillo (19, MCZ). HAITI: Diquini (13, MCZ); Enery, Bata (13, AMNH); hills nr. Port- markings and in details of genitalia. au-Prince (13, MCZ); Quest (13 19, MCZ). Male. Palpus (Figs. 191, 265, 266): Embolus rectangular, not narrowing so 3. dithalea new abruptly near the opening as in galathea; Pelegrina species 132, 133, 192, 270-275; 3 retrolateral ramus an angle not prolonged Figures Map into a hook (Figs. 191, 265). Markings (Fig. Holotype male and paratype female in MCZ, with 264): Cheek band very weak. Clypeus label "ARIZONA: Santa Cruz Co., Sycamore Can- brown, with hairs overhanging chelicerae yon, ca. 9 mi [14 km] W of Peiia Blanca Lake, W of ca. 4000 ft. el. [1,220 m], 19 Jun 1985 dark with a few white medially. White Nogales, W. Maddison 85-060, sweeping in canyon where forehead band contacts AMEs dorsally stream flowing." 10:30 to 12:30. Chelicerae with small me- dial patch of pale scales. Cymbium usually Etymology. An arbitrary combination lacking white scales. Abdomen often show- of letters, to be treated as a noun in ap- ing a trace of the longitudinal white bands position. of females. Measurements: Body length Diagnosis. Similar in markings to gal- 2.8(3.0)4.2 mm; carapace length 1.5(1.6)2.1 athea, from which it differs by the em- = mm, width/length 0.78(0.79)0.80; n 53 bolus that lacks the hooklike retrolateral from Havana, Cuba. ramus and that widens toward the tip. Em- Female. Epigynum (Figs. 237, 267, 268): bolus resembles that of chaimona, but the Flaps long, fairly convex, dark, not trun- rami are farther apart (Figs. 192, 206). cated behind as in galathea. Surface be- Male. Palpus (Figs. 192, 271, 272): Em- hind flaps more or less concave, rising bolus widens slightly from near base to gradually so that posterior mound restrict- near tip. Rami subequal, though retrolat- ed to guide area or absent (Fig. 237). First eral is more prominent. Markings (Figs. curve of duct narrow; second curve pro- 132, 270): On carapace, white bars from ceeds medially. Markings (Fig. 269): side bands to fovea usually strong and fused Carapace covered above thinly with white into inverted V mark. Cheek band weak. scales. Clypeus densely covered with white Clypeus brown, hairs overhanging chelic- scales. Abdominal white spots arranged into erae white centrally, dark laterally. White two median longitudinal white bands on forehead band contacts AMEs dorsally brown background. Measurements: Body 10:30-12:30. Chelicerae with small medial length 3.5(3.7)4.8 mm; carapace length patch of white scales. Cymbium with few 1.5(1.6)1.9 mm, width/length 0.76(0.76) white scales. Legs fairly distinctly annu- 0.79; n = 59 from Havana, Cuba. late. Abdomen shows traces of white spots Pelecrina Jumping Spiders • Maddison 269

of female. Measurements: Body length in June. At Kitt Peak, beating oaks and 3.6(4.0)4.2 mm; carapace length 1.9(2.0)2.1 other shrubs and trees. mm, width/length 0.77(0.78)0.81; n = 53 from Arizona. Sycamore Canyon, 4. Pelegrina edrilana new Female. species Epigynum (Figs. 273, 274): 4 Figures 4, 193, 276-281 ; Map Flaps slightly convex; posterior edge not standing so high above surface behind them Holotype male with one immature in AMNH with label "MEXICO: Tlalpam, D.F. [Distrito as in galathea. Surface rises immediately Federal], Apr.17. 1946, J, C. Pallister." into gentle mound covering all of poste- rior. Second curve of duct proceeds me- Etymology. An arbitrary combination dially. Markings (Figs. 133, 275): Cara- of letters, to be treated as an adjective. pace covered by scales mostly gray-white, Diagnosis. An enigmatic species from and some brown scales around fovea, and southcentral Mexico with a palpus in some just medial to posterior eyes. Clypeus ways resembling each of galathea, proter- densely covered with white scales. Abdom- va, and peckhamorum. The swollen base of inal markings gray-brown with large white the erect portion of the embolus is wider than in spots and small dark spots, much like gal- galathea, thought not so extreme athea (Fig. 275). Measurements: Body as in proterva. The retrolateral ramus is length 3.9, 4.3, 5.3 mm; carapace length wider than in galathea, though not so long 2.0, 2.1, 2.1 mm, width/length 0.75, 0.77, and hooked as in proterva. = 0.79; n 32 from Santa Cruz and Pima Male. Palpus (Figs. 193, 277, 278): Em- Counties, Arizona. bolus swollen at the base of the erect por- Male /Female Matching. The two sexes tion; narrowing distally near opening. were co-collected in Sycamore Canyon and Retrolateral ramus extended into short stout hook Kitt Peak, have similar markings on the (Figs. 193, 277). Markings (Fig. 276): abdomen, and are similar in markings and Cheek band fairly dense but not so dense form to galathea. as proterva. Clypeus brown; hairs over- Courtship (2<5 observed from Sycamore hanging chelicerae dark except for few = white Canyon, Arizona). Crouch (n 8, 26): Body medially. White forehead band con- low = tacts and horizontal (n 2, l6). First legs AMEs dorsally 10:30-12:30. Chelic- = fairly wide to bowed and parallel (n 6, erae with narrow medial patch of pale = = scales from base to V2 2(5), low (n 8, 2(5) to raised a bit (n 4, length. Cymbium = 1(5), waved on series (n 8, 26) at low lacking white scales. Measurements: Body = amplitude (n 4, 26). held down length 3.7(3.7-3.8)3.9 mm; carapace = Palpi (n length 2, 16), waved up and down on series so 1.7(1.8)1.9 mm, width/length 0.72(0.76) = as to drum on substrate (n = 2, still on 0.78; n 5(5 from Distrito Federal and = 16), pause (n 2, 16). Durango, Mexico. Female. Distribution (Map S). Southern Arizona. Epigynum (Figs. 4, 279, 280): Flaps long and convex, turning slightly in- Records. UNITED STATES: ARIZONA: Pima Co.: ward, shiny and generally pale. Surface Quinlan Mtns., picnic area nr. Kitt Peak Observatory, rises to mound quickly behind in 1,900-2,000 m el, 20 June 1985 (3<3 19, MCZ); Santa flaps; Cruz Co.: Sycamore Canyon, ca. 14 km W of Pena many specimens the mound has two dis- Blanca Lake, ca. 1,200 m el., 19 June 1985 {12S 19, tinct front corners (Fig. 280). Females MCZ); Santa Rita Mtns., Madera nr. Canyon, Bog from Oaxaca (Figs. 4, 279), which may ca. 1,.500 m 17 1985 Springs Cmpg(J., el., June (19. a MCZ) represent distinct species, have some- what longer epigynal flaps and a gentler Natural History. In oak woodland at all mound on the posterior surface. First curve three Arizona localities. At Sycamore Can- of duct wide; second curve proceeds yon, beating vegetation, especially shaded, obliquely anteriorly. Markings (Fig. 281): deep in canyon where stream still flowing Carapace covered with white scales. Clyp- 270 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

eus densely covered with white scales. Ab- Metaphidippus capitatus: -Bonnet, 1957: 2810, in domen marked somewhat as proterva, with part. brown background having white side bands Notes on Walckenaer's and central spots. Measurements: Body Synonymy. (1) length 3.4(4.8-5.2)5.4 mm; carapace length first description of Attus protervus (p. 443, after Abbot's 1.7(1.9)2.0 mm, width/length 0.75(0.77) fig. 402) probably refers to 0.80; n = 79 from Distrito Federal, Mexico. this Pelegrina; his second (p. 465, after Abbot's to in- Male/Female Matching. Males and fe- fig. 463) probably Maevia males were co-collected and have a com- clemens (see Walckenaer, 1837: 425; 1941: Attus mon distribution; they have similar size Chamberlin and Ivie, fig. 23). and markings. attentus Walckenaer (species number 61, Abbot's also be P. Distribution (Map 4). Mexico. Most fig. 157) may proterva. specimens from the Distrito Federal; also Attus capitatus Hentz, though considered known from Durango, Oaxaca, and San a synonym of proterva by Chamberlin and Luis Potosi. Ivie (1941), might equally well be P. gal- athea, Eris militaris, or another species. Records. MEXICO: SAN LUIS POTOSI: Guana- (2) Euophrys concolor Banks was synon- juato border on Hwy 57, ca. lOOMS'W, 21°35'N (19, ymized with P. proterva by Edwards MCZ); DURANGO. Pales Colorados, 5 August 1947 because one of the two {16 15, AMNH); DISTRITO FEDERAL: Contreras, (1980), apparently 2,500 m, 23 July 1947 (29, AMNH); Mexico City, specimens in the type vial is a 2 F. pro- 1941 8 1947 January (1

Coll. , examined. July, R. V. Chamberlin, Roewer, and narrower carapace and by (females) 1954: 1206. Bonnet, 1956: 1392. the abdominal markings and flat epigynal Metaphidippus protervus: —Chamberlin and Ivie, 1973: 43- flaps. Male much like those of 1944: 204 (not fig. 23). Kaston, 117, figs. markings 46, 69. peckhamorum, from which proterva dif- Pelegrina Jumping Spiders • Maddison 271

= fers in having a narrower embolus and width/length 0.75(0.76)0.79; n 69 from flatter epigynum. Palpus much like that of Iowa and Ontario. the central Mexican edrilana, differing in Geographical Variation. Southern fe- details. males have a darker carapace with distinct Male. Palpus (Figs. 3, 6-9, 283, 284): side bands, a face more sparsely covered Erect portion of embolus inflated basally, with pale scales, and an abdomen with the wide and transparent centrally. Terminal central pale spots coalesced medially into portion near opening much narrower than a chevron stripe, which is also seen in some basal portion. Retrolateral ramus a long southern males. Some males from Florida, curved hook (Fig. 283). Markings (Figs. Georgia, and South Carolina have a nar- 134, 282): Carapace with strong white rower embolus and lack the white scales markings, including forehead and side between the AMEs on the clypeus. bands. Cheek band dense and distinct from Chromosomes. 2nS = 26 acrocentrics -t- side bands. Clypeus with prominent dia- XXO (13, Mollis, New Hampshire). mond of white scales between AMEs and Courtship (106 observed from Shenan- overhanging chelicerae; lateral to this the doah Co., Virginia; Middlesex and Barn- clypeus and hairs overhanging chelicerae stable Counties, Massachusetts; Dorchester are dark. White forehead band contacts and Caroline Counties, Maryland; Thun- AMEs dorsally; setae ringing AMEs white der Bay, Ontario; Binscarth, Manitoba; see 7:00-12:00 and 2:00-4:00. Chelicerae also Peckham and Peckham, 1889: 45, fig. lacking pale scales. Femur of palpus dis- 18). Has the crouch display with body of- tinctly paler than more distal segments. ten high and first legs low. Raisedspread = = Cymbium with generally dense patch of (n 13, 53). Crouch (Fig. 134; n 15, 73): = white scales centrally. Second, third, and Body horizontal, held low (n 1), at about = = fourth legs with femur bases abruptly pale. normal height (n 6, 33) or high (n 8, = Abdominal dorsum usually light brown 3 3). First legs forward and low (n 15, = with dark spots between side bands. Mea- 73), either horizontal (n 3, 23) or even = surements: Body length 3.3(3.6)4.2 mm; lower than body (n 10, 43), bowed and = carapace length 1.6(1.6)1.9 mm, width/ parallel (n 4, 23), or slightly spread (n = = length 0.74(0.78)0.81; n 73 from Ontar- 3, 13). On one observation the legs were io, Iowa, and Saskatchewan. slightly raised at first but just before he Female. Epigynum (Figs. 238, 285, 286): touched 9 they were lower than body. First = Flaps fairly flat, dark, and con- legs waving little if a all (n 2, 13) or not slightly = = vergent. Surface smooth, rises fairly at all (n 4, 13). Palpi held down (n 7, = abruptly behind flaps into wide though 43) or forward (n 7, 33); moved forward = gentle mound covering most of posterior. (n 10, 53) and waved up and down on = First curve of duct broad; second curve each series (n 15, 73). Abdomen de- = proceeds obliquely anteriorly. Markings pressed on each series (n 1), twitches at = (Figs. 2, 135, 287): Carapace covered with least occasionally (n 4, 13). Repertoires: white and some brown scales. Face some- 33 raisedspread only; 53 crouch only; 23 what darker than galathea, relatively thin- raisedspread and crouch. Several times the ly covered with white scales, especially in display proceeded directly from the southern females. Legs only slightly an- raisedspread stage to touching the female = nulate, beige to light brown with darker without a distinct crouch display (n 8, markings reddish brown markings. Ab- 43). domen pale on sides, with two reddish Distribution (Map 5). Across much of brown longitudinal bands above broken by Canada and northeastern United States, oblique to transverse white stripes. Mea- south in the east to Florida and Texas. surements: Body length 4.4(5.1-5.3)5.6 Although the following records include mm; carapace length 1.6(1.9)2.0 mm. only few from Canada, this is due to my 272 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

examining only collections at museums in P. galathea. Dondale (1961) describes the the United States; proterva is actually very life history of P. proterva in Nova Scotia. common throughout much of Canada. 6. Pelegrina peckhamorum Records. Manv in MCZ and specimens, especially (Kaston, 1973) from: CANADA: NOVA SCOTIA: Kentville; AMNH, new combination QUEBEC: St. Louis de France, Quebec City; ON- 137, 195,239, TARIO; Martin River 58 km N of North Bay, Ottawa, Figures 126, 136, Belleville, Barrie, 36 km E of Thunder Bay, Sudbury 288-293; Map 8 Dist.: Espanola, 15 km E of Espanola. nr. Bruce Mines Pte. Baril nr. Sault St. Marie, 14 km S of Au (Parry Metaphidippus peckharrioruin Kaston, 197.3; 115, figs. Sound Dist.), 20 km E of Manitoba border on Hwy 39-42, $9. Holotype 5 and paratype 9 in AMNH 17; Lake Temagami, Port Credit; SASKATCHE- with labels "Holotype S + allotype 9, Metaphidip- MANI- n. B. Kaston WAN; Waskana Creek, North Battleford; pus peckhamorum sp., det by J. (1949)" N. TOBA: Sandilands Provincial Forest, Binscarth, Gyp- and "col. by B. Malkin, Lakehurst, J. 25 May sumville; BRITISH COLUMBIA: Salmon Arm. 1941," examined. Brignoli, 1983; 643. UNITED STATES (county records); MAINE; Han- cock, Lincoln; NEW HAMPSHIRE; Belknap, Carroll, Diagnosis. A relatively rare eastern spe- VER- Cheshire, Grafton, Hillsborough, Sullivan; cies with male body form and markings MONT; Addison, Caledonia, Chittenden, Rutland, much like but Windham, Windsor; MASSACHUSETTS; Barnsta- very proterva outstanding for its broad embolus. The female is ble, Berkshire, Dukes, Esse.x, Franklin, Hampden, very Middlesex, Nantucket, Norfolk, Suffolk, Worcester; best distinguished from other eastern spe- ISLAND; CONNECTICUT: RHODE Washington; cies by the indistinct markings and large, Fairfield, Middlesex, New Haven, New London, Tol- slightly concave epigynum, with flaps that land; NEW YORK; Hamilton, Nassau, New York City are more convex than in and that Thompkins, Westchester; NEW JERSEY: Bergen proterva Cape May, Hunterdon, Passaic; PENNSYLVANIA have the posterior edge not truncate as in Monroe Adams, Bucks, Carbon, Centre, Forest, galathea. Montgomery, Schuylkill, Warren, York; OHIO Male. Palpus (Figs. 195, 289, 290): Em- Champaign; MARYLAND: Caroline, Charles bolus broad, but still broad Dorchester; WEST VIRGINIA; Mercer; VIRGINIA very tapering retrolateral Fairfax, Rockingham, Shenandoah, Suffolk, Wash- at tip. Rami well separated; ington; KENTUCKY; Hardin, Rowan; TENNESSEE; ramus not elongate as in proterva. Mark- Grundv, Sevier, Unicoi; NORTH CAROLINA; Bun- ings (Figs. 136, 288): Cheek band dense combe! Durham, Macon; SOUTH CAROLINA; Hor- and discrete. Clypeus with prominent di- ry; GEORGIA: Cobb, Polk, Thomas; FLORIDA; Ala- of white scales between AMEs and chua; ALABAMA: Clarke, Dekalb, Jackson; MICH- amond IGAN; Berrien, Calhoun, Charlevoix, Cheboygan, overhanging chelicerae; lateral to this the Crawford, Eaton, Emmet, Genesee, Hillsdale, Isa- clypeus and hairs overhanging chelicerae bella, Jackson, Kent, Lake, Livingston, Mackinac, are dark. White forehead band contacts Marquette, Midland, Muskegon, Osceola, Sanilac, AMEs setae AMEs white Washtenaw, Wayne; INDIANA: Jasper, Marion, dorsally; ringing Starke; WISCONSIN; Ashland, Chippewa, Crawford, 7:00-12:30 and 2:00-4:00. Chelicerae Dane, Door, Douglas, Grant, Green Lake, Iowa, Jef- lacking pale scales. Femur of palpus dis- ferson, Lincoln, Manitowac, Marathon, Price, Rich- tinctly paler than more distal segments. land, Rusk, Sauk, Shawano, Waushara; IL- Taylor, with white scales LINOIS; Cook, Mason; MINNESOTA: Cymbium centrally. Leg Champaign, Measure- Blue Earth, Freeborn, Marshall, Olmsted, Steele, Wi- femora distinctly paler basally. nona; IOWA; Boone, Cla\ ton, Hancock, Winnebago, ments: Body length 3.0(3.6)3.7 mm; car- MISSOURI; Woodbury; Boone, Cole, Jackson, John- apace length 1.4(1.7)1.8 mm, width/length son, St. Louis; NEBRASKA; Lancaster, Loup, Saline; = 0.74(0.76)0.78; n 53 from Barnstable KANSAS: Decateur, Rile\; TEXAS: Anderson, Den- Massachusetts. ton, Hardin, Sabine, San Jacinto; MONTANA; Ra- County, valli, Stillwater; COLORADO: Fremont, Larimer. Female. Epigynum (Figs. 239, 291, 292): Large. Flaps long, fairly convex, usually Natural History. Found on various trees convergent. Surface rises very gradually and shrubs, usually in or near forests; less behind flaps; most of posterior area con- often found in fields and on herbs than is cave. First curve of duct broad; second Pelegrina Jumping Spiders • Maddison 273

curve proceeds obliquely anteriorly. 62, MCZ), 24 km W of Prairie Grove (53 12, MCZ); TEXAS: Leon Co.: SW of Oakwood (13, AMNH). Markings (Figs. 137, 293): Carapace cov- ered with scales. cov- yellowish Clypeus Natural History. May specialize on oaks. ered with white scales. thinly yellowish On Cape Cod, Massachusetts, collected by Abdomen more uniform in color than pro- beating oaks and cranberries in understory terva, brown with Mea- light pale spots. of pine forest (1 record), sweeping oak- surements: 3.6(4.0)5.4 Body length mm; pitch pine (2 records), and beating oaks (1 carapace length 1.7(1.9)2.1 mm, width/ = record). length 0.73(0.77)0.78; n 52 from Mas- sachusetts and Arkansas. 7. Pelegrina neoleonis new species Male/Female Matching. Males and fe- Figures 138, 196, 294-298; Map 6 males have been co-collected in New Jer- Holotype male and paratype female in MCZ with and other- sey, Arkansas, Massachusetts; label "MEXICO: NUEVO LEON: Chipinque Mesa wise, they are the only unmatched 69 in just S of Monterrey, ca. 4500 ft. [1,370 m]; ca. the northeast. 100.4°W 25.6°N, 2 Jun 1983 W. Maddison & R, S. Anderson 83-034, and forest un- Courtship (43 observed from Cape Cod, beating sweeping derstory." Massachusetts). Has the crouch display with body high and first legs low as in P. pro- Etymology. After the state from which = terva. Raisedspread (n 17, 33). Crouch most known specimens come. = (Fig. 126; n 12, 33): Body held normal- Diagnosis. A Mexican species similar to = = high (n 6, 13) or high (n 6, 23). First tristis with a distinctive long, curved retro- = legs held horizontal (n 12, 33) and lower lateral ramus on the embolus. The erect = than body (n 3, 13), waved little if at all portion of the embolus is narrower than in = = (n 9, 23). Palpi held down (n 12, 33); tristis. No characters have yet been found = still on pause (n 2, 13), waved on series to distinguish the female from that of tris- = = (n 4, 23) up and down (n 5, 13), spe- tis, except locality. = cifically from down to forward (n 2, 23), Male (from Nuevo Leon). Palpus (Figs. = with medium-high amplitude (n 5, 13). 196, 295): Embolus distinctive; broad, with = Abdomen still on series (n 4, 23) but retrolateral ramus extended into long hook, = twitched on pause (n 4, 23). Repertoires: much as in tristis, but ramus bears small 13 raisedspread only; 23 raisedspread and bump and is blunt at tip; prolateral ramus crouch; 13 crouch only. obtuse or only slightly acute. Markings Distribution (Map 8). Known from Mas- (Figs. 138, 294): Cheek band weak. Clyp- sachusetts, New York, New Jersey, Ohio eus brown; hairs overhanging chelicerae (Kaston, 1973), Indiana (Kaston, 1973), dark. White forehead band contacts AMEs Tennessee, Arkansas, and Texas. dorsally 10:30-12:30. Chelicerae lack pale scales. Femur of palpus distinctly pale than Records. UNITED STATES: MASSACHUSETTS: more distal segments. Cymbium with none Barnstable Co.: Chatham (13 3 2, MCZ), South Chat- to few white scales. Femur of third leg ham (6<5, MCZ), nr. North Truro at junction of Hwy basal Vs. 6 and Head of the Meadow Road (103 32, MCZ); pale on Measurements: Body Dukes Co.: Bluffs Oak (1<3, MCZ); NEW YORK: Dav- length 3.6, 3.7 mm; carapace length 1.7, isville; Suffolk Co.: Riverhead Coram (26, AMNH), 1.8, 1.9 mm, width/length 0.75, 0.75, 0.76; (2(3 12, NEW Co.: 11 AMNH); JERSEY: BurHngton n = 33 from Nuevo Leon and San Luis km W of New Gretna (4<5 22; AMNH), Lebanon State Potosi. Forest (1<5, AMNH); Middlese.x Co.: Old Bridge (1<5, AMNH); Morris Co.: Chatham, Great Swamp (12, Female. Epigynum (Figs. 296, 297): AMNH); Ocean Co.: Lakehurst (223 112, AMNH), Flaps large and dark, flat and inwardly Lake Horicon nr. Lakehurst (32, AMNH), 6 km W rotated. Surface gently convex, highest of Lakehurst (23 32, AMNH); TENNESSEE: Knox medially behind flaps, except for surface Co.: University of Tennessee farm 3 (13, AMNH); ARKANSAS: Washington Co.: 24 km S of Prairie diving deeply under flaps. First curve of Grove in Cove Creek Valley of the Boston Mtns. (283 duct very broad, expanded to the side and 274 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Records. MEXICO: SAN LUIS POTOSI; 21 km W posterior so that second curve begins well of Xilitla on Hwy 120, 99°05'W, 21°18'N, 12 June of posteriormost portion of flap posteriad 1983 (13, MCZ); NUEVO LEON: Chipinque Mesa the (in contrast with sympatric clavator); just S of Monterrey, 100. 4°W, 25.6°N, 2 June 1983 second curve proceeds anteriorly. Inner {1$ 12, MCZ); Cerro Potosi, ca. 100°14'W, 24°52'N, 4 1983 HIDALGO: Pachuca surface of third curve rough, with numer- June (15 12, MCZ); (12, MCZ); OAXACA: 50 km NW of Oaxaca, 97°00'W, ous projections. Markings (Fig. 298): Car- 17°I4'N, 6 August 1983 (22, MCZ). apace dark above, covered with transpar- ent reflective scales; sides covered thinly Natural History. Beating oaks and pines with white scales. Clypeus densely covered in oak-pine area (3 records); sweeping with white scales. Legs brown. Abdomen shrubs, cloud forest (1 record). Elevations at four locations in Leon and San fairly dark, with only small pale spots. Nuevo Measurements: Body length 4.3(5.9- Luis Potosi from 1,400 to 2,900 m. 6.1)6.2 mm; carapace length 1.9(2.2- 8. Pelegrina tristis new species 2.3)2.3 mm, width/length 0.74(0.77- = Figures 197, 299-303; Map 7 0.79)0.83; n 4$ from Nuevo Leon, Hi- dalgo, and Oaxaca. Holotype male and paratype female in AMNH with Male/Female Matching. This associa- labels "ARIZONA: Cochise Co., Round Park, Chir- icahua Mtns., June 28, 1967. 9300 ft. [2,840 m], tion is indicated by co-collecting in Nuevo Gertsch, Hastings." Leon; by the large epigynal flaps, which would be expected in a species with such Etymology. Latin adjective for "sad," a robust embolus; and by the similarity of referring to the large size of the teardrop- male and female with those of tristis. shaped flaps over the epigynal openings. Geographical Variation. The single male Diagnosis. A large, dark, plainly marked from San Luis Potosi differs from those of species known from southern Arizona, Nuevo Leon in having a sharper retrolater- similar in genitalia to neoleonis and sa- al prong on the embolus, more extensive binema. The erect portion of the embolus white scales surrounding AMEs, a small is broader than in either of those species, patch of pale scales medially on chelicerae, and the rami are sharper than in neoleonis. and the femur of palpus relatively dark. Females are generally not so yellow as in Courtship (33 observed from Chipinque sabinema, and the epigynal openings are Mesa, Nuevo Leon; Cerro Potosi, Nuevo deeper, in that the surface descends more Leon; and Xilitla, San Luis Potosi). Raised- deeply under the anterior part of the flaps. = = spread (n 3, 2(5). Crouch (n 6, 33): Male. Palpus (Figs. 197, 300): Embolus = Body held in normal to low position (n extremely broad, so that retrolateral mar- = 1). First legs bowed and forward (n 4, gin joins without angle to retromargin of = 23), raised to ca. 30° (n 2, 23), or hori- embolar base. Both rami sharply pointed; = zontal (n 1), or femora low but tips curl retrolateral ramus extended into long hook, = upward (n 1). Leg tips not touching (n lacking subterminal bump. Markings (Fig. = = 2, 23), apparently not waved (n 3, 23), 299): Carapace dark, with reduced fore- = or waved only slightly (n 2, 13) on series head band. Cheek band very weak to ab- = = (n 1). Palpi down (n 6, 33), over che- sent. Clypeus brown, with dark hairs over- = licerae (n 1) or curled under tips of che- hanging chelicerae. White forehead band = = licerae (n 1), waved (n 6, 33) up and absent or much reduced, fails to contact = = down (n 1) or outward (n 1) on series AMEs, which are ringed with dark above. = = (n 3, 23) ca. 5-7 c/s (n 1). Abdomen Chelicerae lacking pale scales. Palpus al- = depressed a bit on series (n 1), or at end most uniformly brown, femur not distinct- = of series (n 1). Repertoires: 13 crouch ly paler. Cymbium lacking white scales. only; 23 raisedspread and crouch. Legs relatively uniform brown, femora en- Distribution (Map 6). Northeastern tirely dark. Measurements: Body length Mexico south to Oaxaca. 3.7(4.3)4.6 mm; carapace length 1.8(2.1)2.1 Pelegrina Jumping Spiders • Maddison 275

= mm, width/length 0.75(0.76)0.82; n 45 Notes on Specific Distinctness. Pele- from Chiricahua and Santa Catalina Mtns., grina sabinema is much like pervaga, and Arizona. indeed I long considered it only the west- Female. Epigynum (Figs. 301, 302): ern form of pervaga, but the more strongly Flaps large, dark, and convergent, often developed white-black-white carapace far rotated, sometimes as far rotated as in stripes and narrower embolus of pervaga neoleonis (Fig. 297). Just medial to the flap suggest that pervaga may be the sister spe- at the anterior end the surface is pale and cies to kastoni, with sabinema the sister to descends deep under flap (Fig. 302, ar- those two. The embolus and markings of row). Except for this concavity, the epi- P. sabinema are slightly more like those gynal surface is gently convex, highest me- of tristis and neoleonis, which may be con- dially behind flaps. First curve of duct very sidered outgroups. broad; second curve proceeds anteriorly. Diagnosis. Differs from pervaga in hav- Markings (Fig. 303): Carapace covered ing less swollen carapace sides, an abdo- above thinly with white to dark transpar- men lacking the pale central stripe on the ent reflective scales. Clypeus densely cov- abdomen, wider embolus, weaker male ered with white scales. Abdomen light to cheek band, less dense band of dark hairs medium brown with small central pale beneath male carapace side bands, darker spots. Narrow dark brown spots beside and more robust epigynal flaps, and yellow these pale spots form longitudinal dark female legs. Differs from tristis in having stripes. Measurements: Body length yellow legs, yellow male chelicerae, nar- 5.0(5.7)6.2 mm; carapace length 1.8(2.0)2.3 rower embolus, dense covering of pale - mm, width/length 0.77(0.78)0.82; n 59 scales on female carapace, and shallower from Chiricahua, Huachuca, and Santa epigynal openings. Rita Mtns., Arizona. Male. Palpus (Figs. 198, 305): Embolus Male/Female Matching. This match- very wide at base of erect portion, thought ing is indicated by microsympatry in Chir- still with a distinct angle between retro- retro- icahua Mtns., by robust embolus and flaps, margins of erect portion and base; by similar large size, and by similarity of lateral ramus long and blunt. Carapace genitalia to male and female of sabinema, often broad though sides not swollen as in which are reasonably surely matched. pervaga. Markings (Fig. 304): Cheek band runs and be- Distribution (Map 7). Southern Arizona. weak, horizontally posteriorly neath band of dark hairs beneath white Records. UNITED STATES: ARIZONA: Santa Rita side bands. brown, hairs over- Mtns.: Madera Canyon (49, AMNH, MCZ); Huachuca Clypeus Mtns.: Garden Canyon (19, AMNH); Santa Catalina hanging chelicerae dark. White forehead Mtns.: Bear Wallow to Mt. Lemmon (19, AMNH), band contacts AMEs rather far medially, Chiricahua Mtns.: Round Southwestern Re- Park, from 9:00 to 12:00. Chelicerae yellow, search Station 8 km W of Portal, Barfoot Park, and lacking pale scales. Palpus yellow with Rustler's Park (3<3 49, AMNH). white scales on femur, tibia and cymbium Natural Collected at 1,500- History. interrupted by dark hairs on patella and 2,800 m elevation Females have (3 records). base of cymbium. Legs uniformly yellow. been collected in June (2 records), July (4 Abdomen brown centrally with white side and records), August (3 records). bands, showing trace of paired dark spots of female. Measurements: 9. Pelegrina sabinema new species Body length 9 3.3(3.6)3.8 mm; carapace length 1.7(1.7)1.9 Figures 198, 304-308; Map - mm, width/length 0.78(0.80)0.83; n 55 Holotype male in AMNH with label "ARIZONA, from New Mexico and Arizona. Showlow, July 1967, W. J. Gertsch." Female. Epigynum (Figs. 306, 307): Etymology. An arbitrary combination Flaps flat and large, often far rotated, as of letters, to be treated as an adjective. in tristis and neoleonis, though usually not No. 4 276 Bulletin Museum of Comparative Zoology, Vol. 154,

labels "Dendryphantes pervagus P., 9 Wallace, so dark as in those species. Openings shal- Kansas type" (label is original; handwritten, prob- lower than in tristis; that is, just medial to ably by Elizabeth Peckham) and "G. W. Peckham the at the anterior end the surface is flap Coll.", examined. Roewer, 1954: 1214. — not so pale and does not dive deep under Dendryphantes (Metaphidippus) prevagus [sic]: First curve of Petrunkevitch, 1911: 640. flap. Epigynal surface flat. Metaphidippus pervagus: — Bonnet, 1957: 2817. duct very broad; second curve proceeds in tristis. less anteriorly than Markings Diagnosis. A striking species with swol- covered with (Fig. 308): Carapace well len carapace sides and central pale stripe white to yellowish scales. Clypeus very on the abdomen in both sexes. Very similar densely covered with white scales. Legs to sabinema, from which it is distinguished with brown yellow. Abdomen yellowish by the features discussed under that spe- centrally, paired white spots. Usually cies. paired dark brown spots in posterior half Male. Palpus (Figs. 199, 310); Embolus are beside white spots. Measurements: wide basally; retrolateral ramus long, Body length 4.0(4.2)5.5 mm; carapace pointing distally, having subterminal length 1.6(1.7-1.8)1.9 mm, width/length bump. Carapace sides swollen. Markings - 0.78(0.80-0.81)0.86; n 69 from New (Fig. 309): White carapace side band bor- Mexico and Arizona. dered below by narrow band of black hairs. Male/Female Matching. Male and fe- Below this, the dense white cheek bands males were matched by similar yellow col- do not reach clypeus, which is dark. Hairs or, by robust embolus and flaps, and by overhanging chelicerae dark. White fore- co-collecting and common distribution in head band contacts AMEs far medially, New Mexico and northern Arizona, where from 9:00 to 11:00. Chelicerae yellow, no other unmatched females and males are lacking pale scales. Palpus yellow, with known. white scales on end of femur, on tibia and Distribution (Map 9). New Mexico, cymbium, interrupted by dark hairs on southern and northern Arizona, Colorado, patella. Legs light yellowish brown with western Texas. some darker annulations. Abdomen with central as in fe- Records. UNITED STATES: TEXAS: Jeff Davis longitudinal pale stripe Co.: 24 km NW of Fort Davis (19, AMNH); COL- male. Measurements: Body length Park ORADO: Montezuma Co.: Mesa Verde National 3.7(3.9)4.4 mm; carapace length 1.8(1.8)2.0 NEW MEXICO: Bernalillo Co. (29, = (1<5, AMNH); mm, width/length 0.83(0.85)0.88; n 5<5 AMNH); Lincoln Co.: nr. Ruidoso, Ruidoso Cmpgd. from Erath Co., Texas. (22, AMNH); Los Alamos Co.: nr. Los Alamos (19, AMNH); Sandoval Co.: (19, AMNH), Sandia Mtns., Female. Epigynum (Figs. 240, 311, 312): Juan Tabo area (29, AMNH); Santa Fe Co. (26 19, Flaps fairly large and flat, generally pale. 5 AMNH), Glorieta Mesa nr. Rowe (13 19, AMNH), Surface flat. First curve of duct wide; sec- km N of Galiseo (13, AMNH), Route 66 just E of ond curve anteriorly. 19 km S of proceeds obliquely Edgewood (19, AMNH), Lamy (19, wide and AMNH); Taos Co.: 27 km S of Taos (13 19, AMNH); Markings (Fig. 313): Carapace ARIZONA: Coconino Co.: Flagstaff (13 19, UCB), covered with whitish scales. Clypeus cov- Navajo Co.: Showlow (13, AMNH). ered densely with white scales. Abdomen on middle of Natural History. Collected at 7,000 ft with distinctive pale patch dorsum. Measurements: elevation (2 records), from pinyon pine- Body length 4.3(4.8)5.9 mm; carapace length 1.8(2.0)2.2 juniper (1 record). = mm, width/length 0.80(0.82)0.85; n 59 1 0. Pelegrina pervaga from Erath Co., Texas. & E. (G. Peckham, 1909) Distribution (Map 10). Texas, Oklaho- new combination ma, and Kansas. Figures 199, 240, 309-313; Map 10 Records. UNITED STATES: KANSAS: Wallace Dendryphantes pervagtis G. & E. Peckham, 1909: Co.: Wallace (19, MCZ); OKLAHOMA: Commanche 474, pi. 37, figs. 9, 9a, 9. Holotype in MCZ 19 with Co.: Visitor's Center, Wichita Mtns. (13, WPM); TEX- Pelegrina Jumping Spiders • Maddison 277

Erath Co.; 11 km NE of AS: Stephenville (6<5 79, Female. Epigynum (Figs. 316, 317): TXAM). Flaps rotated a full 90°, with deep openings anterior to in almost Natural History. Collected from juni- just them, resting pits, as in pers in Texas (three records). Dendryphantes nigromaculatus though flaps maintain their prominence as

1 1 . l

raisedspread or early crouch only; IS head in the northeastern United States crouch only. iflaviceps), and a dark southern species Distribution (Map 11). Southern Ari- (exigua). The situation is not nearly so sim- zona, southwestern New Mexico, and Chi- ple as this, however, for hybridization may huahua. occur at the borders of their ranges (dis- cussed under flaviceps) and two species Records. UNITED STATES: ARIZONA: Santa may be confused under the name exigua Cruz Co.: Madera (6

Records. in MCZ and ments: Body length 3.6(3.7)4.3 mm; car- Many specimens, especially AMNH, from: CANADA: NORTHWEST TERRI- apace length 1.7(1.9)2.0 mm, width/length TORIES: Mackenzie: Prelude Lake IIS'SS'W, n = 5<5 from 0.74(0.76)0.79; Neepawa, 62''33'N; Lady Evelyn Falls 117°19'W, 60°57'N; Manitoba. NEWFOUNDLAND: Humber River; PRINCE ED- WARD ISLAND: Tracadie; City; Female. Epigynum (Figs. 241, 321, 322): QUEBEC: Quebec NOVA SCOTIA: Weymouth, Harrington, Baddeck Flaps flat, parallel or only slightly conver- (Cape Breton), North Sydney; ONTARIO: Ottawa, Surface flat. First curve of duct broad; gent. Marten River 58 km N of North Bay, Parry Sound second curve narrow, beginning from pos- Dist.: 14 km S of Pte. Au Baril Station, Algoma Dist.: terior end of first curve and proceeding near Bruce Mines; Sudbury Dist.: Espanola; Thunder toward the midline; flowerlike Bay Dist.: 7 km E of Nipigon; Kenora Dist.: Granite anteriorly Lake Tema- on dorsal face of duct. Lake; Lakefield, Chapleau, Sowerby, gland openings 56 km E of Hearst, Sioux Lookout, Gawas Bay, cov- gami, Markings (Figs. 143, 323): Carapace Kamiskotia Lake, Point, Moberly, Uxbridge, ered with transparent brown scales with a Spanish River, Iron Bridge, St. Williams, Batchawana, brassy sheen. Beige spots on forehead be- Nipigon, Dorset, L. Opeongo, Cababogie (Tweed), Pancake Bay nr. Batchawana, Emo, tween and beside AMEs recall white spots Haileybury, Fairbank Lake Province Park, Nestorville, Golden of male and are than in usually stronger Lake, Minden, SouthTea Lake (Algonquin); MANI- exigua. Setae directly above AMEs brown. TOBA: Kettle Rapids, Cedar Lake, Lyons Lake, 19 AL- Clypeus generally densely covered with km E of Neepawa, Sandilands Provincial Forest; BERTA: Banff, Athabasca Land- white scales. Abdomen with brassy sheen Edmonton, Jasper, ing, Fitzgerald, North Lake Athabasca; BRITISH and fourth of white spots formed into pair COLUMBIA: Wells Cray Park, Columbia Lake, distinct chevron. Measurements: Body Salmon Arm, Arrow Lakes. UNITED STATES length 4.4(4.7)4.8 mm; carapace length (county records): MAINE: Aroostook, Penobscot, Pis- NORTH CAROLINA: 1.8(1.9)2.0 mm, width/length 0.74(0.77) cataquis, Washington; Avery, Buncombe; MICHIGAN: Allegan, Baraga, Calhoun, 0.77; n = 52 from Neepawa, Manitoba. Charlevoix, Cheboygan, Chippewa, Crawford, Delta, 2n<5 = 26 acrocentrics + Chromosomes. Emmett, Grand Traverse, Ingham, Mackinac, Mar- XXO (16 Nipigon, Ontario; 16 Edmonton, quette, Oakland, Washtenaw; WISCONSIN: Chip- Alberta). pewa, Lincoln; ILLINOIS: Lake; MINNESOTA: Clearwater, Hennepin; MONTANA: Jefferson, Ra- Courtship (43 observed from Edmon- valli, Sanders; IDAHO: Fremont; WYOMING: Crook, ton, Alberta, and Manitoba). Neepawa, Lincoln, Park, Teton; COLORADO: Archuleta, Boul- With unusual alternate waving of palpi NEW MEXICO: Sandoval, San Miguel, = der, Gilpin; during crouch display. Raisedspread (n Taos; WASHINGTON: Okanagan, Stevens. = 1). Crouch (n 7, 45): Body held at normal = Natural A conifer dweller, col- height (n 5, 2S) or somewhat raised (n History. = lected from (11 records from On- 1). First legs forward and horizontal (n spruce = = tario, Manitoba, and Alberta), including 2, 2(3) with tips raised (n 1) or not (n = = white (2 records); pines (4 records 1), or whole leg raised slightly (n 5, spruce = from Ontario, British Columbia, and 2(5). First legs apparently not waving (n = pine (1 2, 23). Palpi held down (n 1) and some- Michigan), including lodgepole = and (1 record); junipers what forward (n 2, 23). Palpi wave (n record) jackpine = = records from Ontario); and larch (2 re- 6, 33) up and down alternately (n 5, (3 = cords from Ontario and Illinois). 23), fairly slowly, on series (n 1); spe- cifically, palpi tips wave in small circles (n = 13. 1), left clockwise and right counterclock- Pelegrina flaviceps = wise or vice versa (n 1). Abdomen (Kaston, 1973) combination twitches with low amplitude on series (n new = 144, 145, 202, 242, 324-328, 1). Repertoires: 33 crouch only; 13 Figures 13 raisedspread and crouch. 340, 341; Map Distribution 12). Across much of (Map Metaphidippus flaviceps Kaston, 1973: 110, figs. 15- Canada and northeastern United States, 20, (39. Holotype S and paratype 2 in AMNH with S + south along the Rocky and Appalachian labels "Holotype allotype 2, Metaphidippus det B. Kaston and Mountains. flaviceps n.sp., by J. (1949)" 280 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

"Clarendon, Vt., 2 Sept. 1939, E. M. Greenspan," distinct. Lacks white spots on forehead. examined. Brignoli, 1983: 643. Cephalic area appears yellow in alcohol, be dark in life (Fig. 144) and Notes on Specific Distinctness. Appar- though may is covered with small dark hairs. Cheek ent hybridization between this species and band dense and wide. brown, hairs the other two in the group makes their Clypeus overhanging chelicerae dark. Setae sur- separation doubtful. Males from northern AMEs brown to brown en- Massachusetts (Pepperell, Groton, East rounding light Chelicerae with deep, Templeton), southern New Hampshire tirely. yellow long, dark black spot medially, and brown spot (Hollis), and Ithaca, New York, show grades near the base. lateral to the of intermediacy between exigua and flav- laterally Just in medial black spot the surface is flat to iceps. Of 25 males collected at one site concave. Palpus light brown Pepperell, 9 have dark legs and a flat ce- slightly throughout, not noticeably phalic area (as in exigua farther south), 10 cymbium darker. lacks white scales. Legs have dark legs and a bulbous cephalic area, Cymbium pale yellow with thin black prolateral lon- 1 has yellow legs and a flat cephalic area, stripe on all femora except some- and 5 have yellow legs and a bulbous ce- gitudinal times lacking on posterior legs. First leg phalic area (as in flaviceps farther north). with of white hairs. Measurements: At this site, even those with yellow legs fringe 3.7(3.8)3.8 mm; carapace and bulbous carapace have a wider cara- Body length 1.8(1.8)1.9 mm, width/length pace than is usual for flaviceps. Courtship length n = 53 from Co., behavior of the apparent hybrids from 0.72(0.73)0.74; Sagadahoc Maine. Pepperell appears like that oi flaviceps and Female. 326, 327, 340, exigua (35 observed). In Michigan and Epigynum (Figs. to Maine, on the northern edge of the range 341): Flaps parallel convergent. Epi- surface flat. First curve of duct broad, of flaviceps, males otherwise like fla- gynal not so as in second curve vipedes have been found with the bulbous long flavipedes; broader and more medially directed than cephalic area. in not nearly so broad Diagnosis. Males can be distinguished flavipedes, though as in flowerlike on from those of flavipedes and exigua by the exigua; gland openings dorsal face of duct. Sometimes bulbous cephalic area, dark lateral spots Carapace: with swollen area. on the chelicerae (Fig. 324), carapace dust- cephalic Markings (Fig. dark on fore- ed with pale scales (Fig. 144), and pale 328): Carapace uniformly of scales behind yellow legs. Females can be difficult to head, lacking patches pale distinguish from those of flavipedes and anterior eyes. Clypeus densely covered exigua; they are generally paler, lack white with white scales. Chelicerae with prom- spots on the forehead, often have narrow inent medial black spot reminiscent of dark lines on the femora, and may have males. Some females with thin longitudi- the cephalic area slightly swollen. The epi- nal lines on leg femora. Abdomen usually gynal flaps and ducts are intermediate be- indistinctly marked, sometimes with dark tween those of flavipedes and exigua. The areas formed into longitudinal bands on medial black spot on the chelicerae is much either side of middle paler area. Measure- more distinct in flaviceps females than in ments: Body length 3.7(4.0)4.6 mm; car- females. apace length 1.7(1.8)1.9 mm, width/length flavipedes = Male. Palpus (Figs. 202, 325): Embolus 0.74(0.74)0.75; n 59 from Sagadahoc Co., divided deeply into two rami; prolateral Maine, and Durham, New Hampshire. ramus of embolus not as twisted as in ex- Courtship (43 observed from Reid State igua. Carapace is swollen dorsally in ce- Park, Maine). With unusual alternate cir- phalic area; carapace narrowest of group, cling of palpi during crouch stage. Raised- = until 2 or 3 especially at front. Markings (Figs. 144, spread (n 2, 2(5): continued 324): Carapace and abdomen dusted with body lengths from female when male went = = pale scales, so as to make side bands in- into crouch stage (n 2, 26). Crouch (n Pelegrina Jumping Spiders • Maddison 281

= "Nathan Banks Coll.", e.xamined. Roewer, 1954: 10, 46): Body held high (n 6, 2S) or low = 1209. (n 1). First legs forward and bowed (n = = Dendryphantes virginis Chamberlin, 1925a: 2.33. 5, 3(3), tips close (n 2, 15); legs move material said to be in MCZ Chamberlin = Type by more parallel as he gets close (n 1). Fe- but no holotype found therein. Paratypes from = District of Columbia Bladen- mur low but raised distally (n 5, 3<3), or Woodridge, {33 19), burg, Maryland {2S), examined. Bonnet, 1956: 1402. legs may be horizontal and lower than body — = Metaphidippus virginis: Muma, 1944; 11. = 4, down and forward (n (n 1(5). Palpi Metaphidippus exiguus: — Kaston, 1945: 10. Kaston, flicker = for few 3, IS). Palpi (n 9, 4$) 1973: 112, figs. 26-29, <59. = — 1957: 2813. seconds (n 4, 3(5), then pause for few Metaphidippus flavipedes: Bonnet, = seconds (n 3, 2(5). Flicker-pause cycle = 3-4 = or more repeats (n 5, 36), (n 1) Notes on Synonymy. The holotype fe- than 8 times = flicker is of (n 1). Palpus male of D. exiguus is peculiar in having low and = amplitude high frequency (n the palpus tarsi swollen, as in antepenul- and is on 2, 2(5) superimposed larger up timate instar males, yet has a well-devel- and down circular wave; slightly right pal- oped epigynum. There is doubt as to the pus moving up as left down and vice versa for the = placement of this specimen, epi- (n 2, 2(5). Abdomen twitched occasion- = gynum is much more like that of flaviceps = when flickered (n ally (n 9, 43) palpi than of southern species that has been called For much of male stood 5, 3(5). display, exiguus, with the flowerlike glands on the without when he did walking, thought face of the duct, not hidden by a fold, and and abdomen were flickered palpi during the duct not nearly so broad (Fig. 342) as series = and = 3, (n 1) during pause (n 26). is usual in the southern species. However, male to mount after about Once, proceeded the markings of the type are like those of 3-4 flicker of crouch = palpus cycles (n the southern species (white scales between 2S crouch 2S raised- 1). Repertoires: only; anterior eyes on forehead, no stripes on and crouch. spread legs, wide carapace, abdominal markings Distribution Northeastern (Map 13). more uniformly dark). Males collected United States and southeastern Canada. from Ithaca, New York (including a male label Records. Over 1003 1009 in AMNH and MCZ from: in the MCZ with a Bryant indicating CANADA: QUEBEC: Lake Champlain; ONTARIO: that it was collected with Banks's female STATES Toronto, Newmarket, Ottawa. UNITED type of exiguus), are in most respects like records): MAINE: Cumberland, Hancock, (county exigua but have a head bump like flavi- Penobscot, Sagadahoc, Waldo; NEW HAMPSHIRE: It therefore that the Carroll, Cheshire, Coos, Grafton, Strafford; VER- ceps. appears type MONT: Caledonia, Rutland, Windham; NEW YORK: locality of exigua is along the hybrid zone Albany, Cortland, Essex, Franklin, Greene, Hamil- with flaviceps (see notes under flaviceps). ton, Lewis, Nassau, Oneida, Onondaga, Schoharie, Specimens of pure flaviceps occur in towns Steuben, Yates; PENNSYLVANIA: Pike; Tompkins, near Ithaca. Until variation at the type WEST VIRGINIA: Preston; MICHIGAN: Charle- is better Kaston's voix, Kalkaska. locality understood, ap- plication of the name exigua to the south- Natural A conifer dweller, col- History. ern species will be maintained. lected from spruce (2 records), junipers To further complicate the application and and pine (1 record), spruces, firs, pine of the name exigua, two distinct color forms and hemlocks (1 record), (1 record). are found in the south that may very well represent distinct species. The typical (dull) 14. Pelegrina exigua (Banks, 1892) form (Figs. 146-147, 329, 333) is more new combination uniformly dark in carapace and append- Figures 127, 146-149, 203, 243, ages; the striped form (Figs. 148, 149, 334, 329-337,342; Map 14 335) has much more extensive white mark- ings and distinct markings on the legs. The Dendryphantes exiguus Banks, 1892: 7.5, pi. 5, fig. two forms perfectly discrete: all 30, 9. Holotype in MCZ 19 with labels "Dendry- appear are sorted to phantes exiguus Bks type," "Ithaca, N.Y.," and undamaged specimens easily 282 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

one or the other. However, no clear dif- hanging chelicerae dark. Setae surround- ferences in genitaha have been found, both ing AMEs white laterally 2:00-4:00, often forms having the embolus and spermathe- medially 9:00-10:00; brown otherwise. cae as considered diagnostic for exigua. Chelicerae yellow with front surface The spermathecal ducts may be more con- slightly concave and with black patch on voluted in the dull form, but an adequate inner margin shorter than in flaviceps but and convincing characterization of any dif- nonetheless deep, distinct, and wide; che- ference has proved elusive. The only con- licera lacks prominent basal lateral brown vincing structural difference so far noted spot. Palpus brown, basal segments not dis- is in the width of the dark depression on tinctly paler. Patella, tibia, and cymbium the inner margin of the male's chelicera. lack white scales. Legs more or less uni- The one striped male observed had a formly dark, without distinct longitudinal courtship display as in dull males, to the lines. Abdomen dorsum brown, with weak level of detail observed. Only the dull form side bands. Striped form (Figs. 148, 334): has been seen from some coastal states Carapace with white spots between eyes (New York, New Jersey, North Carolina), of anterior eye row as in flavipedes. Cheek while only the striped form has been seen band thin but long. Clypeus with patch of from Missouri and Illinois. However, the white scales between AMEs, hairs over- two forms occur sympatrically in Virginia, hanging chelicerae white centrally and Maryland, and Arkansas. In Virginia and dark laterally. Setae surrounding AMEs Maryland, they have even been caught white laterally 2:00-4:00, medially 7:00- from conifer trees in the same field. In 11:00; brown otherwise. Chelicerae yellow Massachusetts, there is variation in the with black depression on inner margin nar- amount of white markings but the varia- rower than in dull form. Basal segments tion does not appear to sort itself into two of palpus paler than cymbium. Palpus fe- discrete forms. Until stronger evidence is mur, patella, and tibia with white scales; available to separate the forms, they will cymbium lacking white scales. First leg be kept under the name P. exigua. The femur and patella dark ventrally but with name D. exiguus and apparently also D. white scales dorsally; tibia mostly dark es- virginis apply to the dull form. In the fol- pecially on anterior surface. Other legs pale lowing description the markings of the two with dark markings. Abdomen dorsum forms will be described separately. dark brown, with strong white side bands. Diagnosis. Males of exigua can be dis- Measurements: Body length 4.4(4.5)4.9 tinguished from flavipedes and flaviceps mm; carapace length 1.9(2.0)2.2 mm, = by the mostly dark brown legs, the thin width/length 0.81(0.84)0.84; n 5<5 from retrolateral ramus (Chamberlin, 1925a), Massachusetts, Maryland, Virginia, Ken- and the more twisted prolateral ramus. The tucky, and North Carolina. black spot on the chelicerae is much broad- Female. Epigynum (Figs. 243, 331, 332, er than in flavipedes. Females of exigua 336, 337, 342): Flaps convergent, poste- have more strongly convergent epigynal riorly at about 45° rotation. Epigynal sur- flaps and an extremely broad second curve face flat. Ducts most notable for the very of the internal ducts. broad second curve such that the anterior Male. Palpus (Figs. 203, 330): Embolus edge of this curve begins near anterior end divided deeply into two rami; retrolateral of flap; flowerlike gland openings along ramus thinner than prolateral; prolateral anterior edge of curve. Markings: Dull ramus strongly twisted. Markings: Dull form (Figs. 147, 333): Carapace brown, form (Figs. 146, 329): Carapace relatively covered with transparent brown scales ex- dark, area above first eye row brown. cept for some beige to white scales. Like Cheek band much reduced, to streak be- flavipedes, there are patches of pale scales side ALEs. Clypeus brown, hairs over- behind and between eyes of anterior row; Pelecrina Jumping Spiders • Maddison 283

scales surrounding AMEs dark dorsally. Distribution (Map 14). Eastern United Clypeus densely covered with white scales. States except for far north. Abdomen almost uniform brown, with Records. Many specimens, especially in MCZ and chevrons not so distinct as in flavipedes. AMNH, from: UNITED STATES (county records): Striped form (Figs. 149, 335): Carapace Didl form: MASSACHUSETTS: Barnstable, Essex; often darker than in dull form, covered CONNECTICUT: Fairfield, New Haven; NEW YORK: Nassau, Suffolk, NEW with transparent brown scales except for Tompkins; JERSEY: Burlington, Hunterdon, Ocean; WEST VIRGINIA: some beige to white scales. Behind and Jefferson; VIRGINIA: Augusta, Bedford, Brunswick, between anterior eyes are patches of pale Essex, Fairfax, Norfolk, Page, Shenandoah, Wash- scales; scales surrounding AMEs dark dor- ington; MARYLAND: Caroline, Montgomery, Prince Georges; DISTRICT OF COLUMBIA: Woodridge; sally. Clypeus densely covered with white KENTUCKY: Rowan; TENNESSEE: Unicoi; NORTH scales. with some dark Legs pale markings; CAROLINA: Burke, Craven, Durham, Rockingham first tibia dark as in males. Abdomen dark SOUTH CAROLINA: Oconee; GEORGIA: Thomas ALABAMA: DeKalb, Tuscaloosa; MISSISSIPPI: Scott centrally, with contrasting pale side bands. ARKANSAS: Bradley, Calhoun; TEXAS: San Augus- Measurements: Body length 5.3(5.7)5.8 tine. Striped form: VIRGINIA: Augusta, Essex, mm; carapace length 2.0(2.1)2.2 mm, MARYLAND: - Washington; Georges, Montgomery; width/length 0.79(0.80)0.82; n 59 (dull ALABAMA: Madison; ILLINOIS: Hardin; MISSOU- RI: Boone, ARKANSAS: Form not form) from Rowan Co., Kentucky. Cole; Washington. distinguished: MASSACHUSETTS: Barnstable, Es- (73 observed from Shenan- Courtship sex, Middlesex, Norfolk, Suffolk. doah Co., Virginia; Caroline and Mont- gomery Counties, Maryland; all dull form Natural History. Usually found on co- except one S striped from Montgomery nifers, known from pines (8 records), ju- Co.). With unusual alternate waving of nipers (8 records), occasionally on other palpi during crouch display. Raisedspread plants such as oak (1 record) and walnut = = (n 16, 7<5). Crouch (n 10, 53): Body (2 records). = = low (n 3, 23) or raised (n 1). Male walks in series with long pauses during 1 5 . Pelegrina mon tana = new combination which palpi are waved (n 1), or walk is (Emerton, 1891) more or less continuous with constant pal- Figures 1, 154, 204, 244, 343-347; 16 pus and leg waving and abdomen twitch- Map = First ing (n 2, 13). legs forward and montanus 189L 11. = = Dendryphantes Emerton, Types parallel (n 9, 43), slightly bowed (n 2, in MCZ 2(J 29 with labels "Dendryphantes mon- = in 23) or stretched forward (n 7, 33), with tanus [underlined red], Mt. Washington, N. H." = and "J. H. Emerton Coll.", examined. G. & E. tips almost touching (n 3, 23) or not (n Peckham, 1909: 459, pi. 37, figs. 4, 4a-c, 69. Roew- = 2, First horizontal = 13: 13). legs (n 2, er, 1954: 1213. Bonnet, 1956: 1396. apparently in intense display), or slightly Dendryphantes (Metaphidippus) montanus: — Pe- = = 1911: 636. raised (n 6, 33), to ca. 30° (n 2, 23). trunkevitch, Metaphidippus montanus: —Chickering and Bacorn, Tips of first legs moved side to side (both 1933: 526. 1973: <59. = Kaston, 115, figs. 37, 38, same direction) during leg waving (n 5, = 33), in phase with palpus waving (n 1). Diagnosis. A large, dark northern and = ^ Palpi forward (n 3, 33) and down (n montane species. The embolus that ex- = 1) or slightly raised (n 2, 23). Palpi waved pands near tip and the rough epigynum = = alternately (n 5, 33) up and down (n with ridges behind the flaps are diagnostic. 5, 33) several times at about 2 left and 2 Male. Palpus (Figs. 204, 344): Embolus right palpus waves per second for 2-5 sec narrow near base and flares at tip, almost = = (n 1). Abdomen twitched (n 3, 23) spoon-shaped, concave dorsally. Retrolat- = while palpi waved (n 2, 23). Repertoires: eral ramus reduced. Small denticles cover 23 raisedspread only; 53 raisedspread and the embolus surface basally. Markings crouch. (Figs. 1, 154, 343): Carapace with diffuse 284 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

white side band and white forehead band. in 1949 as montana; the male has the tip Cheek band very weak to absent. Clypeus of both emboli broken (see Cutler, 1979) brown; hairs overhanging cheHcerae dark. and thus resembles montana. White forehead band contacts AMEs dor- Records. Most in MCZ, from: CANADA: YUKON: sally 10:30-1:00. Chelicerae lacking pale Tklo-Klut, 10 km E Old Crow {U 42), King Edward scales. white scales. Ab- Cymbium lacking range N of Old Crow (iS); NEWFOUNDLAND: In- dor- domen more or less uniformly dark dian River (23 19), Terra Nova National Park (1(5); Fox sally, much darker than insignis; anterior QUEBEC: Anticosti, Bay (23 1$), Lake Mistassini, Ayikwapit Peninsula (1

— Metapf dippus capitatus: Bonnet, 1957; 2810, in 0.77(0.77)0.79; n = 56 from New Bruns- part. wick, Massachusetts, Minnesota, and Sas- Notes on Synonymy. The specimen cit- katchewan. ed above be labeled as Female. may incorrectly Epigynum (Figs. 245, 351 , 352): for Banks's short holotype, description and fig- Flaps and divergent or parallel. Sur- ures seem to depict females of proterva face behind flaps raised into bulge only because of the white pubescence (not yel- medially and posteriorly, so that surface as in rises low what we now call "insignis"), gradually behind flaps, or if surface the red-ringed leg segments (not uniform- rises abruptly it does so at some distance ly yellowish), the large reddish spots on behind flaps. Posterior bulge often consid- the abdomen (not smaller and black), and erably higher than level of flaps. Notch the long, parallel epigynal flaps (not short triangular and sharp. First curve of duct and divergent). Nonetheless, since the narrow; second curve proceeds obliquely or specimen marked holotype is clearly of the posteriorly medially. Markings (Figs. species we now call 'Hnsignis," it seems 151, 353): Carapace covered densely with best to leave the matter as it stands and yellowish white scales. Clypeus densely presume that Banks was somewhat erro- covered with yellow scales. Legs more or neous in this description. less uniformly yellow. Abdomen dorsum Diagnosis. Notable for the yellowish with prominent paired black spots, oth- markings with strong black spots on the erwise yellow-brown to red-brown with abdomen. Pelegrina montana and cle- paired spots and lateral markings of yellow mata are similar but the long spatulate and white scales. Measurements: Body embolus and epigynal topography of in- length 3.8(4.1)5.3 mm; carapace length signis are distinctive. 1.7(1.9)2.1 mm, width/length 0.74(0.78) n = Male. Palpus (Figs. 205, 349, 350): Em- 0.79; 59 from Saskatchewan and Min- nesota. bolus flares slightly in distal half; usually bent slightly toward the retrolateral. Ret- Chromosomes. 2nS = 26 acrocentrics + rolateral ramus reduced. The tip of the XXO (is from Taber, Alberta). embolus is very thin and sometimes par- Courtship (26 observed from Taber, Al- tially or entirely broken off (see Cutler, berta, and North Battleford, Saskatche- 1979). Small denticles cover the embolus wan). With triangular leg position during = surface basal to opening. Markings (Figs. crouch display. Raisedspread (n 1). = 150, 348): Carapace often suffused with Crouch (Fig. 128; n 8, 23): Body at low = = brassy scales dorsally, with patches of white (n 5, 1(5) to normal (n 3, 1$) height. = between the fovea and posterior eyes in First legs horizontal and forward (n 8, = addition to ample white side bands and V 2(5), spread slightly (n 3, 1<5) or bowed = mark behind AMEs. Cheek band weak. with tips close (n 5, 1<5). First legs ex- Clypeus brown; hairs overhanging chelic- tended forward on series and waved slight- erae dark. White forehead band contacts ly; on pauses, first legs held back so as to AMEs 10:30-12:30. Chelicerae form a roughly triangular shape as in ver- dorsally = with small patch of yellowish scales me- ecunda and aeneola (n 5, 13), though dially near base. Palpus femur covered with the leading leg is sometimes held in during = = white scales; more distal darker series (n 3, 1<5). Palpi down (n 8, 23) segments = and without white scales. Cym- and tucked in (n 5, 13); waved up and usually = = bium with none to a few white scales. Ab- down (n 8, 23), on series (n 5, 13). domen shows paired black spots of female; Repertoires: 13 crouch only; 13 raised- anterior paired medial white spots distinct; spread and crouch. brassy sheen medially. Measurements: Distribution (Map 17). New Brunswick Body length 3.4(3.6)4.1 mm; carapace west to Alberta, south to New York and length 1.7(1.8)2.1 mm, width/length Colorado. 286 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Records. In AMNH, MCZ, and WPM from: CAN- of dithalea, but the rami are closer to- ADA: NEW BRUNSWICK: nr. ONTAR- Chipman; gether (Figs. 206, 192). The female of IO: Barrie; 5 km S of Richmond near Ottawa; Cam- chaimona is known. SASKATCHEWAN: Wells Lake nr. Alberta poorly bridge; Male. Embolus border; North Battleford; Estevan; ALBERTA: Islay; Palpus (Figs. 206, 355): 8 km W of Writing-on-Stone Provincial Park; Med- rectangular, widening gradually to base on icine Hat; betw. Cereal and Oven; Taber. UNITED retromargin so that there is no distinct an- STATES (county records): MAINE: Hancock, Lin- gle between erect portion and base. Rami coln; VERMONT: Windham; MASSACHUSETTS: small and not much Erect Middlesex, Norfolk, Worcester; CONNECTICUT: separated. por- Fairfield, Hartford, New Haven, New London, Tol- tion of embolus arises more retrolaterally land; NEW YORK: Seneca, Tompkins; NEW JER- than in dithalea. Embolus surface with SEY: Ocean; MICHIGAN: Hillsdale, In- Bergen, small denticles basally. Markings (Fig. gham, Lenawee, Midland, Oakland; WISCONSIN: 354): Cheek band generally weak. Clypeus Dane; ILLINOIS: Champaign; MINNESOTA: Hen- hairs chelicerae dark nepin, Olmsted, Polk, Ramsey; IOWA: Boone; brown; overhanging NORTH DAKOTA: Burleigh, Nelson, Ransom; except white medially. White forehead COLORADO: Fremont. band contacts AMEs dorsally 10:30-12:30. Chelicerae with small medial patch of Natural History. On Chamaedaphne , white scales. with none to a few Betula, and other vegetation in bogs in Cymbium white scales. Measurements: New Brunswick and Ontario; in oldfields Body length 3.9(4.1-4.2)4.3 mm; carapace and prairies in Ontario, Alberta, Minne- length 1.6(2.0)2.1 mm, 0.77(0.78) sota, and Wisconsin. In an open habitat in width/length 0.80; n = 55 from Chiricahua Mtns., Ari- North Battleford, Saskatchewan, P. pro- zona. terva was common and restricted to the Female. 356, 357): taller shrubs (taller than 1 m), whereas Epigynum (Figs. Flaps fairly short, or parallel. P.insignis was common and restricted to convergent Surface raised inside the short shrubs (shorter than 0.5 m) among just flaps midway along their length; behind flaps surface low, the grasses. In habitats in Alberta and Mas- rises into mound sachusetts, P. insignis has also been found prominent posteriorly. First curve of duct narrow; second curve on short shrubs and herbs. In late June near Templeton, Massachusetts, numerous fe- proceeds medially. Markings (Fig. 358): with white scales males were found with egg sacs in nests in Carapace dorsally. Clyp- eus covered with white scales. Ab- living but curled leaves of goldenrod and densely other herbs. domen pale laterally and dark centrally with paired white spots. Measurements: 17. Pelegrina chaimona new species Body length 4.6, 5.0, 54. mm; carapace Figures 206, 354-358; Map 18 length 1.8, 1.8,2.1 mm, width/length 0.78, = — 0.78, 0.79; n 32 from Arizona and Chi- Metaphidippus n. sp. nr. verecundus: Jung and Roth, huahua. 1974: 33 (specimens identified by W. J. Gertsch, With the fe- examined). Male/Female Matching. Holotype male in AMNH with label "ARIZONA: 5 males of chalceola, dithalea, and kastoni mi [8 km] W of Portal, Cochise County, SWRS well associated with males, there remain [Southwestern Research Station], June 9, 1968, V. two males (chaimona and huachuca) and Roth." two females unmatched in southern Ari- Etymology. An arbitrary combination zona. The matching of these is in doubt, of letters, to be treated as an adjective. but the following evidence supports the Diagnosis. Male closely resembles P. matching of the females described above montana, but erect portion of embolus is with the male of chaimona: the flaps of more or less parallel-sided. The embolus is the female are of typical robustness, as shorter and stouter than that of insignis would be expected to match the embolus and has a distinct retrolateral ramus. The of chaimona; the first curve of the epi- embolus bears some resemblance to that gynal duct is narrow like the related in- Pelegrina Jumping Spiders • Maddison 287

signis and montana; the cephalic plate is Embolus lacks the spoonlike dorsal con- somewhat rugose as in males; these females cavity present in the montana group. Te- have only been found in extreme eastern gulum swollen prolaterally into prominent Arizona and south, as have the males. bump. Markings (Figs. 152, 359): Cara- Distribution (Map 18). Southeastern Ar- pace with white side bands and white V izona, Chihuahua, and Nuevo Leon. mark behind AMEs; sometimes with a weak white band. Cheek band Records. UNITED STATES: ARIZONA: Cochise marginal weak. hairs Co.: Chiricahua Mtns.: Southwestern Research Sta- Clypeus brown; overhanging tion 8 km W of Portal (76 12, AMNH), Cottonwood chelicerae dark. White forehead band con- Creek, Rucker Canyon (15, AMNH), Cave Creek tacts AMEs dorsally 10:30-12:30. Chelic- (1<5, AMNH). MEXICO: CHIHUAHUA: Pri- Canyon erae occasionally with a few white scales mavera (13, AMNH), San Jose Babicora (26, AMNH), medially. Palpus femur and distal seg- summit NE of San Jose Babicora (19, AMNH), Ma- ments all with at least dera (12, AMNH); NUEVO LEON: Cerro Potosi (1<5, including cymbium MCZ). some white scales. Abdomen without cen- tral paired white spots; some males show Natural History. Collected from Chrys- paired black spots. Measurements: Body othamnus (IS) and sweeping herbs (13) at length 3.8(4.0)4.2 mm; carapace length elevations from 1,650 to 3,200 m. Males 1.9(2.0)2.2 mm, width/length 0.76(0.77) collected in May (1<5), June (65), July (3<5); 0.79; n = 5(5 from Outlook, Saskatchewan. females collected in July (39). Jung and Female. Epigynum (Figs. 246, 362, 363): Roth (1974) collected this species in their Dark and shiny. Flaps convergent. Entire zone 2 in the Chiricahua Mountains (1,460- area behind flaps raised into a bulge, so 1,700 m elevation). that surface rises abruptly behind flaps. Notch or rounded and of- 1 8. Pelegrina clemata triangular short, ten half the of First curve (Levi & Levi, 1951) new combination only length flaps. of duct second curve medi- Figures 152, 153, 207, 246, narrow; goes 359-364; Map 19 ally. Markings (Figs. 153, 364): Carapace covered with white scales. Clypeus densely clematus Levi and 1951: Metaphidippus Levi, 232, covered with white scales. Abdomen dor- 42 39 be $9. figs. 37, 39, 40, (fig. may insignis), sum brown with white and Holotype $ and paratype 2 in AMNH with label paired spots lateral with black "Metaphidippus clematus Levi 9 S, $ holotype 2 markings; usually paired allotype. Medicine Hat, Alta., Aug. Carr," exam- spots, though not so distinct as insignis. ined. The first three pairs of white spots may be Dendryphantes clematus: — Roewer, 1954: 1209. joined to form two longitudinal bands. Diagnosis. Markings whitish instead of Measurements: Body length 4.7(5.8)5.8 it from Em- mm; carapace length 1.9(2.1)2.1 mm, yellow distinguish insignis. = bolus narrower at tip than in montana, width/length 0.73(0.76)0.79; n 52 from chaimona, and dithalea, much as Outlook, Saskatchewan. insignis, = in baila and chalceola, though these have Chromosomes. 2n3 26 acrocentrics + different markings and a wider carapace XXO (IS from Richfield, Utah). than clemata. The tegulum's prominent Courtship (143 observed from Outlook, bulge is unusual. The dark epigynum with Saskatchewan; Morrin, Alberta; and Piute = a prominent shiny mound behind the flaps Co., Utah). Raisedspread (n 21, 116). = is distinctive. Crouch (n 29, 96): Body at normal height = = Male. Palpus (Figs. 207, 360, 361): Erect (n 8, S6) to low (n 2, 16). First legs = portion of embolus straight, tapering or of forward and horizontal (n 15, 76) or = equal width from base to tip, with small slightly raised (n 8, 36), or raised to 45° = rami at tip; prolateral ramus is more prom- with femur low (n 3, 25); nearly parallel = = inent than retrolateral ramus, which is (n 5, 26), or spread fairly wide (n 4, = small and not projecting retrolaterally. 26). First legs not waved (n 11, 46). Palpi 288 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

= aeneolus: — Pe- down (n 23, 86) and beside chelicerae Dendryphantes (Metaphidippus) = = trunkevitch, 1911: 622. (n 3, 16) or curled under (n 5, 15); Dendryphantes uteanus Chamberlin and Gertsch, flickered series = during (n 23, 85), vig- 1929: 6. in AMNH 1<5 = = 110, figs. 50, 51, Holotype orously (n 5, 13) up and down (n 4, with labels "Dendryphantes uteanus, S holotype," = aeneolus S 2(5), still on pause (n 4, 26). Abdomen "Metaphidippus (Curtis), wlll.n40, = = HOLOTYPE Dendryphantes uteanus Chamb. & twitched (n 15, 56) series (n 9, during "Utah: Lamb's Can 6-10-28 W. and = Gert.", J. G.", 3(5), or perhaps at end of series (n 3, 16), = "28Ff. N40:W1 11, "examined. Roewer, 1954: 1216. still on pause (n 4, 23); twitched contin- Bonnet, 1956: 1402. NEW SYNONYMY. = — uously when walking to mount (n 1). Metaphidippus aeneohis: Chamberlin and Ivie, 1941: 26. 1957: 2809. Repertoires: 3(5 raisedspread only; 23 Bonnet, crouch only; 83 raisedspread and crouch. of the Distribution (Map 19). Western Canada Diagnosis. A common species and United States. western United States and Canada. Males are notable for their dark markings, with Records. In MCZ, AMNH, UWBM, MSU, and WPM reduced white scales on the The from: CANADA: SASKATCHEWAN: Outlook; AL- carapace. BERTA: Medicine Hat; Morrin Recreation Area on rami of the embolus are more widely sep- Red Deer River; BRITISH COLUMBIA: Bull River, arated than in clemata, balia, or chalceola. ca. 115.5° W, 49.5° N; Cranbrook. UNITED STATES Females can be identified by carapace and MONTANA: Sanders; WYOMING: (county records): abdominal markings, the epigynal topog- Carbon, Lincoln, Park, Sheridan, Teton; COLORA- and The sur- DO: Dolores, Gunnison, Huerfano, Lake, Larimer, raphy angled flaps. epigynal Saguache; UTAH: Box Elder, Cache, Millard, Piute, face rises more abruptly behind the flaps Rich, Sevier, Summit, Utah, Weber; IDAHO: Bear than in balia, but not so raised in a bulge Lake, Blaine, Cassia, Fremont, Gem, Onei- Canyon, as in insignis or clemata. The epigynal da, Payette; NEVADA: Elko; NEW MEXICO: Taos; flaps are much more robust than in the WASHINGTON: Grant, Kittitas, Yakima; ORE- from GON: Baker, Deschutes, Grant, Harney, Malheur; sympatric Metaphidippus mannii, CALIFORNIA: Contra Costa, Eldorado, Los Angeles, which aeneola also differs in markings, in- Mono, Placer, Sierra, Siskiyou. cluding having white scales between AMEs. Male. 208, 209, 366-372): Natural History. From sagebrush (Ar- Palpus (Figs. Embolus toward the retrolateral, temisia) in Washington, Oregon, Colora- leaning broadest at tip; two rami widely separated. do, Utah, and Wyoming (10 records). Also 156, 365): Carapace dark taken from field vegetation (Washington), Markings (Figs. and with few or no white scales Purshia (Oregon), Chrysothamnus (Cali- shiny, beneath ALEs and small fornia), Haplopappus, and Sarcobatus except eyes (oc- casional 66 have white side bands, (Utah), 1 record each. though usually sparse). Clypeus brown; hairs over- 19. Petegrina aeneota hanging chelicerae dark brown. Forehead (Curtis, 1892) new combination band absent; setae ringing AMEs white at Figures 156, 157, 208, 209, 247, least laterally 2:00-3:00; otherwise, thin 365-377; Map 21 scales brown to white. Chelicerae lack pale scales. white scales. Ab- Dendryphantes aeneolus Curtis, 1892: 332. Types in Cymbium lacking domen white side bands often faint MCZ 3(5, 62 with labels "Dendryphantes aeneolus dark; Curtis 1892. California. Co-type. S 9" (label is orig- or absent posteriorly. Measurements: Body Elizabeth inal; handwritten, probably by Peckham) length 4.2(4.3)4.7 mm; carapace length and "G. W. Peckham Coll.", examined. Curtis (1892: 2.0(2.1)2.2 mm, width/length 0.75(0.79) 335) implies that the type locality is the San Fran- 0.81; n = 73 from Nevada, and cisco Bay area. G. & E. Peckham, 1909: 468, pi. Oregon,

36, figs. 1-lb, and pi. 38, 6, 6a, <59. Roewer, 1954: Utah. 1205. Female. Epigynum (Figs. 247, 373-376): Banks, 1895: 96. in MCZ Dendryphantes bifida Types Flap with inner edge angled where the 29, 3 pS with labels "Dendryphantes bifida Bks. posterior half of flap bends down into de- type", "Olympia, Wash" and "Nathan Banks Coll.", Surface rises almost im- examined. This type series is incomplete, for Banks pression. abruptly, described the adult male. mediately posterior to the flaps, but pos- Pelegrina Jumping Spiders • Maddison 289

terior portion of epigynum is fairly flat. Courtship (93 observed from Yakima Notch often triangular with a sharp an- and Kittitas Counties, Washington, and terior point, but many 99 have rounded Riverside Co., California). First legs make notch. First curve of duct narrow; second shape during crouch display, as triangular = curve proceeds medially. Markings (Figs. in P. verecunda. Raisedspread (n 21, = = 157, 377): Carapace only thinly covered 53). Crouch (n 22, 83): Body low (n = with white scales; bronze scales also es- 6, 33). First legs horizontal (n 17, 73) but pecially on cephalic plate. Most females femora held back and to sides and distal with inverted T marking on cephalic plate segments pointed forward and with tarsi consisting of white band of scales starting nearly touching so as to make a triangle = between AMEs, proceeding posteriorly, shape (n 12, 53), though occasionally then spreading laterally to behind small femora held forward and tips not touching = eyes. Otherwise, bronze behind AMEs and (n 3, 13). First legs flickered on series (n = = ALEs, and bronze between posterior eyes. 9, 43), but only slightly (n 3, 13) or == Clypeus densely covered with white scales, not waved (n 7, 33); also flickered when = paler between AMEs than in balia and very close (n 6, 43) at which time legs = = mannii, which have orange scales. Abdo- extended (n 4, 33). Palpi tucked in (n = men white markings usually small except 6, 23), flickering on series (n 6, 23) or = often large central pale spot. Background when male very close (n 2, 23). Reper- dark, often bronze or gray, occasionally toires: 13 raisedspread only; 43 crouch only; with paired dark spots on either side of 43 raisedspread and crouch. the midline. Measurements: Body length Distribution (Map 21). Western United 4.5(4.8)6.1 mm; carapace length 2.0(2.1)2.3 States, extending into Canada and Mexico. = mm, width/length 0.77(0.79)0.85; n 79 from and Arizona. Records. Many specimens, especially in MCZ, Oregon AMNH, UWBM,'and UCB, from: CANADA: BRIT- Variation. In western SS Geographical ISH COLUMBIA: Alice Lake Province Park; Creston; (aeneola proper, British Columbia, Wash- Furry Creek; Massett; Victoria; Wellington, Quali- ALBERTA: Waterton Lakes National ington, Oregon, California), embolus nar- cum, Nanaimo; Park. UNITED STATES (county records): SOUTH row with prolateral face gently curved or DAKOTA: Custer, Horsethief, Jackson, Pennington; bent in eastern 33 slightly (Fig. 208); (form MONTANA: Flathead, Lewis and Clark, Park; IDA- uteanus. South Dakota, Montana, Wyo- HO: Bear Lake, Boise, Bonner, Franklin, Latah, ming, Colorado, Utah, New Mexico, Ari- Oneida, Payette, Washington; WYOMING: Sheri- COLORADO: Custer, Fre- zona), embolus is wider with prominent dan; Alamosa, Douglas, mont, Hinsdale, Juab, La Plata, Larimer, Logan, on prolateral face just basal to the angle Montezuma, Utah; UTAH: Box Elder, Cache, Davis, and rami more opening, divergent (Fig. Grand, Juab, Piute, Salt Lake, Uinta, Weber; NE- 209). The few specimens available from VADA: Washoe; NEW MEXICO: Bernalillo, Lin- the intermediate area (Idaho, Nevada, and coln, Otero, Rio Arriba, Sandoval, San Miguel, Santa Torrance, Valencia; ARIZONA: southeastern California) and occasional Fe, Taos, Apache, Cochise, Coconino, Graham, Mohave, Yavapai; from show some inter- specimens Oregon WASHINGTON: Asotin, Chelan, Clallam, Columbia, differences in 63 and gradation. No other Douglas, Grant, Grays Harbor, Jefferson, King, Kit- none in 99 have been found between west- titas, Pierce, San Juan, Skagit, Skamania, Snohomish, Walla OREGON: ern and eastern populations; hence, 1 con- Stevens, Thurston, Walla, Yakima; Benton, Deschutes, Douglas, Grant, Harney, Jackson, sider them conspecific. SS from the Colum- Jefferson, Josephine, Klamath, Lake, Lane, Marion, of from south- bia basin Washington, some Multinomah, Umatilla, Union, Wallowa; CALIFOR- ern California, and occasional males from NIA: Alameda, Contra Costa, El Dorado, Kern, Las- Oregon and northern California have ex- sen, Los Angeles, Marin, Mendocino, Modoc, Mon- Nevada, Plumas, Riverside, San Ber- tensive but not very dense white side bands terey, Orange, nardino, San San Mateo, Santa Barbara, Santa on the Diego, carapace. Clara, Santa Cruz, Santa Cruz Island, Shasta, Sierra, = acrocentrics + Chromosomes. 2n6 26 Siskiyou, Trinity, Tulare, Ventura, Yuba. MEXICO: XXO (23 from Apple Canyon, Riverside BAJA CALIFORNIA DEL NORTE: Parque Nacion- al Sierra San Pedro Martin. Co., California). 290 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Natural History. Habitat: In Washing- Etymology. After the Greek adjective ton, collected from pines including Pinus balios, meaning "dappled" (Woods, 1966). ponderosa, understory of riparian poplar Diagnosis. A large western species with woodland, understory ferns, Ceanothus, light brown and beige markings. The male alders in bog, lakeside marsh, houses; at can be identified by the broad side bands elevations of less than 50 m to more than of the carapace and the flange on the fang. 1,700 m. More than 15 adults males and The embolus is narrow, very similar to that females were recovered by D. H. Mann of chalceola, but it leans slightly and the and others on snow at 1,700-3,000 m, ap- erect portion broadens gradually into base. parently having ballooned (Pierce Co., The tibial apophysis of balia usually points Washington). In Oregon, commonly col- more distally than in chalceola. The fe- lected from Abies grandis and Pinus pon- male can be identified by the spotted ab- derosa; also from Pseudotsuga, Picea, Pi- domen and the epigynum, which differs nus contorta, hemlock, alder, Salix, brack- from that of the sympatric aeneola in hav- en fern, Calocedrus, Taxus, oak, Ceano- ing the surface rise gradually behind the thus, and Larix occidentalis. Curtis (1892) flaps. reported it common in San Francisco area Male. Palpus (Figs. 210, 379): Embolus gardens on honeysuckle, rose bushes, live narrow and tall, leaning slightly to retro- oaks, and Laurestina. In Nevada, beating lateral; erect portion broadens gradually pinyon pine; in Arizona on pine. Life cy- into base. Retrolateral ramus points retro- cle: In Oregon's Malheur National Forest, laterally. Retrolateral edge of embolic base B. Fichter and A. Moldenke collected 44<5 with membranous fold, unlike chalceola. 779 from 12 to 17 June 1982, 66 252 from Chelicerae robust and divergent; fang with 18 to 23 July 1982, and 819 from 20 to 29 pronounced flange on cutting edge (Fig. September 1982. In the San Francisco area, 378, arrow). Markings (Fig. 378): unlike "males and females appear as adults as most other Pelegrina males in having early as April, but the former become rare markings more spotted than striped. Car- after the first of June and the latter after apace with distinctive, broad creamy white the first of September. The females begin side bands. Cheek band weak and broad. laying eggs in May" (Curtis, 1892: 335). Clypeus brown; hairs overhanging chelic- Curtis reported one or two egg sacs per erae dark with some white medially. White females with about 50 eggs that hatched forehead band fails to contact AMEs dor- on average in 25 days. In Los Angeles sally, which are ringed by dark or at most County, California, the 203(3 examined by thin white setae. Chelicerae lacking pale me were collected from November through scales. Cymbium lacking white scales. Ab- June; the 2199 from February through June domen either mottled as in female, with plus two in September. Behavior: Curtis conspicuous basal band and second lateral (1892) described the spider's entrance to bar, or with side spots fused into very wide and defense of its retreat, its ballooning, cream side bands. Measurements: Body and its reaction to a sluggish pet lizard. length 3.8(4.3)5.1 mm; carapace length Land (1969a, b, 1971) investigated visual 1.8(2.1-2.3)2.3 mm, width/length 0.79 = behavior and eye structure and function (0.81-0.82)0.85; n 66 from California, of P. aeneola. Oregon, and Washington. Female. Epigynum (Figs. 248, 380, 381): 20. Pelegrina balia new species Flaps usually light brown, convergent, surface Figures 210, 248, 378-382; Map 20 though sometimes dark. Epigynal mostly concave except or bump just medial Holotype male and paratype female in UCB with to flaps; surface rises behind flaps gradu- label "CA[lifornia]: S[an]ta. Barbara Co., Ballinger to a medial and First Cyn., 17 mi, [27 km] SE. New Cuyama, el. 3000' ally posterior bulge. [915 m], V-9-1980, C. E. Griswold." curve of duct pale; second curve goes me- Pelegrina Jumping Spiders • Maddison 291

dially; third curve smooth on inner sur- Etymology. An arbitrary combination face. Markings (Fig. 382): Carapace cov- of letters designed to resemble the name ered densely with yellow-white scales, of- of the similar species P. aeneola both in ten with dark streak on thorax side. Clyp- structure and in referring to the bronze eus densely covered with white, between color (Greek, chalceos). AMEs orange or tan (white in some Ore- Diagnosis. A dark, shiny southwestern gon 99). Abdomen mottled with large pale species resembling aeneola and balia. The spots: with pale basal band fused to first narrow, tall embolus is much like that of lateral bar; second oblique bar swollen; balia, from which chalceola differs by the posterior lateral bars inconspicuous; cen- lack of the pronounced flange on the male tral pale spots round, edge or connected fang, and the much darker body with a with dark brown. Measurements: Body bronze sheen in both males and females. length 4.7(5.2)6.1 mm; carapace length Male. Palpus (Figs. 211,384): Erect por- 2.1(2.2)2.3 mm, width/length 0.79(0.80) tion of embolus narrow and tall, straight, = 0.81; n 52 from California. usually broadens abruptly into embolic Male/Female Matching. This match- base so as to leave angle between erect ing is indicated by the similarity in robust portion and base. Retrolateral ramus points carapace and mottled markings and by co- retrolaterally. Retrolateral edge of embolic collecting. base is simple and schlerotized, lacking the Distribution (Map 20). California, north fold seen in balia. Markings (Figs. 139, to Washington and east to Arizona and 383): Carapace dark, with narrow white Colorado. side bands often reduced behind posterior eyes. Cephalic area with transparent Records. About 70 specimens in AMNH, UCB, and bronze scales. Forehead lacks white band, MCZ from: UNITED STATES (county records): in is WASHINGTON: Spokane; OREGON: Baker, Des- though some males there small patch chutes, Harney, Jackson, Klamath, Lane, Wheeler; of pale scales between and behind AMEs. CALIFORNIA: El Dorado, Fresno, Inyo, Kern, Las- Cheek band weak to almost absent. Clyp- sen, Los Mendocino, Modoc, Mono, Angeles, Mariposa, eus brown; hairs overhanging chelicerae Plumas, Riverside, San Bernardino, Santa Barbara, white and tan laterally, to all Shasta, Siskiyou, Sonoma, Stanislaus, Trinity, Ven- centrally tura; COLORADO: Mesa; ARIZONA: Yavapai. brown. Setae ringing AMEs brown dor- sally. Chelicerae robust but vertical, with Natural History. On occi- Juniperus none to a few pale scales medially. Cym- dentalis (3 records), Pseudotsuga (2 re- bium lacking pale scales. Third leg dark cords), macnabiana (1 record), Cupressus on distal % to entirely dark. Abdomen dor- Abies (1 record), and Calocedrus (1 rec- sally brown with paired black spots and in and northern California. ord) Oregon thin white side bands; third and fourth From juniper woodland in California (5 pairs of white spots when present are lat- records). Over the entire range, 35 were erally directed bars. Measurements: Body collected in April, 10 in May, 3 in June length 3.9(4.0)4.9 mm; carapace length and 1 in September. 1.8(2.1)2.3 mm, width/length 0.80(0.82) 0.83; n = 53 from Arizona.

21 . chalceola new Pelegrina species Female. Epigynum (Figs. 385, 386): 20 Figures 139, 211, 383-387; Map Flaps short, slightly convergent, posteri- orly in concavity. Epigynal surface gently Metaphidipptis n, sp. nr. montanus: —Jung and Roth, convex behind Second curve of duct 1974: 33 (specimens identified by W. J. Gertsch, flaps. examined). goes medially; third curve with rough in- male and several immatures in MCZ with Holotype ner surface. Markings (Fig. 387): Brown label "ARIZONA: Santa Cruz Co., Madera upper with bronze sheen. Carapace covered with Canyon, Santa Rita Mtns., ca. 5500 ft [1,680 m], 13 reflective white to tan scales. of Aug' 1983. W. Maddison 83-158 oak v^'oodland, Spots pale beating oaks." scales between anterior eyes similar to fla- 292 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

24 km S of Prairie Grove (13, MCZ); ILLINOIS: Har- vipedes. Clypeus densely covered with din (9). white scales. AME eye ring darkest dor- sally, tan to brown, as in male. Abdomen Natural History. Collected beating oaks Measure- with large paired dark spots. in oak woodland (1<3, Arizona) and from 4.7 ments: Body length 4.4, 4.4, 4.7, mm; juniper (13, Texas) at altitudes from 300 carapace length 2.1, 2.2, 2.3, 2.3 mm, m to 1,650 m = (Arkansas) (Arizona). Jung width/length 0.78, 0.78, 0.81; n 42 from and Roth (1974) collected this species in Arizona. their zones 1 and 2 in the Chiricahua Male/Female Matching. Among Ari- Mountains (1,200-1,700 m). zonan species, the female and male share a distinctive wide box-shaped carapace, 22. Pelegrina furcata dark bronzed appearance, and markings (F. P.-Cambridge, 1901) on abdomen. The AME ring dark above new combination and light elsewhere also unites male and Figures 158, 159, 212, 249, 250, female. The underside of the abdomen is 388-402; Map 22 also fairly pale, distinguishing it from the P.P. 1901: 267, female of huachuca. Metaphidippus furcatus -Cambridge, pi. 24, figs. 8, 8a, 3. Type material in BMNH 23 Variation. Two male Pe- Geographical with label "Dendryphantes furcatus, sp. n. m's, Ori- legrina from Durango tentatively identi- zaba, Mexico (H. S. )" and 23 with label "Dendry- n. Gua- fied as chalceola differ from specimens phantes furcatus, sp. Type 3, Syntype 3., temala (Sarg)," examined. Despite the type label from the United States in being large (body on the latter specimens, the holotype may be better 5.2,5.2 2.5,2.5 length mm; carapace length considered to be among the former, given Cam- mm, width/length 0.82,0.83), with em- bridge's indication of the distribution as Orizaba. bolus shorter and leaning more to the ret- Bonnet, 1957: 2813. —G. & E. Peckham, 1909: rolateral in the distal half, and with a bump Dendryphantes furcatus: 473. Roewer, 1954: 1203. on the side of the chelicera almost as in Dendryphantes mimus Chamberlin, 1925b: 135, figs. the mannii but not so well devel- group 53, 54, 3. Holotype in MCZ 13 with label "Den- oped. dryphantes mimus Chamb., 3 holotype, N. M.: Pe- Courtship (IS observed from Madera cos, R. V. Chamberlin Coll. 1047," examined. = Roewer, 1954: 1212, Bonnet, 1956: 1396. NEW Canyon, Arizona). Raisedspread (n 6): = SYNONYMY. Carapace high (n 6); abdomen down (n = = 6) and trailing (n 2). First legs raised, Diagnosis. A species common in the = forward, spread (n 6), but legs moved Mexican highlands, with a striking court- = to more parallel as he got closer (n 4), ship display and distinctive embolus hav- = waved little if at all (n 6). Palpi down, ing two blunt rami. The epigynum, with = waving little if at all (n 6). Male pro- convex flaps, concave surface, and wide ceeded directly from raisedspread to second curve of the ducts, is distinctive. reaching to touch the female, without go- Male. Palpus (Figs. 212, 389-394): Em- = ing through a crouch display (n 4). bolus heavy and slanting, with retrolateral Distribution (Map 20). Southern Ari- ramus extended and truncate. Markings zona east to southern Illinois. (Figs. 158, 88): dark brown with distinct sheen and contrasting white side bands. Carapace side bands usually connect to Records. UNITED STATES: ARIZONA: Cochise white scales over anterior row to make Co.: Chiricahua Mtns., Southwestern Research Sta- eye of white the tion (83, AMNH); Chiricahua Mtns. (19, AMNH); Hu- a continuous band encircling achuca Mtns., Montezuma Pass (13, AMNH); Santa front of the carapace (though not seen in Cruz Co.: Santa Rita Mtns., Madera MCZ Canyon (1<5, male drawn. Fig. 388). Cheek band mod- IS, AMNH), Santa Rita Mtns. (29, AMNH); TEXAS erately weak. Clypeus brown; hairs over- Denton Co.: Lake Dallas opposite Hatchery (13, MCZ) chelicerae brown to white me- Erath Co.: Stephenville (33, TXAM); ARKANSAS hanging Washington Co.: Boston Mtns., Cove Creek Valley, dially, brown laterally. White forehead Pelecrina Jumping Spiders • Maddison 293

band contacts AMEs dorsally 10:30-12:30 lateral ramus of the embolus is truncated or 1:00. Chelicerae with some pale scales transversely, so that its distal tip makes a medially. Cymbium lacking pale scales. line perpendicular to the axis of the palpus. Measurements: Southern Arizona: Body Females are dark. (3) A third form occurs length 3.5(4.2)4.5 mm; carapace length in northern Arizona (Yavapai Co.), Colo- 1.7(2.1)2.2 mm, width/length 0.74(0.77) rado, and New Mexico, having a wider = 0.77; n 5(5 from Santa Rita Mtns., Ari- embolus and convergent flaps (mimus: zona. Northern Arizona: body length 3.5, Figs. 390, 391, 395, 396). The retrolateral 3.6, 3.9 mm; carapace length 1.7, 1.8, 1.8 ramus of the embolus is truncated trans- = mm, width/length 0.75, 0.78, 0.79; n 33 versely, as in form (2). The difference be- from Yavapai Co., Arizona. tween the northern (mimus) and southern Female. Epigynum (Figs. 249, 250, 395, (jurcata) Arizona specimens is rather strik- 397, 398, 400, 401): Flaps strongly convex ing, for the females are also smaller and and often dark. Surface concave behind paler in the north. Though mimus might flaps, without mound, rising gradually to be considered a distinct species, specimens lip at back edge. First curve of duct nar- in New Mexico present a confusing mix- row; second curve broad initially but nar- ture of characteristics of forms (1), (2), and rows as it proceeds medially. Markings (3). (4) A fourth form is found in western (Figs. 159, 396, 399, 402): Body often with Oaxaca (43 159, 31 km N or Guelatao de slight bronze sheen; variable in markings. Juarez, ca. 96.5°W, 17.5°N, 2,600 m el., 3 Carapace covered with brassy reflective August 1983, W. Maddison & R. S. An- scales, sometimes dark, sometimes mixed derson, MCZ), with very wide embolus, with white. Clypeus only thinly covered dark females, and extremely robust flaps with white scales except in northernmost (Figs. 388, 400-402). This form occurs populations (form mimus). Measure- within 50 km of the widespread form. In ments: Southern Arizona: Body length total, the variation among these popula- 4.5(4.7-4.9)5.4 mm; carapace length tions is confusing, and though several spe- 2.0(2.1)2.2 mm, width/length 0.75(0.78- cies may be present, only one will be rec- = 0.79)0.79; n 62 from Santa Rita Mtns., ognized until better studied. = Arizona. Northern Arizona: Body length Chromosomes. 2n3 26 acrocentrics + 4.0(4.6)5.7 mm; carapace length 1.8(1.8)2.0 XXO (2(5 from Madera Canyon, Arizona). = mm, width/length 0.72(0.75)0.77; n 59 Courtship (10(5 observed from Nuevo from Yavapai Co., Arizona. Leon, Hidalgo, Queretaro, Puebla, Oaxa- Geographical Variation. Four geo- ca, Chiapas, and the Santa Rita Mountains graphical forms might be recognized. (1) of Arizona). Very unusual for the genus, The most widespread form occurs from with vigorous leg waving and body jerking Guatemala north to northern Mexico, with in a stage I will call the semaphore stage. = narrower embolus and thinner epigynal Semaphore (n 24, 103): Body high to = flaps that are divergent or parallel (Jurcata very high (n 24, 103). Male walked si- = = s.s.,; Fig. 393). The retrolateral ramus of dhng (n 19, 83) in series (n 14, 53). the embolus is truncated Most First legs wide, nearly 180° apart, approx- obliquely. = females through this range are well marked imately horizontal (n 21, 83) or below = with pale spots, as in form mimus (Fig. horizontal (n 1), though occasionally not 396). (2) A second form, very similar to much more than 90° apart and raised to = the widespread form, occurs in the Santa 60° (n 1), waved vigorously up and down = Rita, Santa Catalina, and Chiricahua almost to vertical (n 21, 93), though = Mountains of southern Arizona and prob- sometimes only to ca. 40° (n 1), at ca. = = ably in northern Mexico 392, 397- 3-4 c/s (n 2, 23) or 5 c/s (n 1) on each (Figs. = 399). The embolus is also narrow and the sidle (n 9, 43). The leg wave is vertical flaps divergent or parallel, but the retro- and slightly posterior to bring the legs up 294 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

CHIHUAHUA: Canon near Pri- and back (n =7, 45); the left and right sides rery (13); Prieta, = mavera (19); HIDALGO: 4 km NE of Tlanchinol (23 wave in unison (n 4, 36), or occasionally = 69); 3.4 km SW of Cuesta Colorada (43 29); 10 km asynchronously (n 1). Palpi hanging SW of Santa Maria, 21°06'N 8 km N = = 99°00"W, (13); down (n 18, 83) and parallel (n 9, 46) of Encarnacion, 99.12°W, 20.55°N (29); Huachinango = and a bit forward (n 7, 3(3). Palpi wave (19); Apulco (13 29); Champuhuacan (19); Maguey = Verde, 99,12°W, 20.49°N; PUEBLA: 8 km N of Te- with low-medium amplitude (n 5, S6) ziutlan (13 39); near Xicotepec de Juarez, 97°59'W, on each sidle = 4, and down like (n 26), up 20"'17'N 5 km N of 130 on road to = (13) Hvw Naupan pushing and pulling motion (n 1), or ca. 21°15'N = (13); QUERETARO: 99°10'W, (53 39); largely still (n 1, 26). Abdomen trails a GUERRERO: Chilapa (19); VERACRUZ: 3 km N of = = Fortin de las Flores 6 km NE of bit on sidles (n 5, 26), ca. 10-30° (n 3, (13); Coscomatepec = (23); 7 km N of Huatusco (13 39); Orizaba (3399); 1(5), but more or less horizontal (n 16, DISTRITO FEDERAL: Santa Rose (13), Contreras 6 from 56). Occasionally pauses vigorous OAXACA: 23 km SW of Valle Nacional on = (13); Hwy leg waving and jerks whole body (n 11, 175 (43 59); 27 km SW of Valle Nacional on Hwy = 175 31 of Guelatao de 66) approximately 4-5 times (n 3, 3(3) or (13 29); km N Juarez (43 159); = CHIAPAS: 5 km W of San Cristobal de las Casas (33 3 times (n 1) while the first legs are = 39); Grutas de San Cristobal (19); San Cristobal (43 spread wide and horizontal (n 11, 66). = 49). GUATEMALA: locality unknown (23). These jerks came after a few sidles (n Natural Collected from oak 1). The body may be lowered for the jerks History. (7 = = shrubs in (n 1). Reach (n 3, 23): The male pro- records), grasses, herbs, and ceeded directly from this semaphore stage clearings (5 records), pine (3 records), ju- = into the reach to touch the female (n 3, niper (1 record), Ceanothus (1 record), in cloud 23). During reach, body jerked a few times Cercocarpus (1 record), oak-pine = forest zones. at elevations of (n 1). First legs held parallel and forward Collected = (n 2, 23). Palpi held parallel and forward 1,000-1,400 m (8 records, 1,500-2,000 m = (n 2, 23). (7 records), and 2,100-3,000 m (7 records). This description is from the widespread form (1) with the following exceptions. 23. Pelegrina volcana new species Displays of 43 from the distinctive popu- Figures 213, 403, 404; Map 23 lation from Guelatao de Juarez, Oaxaca Holotype male in MCZ with labels: "PANAMA: El (form (4)), showed the same form of sem- Volcan, A. M Chickering" and "R. P. El Volcan, aphore display with legs spread wide, vig- 1950." = Aug. 9-14, orously waved up during series (n 11, An arbitrary combination 43). No whole-body jerks were noted, how- Etymology. of letters to the to ever. One male from southern Arizona referring type locality, be treated as an adjective. (form (2)) showed the same semaphore dis- = Known from only two males play (n 4), though no whole-body jerks Diagnosis. were noted. from Panama; much like bicuspidata but the embolus is not so bent as in that Distribution (Map 22). Throughout the long highlands of Mexico, extending north to species. Erect Colorado and south to Guatemala. Male. Palpus (Figs. 213, 404): por- tion of embolus broadens gradually into base; rami small and subequal. Tibial Records. in MCZ and AMNH: UNITED Mostly apophysis appears double because ridge STATES (county records): COLORADO: Boulder into second more ex- (46), El Paso (!

tacts AMEs dorsally 10:30-12:30. Chelic- Chiapas 3.) Measurements: Body length erae with long medial patch running al- 2.7, 3.2 mm; carapace length 1.3, 1.5 mm, = most at least % length. Palpus femur dis- width/length 0.77, 0.83; n 23 from Chia- tinctly paler than more distal segments. pas and Guatemala. Cymbium with none or few white scales. Female. None matched to male. Cam- Legs fairly distinctly annulate. Anterior bridge's P. ochracea may represent the fe- three pairs of abdominal spots longitudi- male of P. bicuspidata. nally directed, 4 through 6 transverse. Strong abdominal side bands. Measure- Records (Map 23). In addition to the holotype, one ments: Body length 3.5, 3.7 mm; carapace other male is known, from Mexico; Chiapas: pine forest, 24 km NW of Arriaga 94.0rW, 16.25°N, 27 length 1.7, 1.8 mm, width/length 0.76, August 1966 (AMNH). 0.77; n = 2(5 from Panama.

25. ochracea 24. Pelegrina bicuspidata Pelegrina (F. P.-Cambridge, 1901) (F. P.-Cambridge, 1901) new combination new combination 407-409; 23 Figures 214, 405, 406; Map 23 Figures Map Metaphidippus ochraceus F. P.-Cambridge, 1901: Metaphidippus bicuspidatus F. P.-Cambridge, 1901: 272, pi. 25, figs. 6, 6a, 2. Holotype in BMNH 12 269, pi. 24, figs. 13, 13a, b, <5. Holotype in BMNH with label "Dendryphantes ochraceus, sp. n. Type 1(5 with labels "Dendryphantes bicuspidatus, sp. 2, Guat. (Sarg)," examined. Bonnet, 1957: 2816. Type (3, Guatemala (Sarg)" and "1905, 268.", ex- Chickering's M. ochraceus (1946: 312) is not the amined. Bonnet, 1957: 2810. same as this species, nor is it a Pelegrina. Dendryphantes bicuspidatus: —Roewer, 1954: 1191. Dendryphantes ochraceus: —Roewer, 1954: 1198.

Diagnosis. A rarely collected species from southern Mexico and Guatemala. The Diagnosis. The epigynum is similar to that of P. furcata, but the flaps are not distinctive embolus is long and bent, unlike so convex and are shorter. As that of volcana. quite already noted, this could be the female of P. bi- Male. Palpus (Figs. 214, 406): Embolus cuspidata. heavy, abruptly bends basal to opening. Female. 407, 408): Rami small and subequal. Tibial apophysis Epigynum (Figs. Flaps convex, parallel, and fairly short; appears double because ridge prolonged light brown; behind them the surface is into second apophysis, more extreme than gently concave with mound restricted to in furcata (Fig. 389). Markings (Fig. 405): near posterior margin, and median ridge Typical for the genus, with white cheek, from to mound. forehead, and side bands. Cheek band extending flaps posterior First curve of duct narrow; second curve moderately dense. Clypeus brown; hairs proceeds medially. Markings (Fig. 409): overhanging chelicerae white medially, Carapace red-brown, thinly covered with brown laterally. White forehead band con- white scales. Clypeus covered with white tacts AMEs dorsally 10:00-12:30. Chelic- scales, scales surrounding AMEs white. erae with thin patch of pale white scales Legs lacking strong annulae, tan to light medially extending to % length. Palpus fe- brown. Abdomen light brown with pale mur distinctly paler than more distal seg- markings (Fig. 409). Measurements: Ho- ments. Cymbium lacking white scales. Legs lotype body length 4.0 mm; carapace with fairly distinct annulation; back of tib- length 1.6 mm, width/length 0.75. ia 2 uniformly pale; femur 3 dark in distal %. Abdomen dorsum more or less solid Records (Map 23). MEXICO: OAXACA: Oaxaca, brown, without trace of dark spots, sur- Base San Felipe Mtn., 16-17 September 1947 (12, on rounded by discrete white side bands AMNH); CHIAPAS: San Crist6bal, 13 September 1947 abdomen. (Description based mostly on (12, AMNH). GUATEMALA (12, BMNH). 296 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

26. Petegrina morelos new species markings with strong lateral fourth pair of Figures 215, 410-414; Map 23 spots, and they occur sympatrically.

Holotype male in AMNH with label "7 mi [11 km] Records (Map 23). In addition to the male holotype, N Cuernavaca, Morelos, Mexico, July 3, 1941, A M the single female known is from: Mexico: Morelos: " and L I Davis. Cuernavaca, July 1953 (AMNH).

A noun in after Etymology. apposition, 27. Petegrina huachuca new species the type locality. Figures 216, 415-419; Map 24 Diagnosis. Much like furcata, but the male in with label retrolateral ramus of the embolus is pro- Holotype AMNH "ARIZONA: 8000 ft. [2,440 m], Carr Canyon, Huachuca Mts., longed and curves to the prolateral. The June 3, 1952." female matched to the holotype has much more contrasting markings than in furcata Etymology. A noun in apposition, after and epigynal ducts with a much narrower the type locality. second curve. Diagnosis. Most distinctive for its large Male. Palpus (Figs. 215, 411): Embolus branched embolus bearing some resem- with two blunt rami much like those of blance to that of P. furcata. Females furcata, but the retrolateral is long and matched with the male have long flaps in curves to the prolateral. Tibial apophysis a distinctively sculptured epigynum. appears double because ridge prolonged Male. Palpus (Figs. 216, 416): Embolus into second apophysis. Markings (Fig. 410): large and unusual, with retrolateral ramus Carapace side bands dense and discrete. extended retrolaterally as a long blade. Cheek bands weak. Clypeus brown; setae Tibial apophysis appears double because overhanging chelicerae dark except a few ridge prolonged into second apophysis. white hairs medially. White forehead band Markings: Typical for the genus, with contacts AMEs dorsally 10:30-12:30. Che- cheek, side, and forehead bands on the licerae with a few white scales medially. carapace and side bands on the abdomen. Palpus cymbium brown, lacks white scales. Cheek band moderately dense. Clypeus Abdomen with white side bands and paired brown; hairs overhanging chelicerae dark white spots reminiscent of female. Mea- with some white centrally. White forehead surements: Body length 38 mm; carapace band contacts AMEs dorsally. Chelicerae length 1.9 mm, width/length 0.78. with a few pale scales medially. Legs with Female. Epigynum (Figs. 412, 413): indistinct annulation. Measurements: Body Flaps long and convex, dark. Epigynal sur- length 3.7 mm; carapace length 1.8 mm, = face slightly concave, rises gradually be- width/length 0.77; n 16 from Huachuca hind flaps to higher posterior margin. First Mtns., Arizona. curve of duct broad; second curve narrow, Female. Epigynum (Figs. 417, 418): proceeds medially. Markings (Fig. 414): Flaps dark, long, and parallel. Surface Carapace brown, thinly covered with white bulges just medial to flaps about half way scales. Clypeus covered with white scales along their length; behind this the surface though scales surrounding AMEs are or- is concave, rising gradually into pro- ange laterally and medially. Legs annu- nounced medial and posterior bulge. First late. Abdomen strongly marked with four curve of ducts very broad; second curve distinct pairs of white spots on dark back- goesanteriomedially. Markings (Fig. 419): ground; fourth pair large and transverse. Carapace covered with white scales with Measurements: Body length 4.7 mm; car- some light brown. Clypeus densely white; apace length 1.9 mm, width/length 0.80. white between AMEs. AME scales yellow- Male/Female Matching. The female ish white above, white below. Abdomen described is matched tentatively with the brown; central pale spots are laterally di- male, for both are similar to furcata in rected bars, side bands. Measurements: genitalia, they have similar abdominal Body length 4.5, 5.1 mm; carapace length Pelegrina Jumping Spiders • Maddison 297

= arizonensis:— Pe- 2.1, 2.2 mm, width/length 0.79, 0.80; n Dendryphantes (Metaphidippus) 1911; 622. 29 from Santa Catalina and Santa Rita trunkevitch, Dendryphantes mimus:—Chamberlin, 1925b, in part; Mtns., Arizona. 135, fig. 52 9. Male/Female Matching. As discussed Metaphidippus arizonensis: —Bonnet, 1957; 2810. Cutler and 1985; 69. under P. chaimona, there is doubt regard- Jennings, 3, figs. 3-11, Metaphidippus glacialis: —Bonnet, 1957; 2814. ing the male/female association of chai- mona and huachuca. The following evi- Like helenae this has dence supports the matching of the fe- Diagnosis. species unusual for the with the males already described with the male of genitalia genus, erect portion of the embolus arising retro- huachuca: the flaps of the female are long and the far rotated. and convex, the surface has distinct con- laterally epigynal flaps Differs from helenae in a sharp cavities and bulges, and the first curve of having pointed embolus, short tibial apophysis, and the duct is very broad, as would be ex- rotated 180°. pected to match the robust embolus of hu- flaps only Male. Em- the is smoother than Palpus (Figs. 217, 421, 422): achuca ; cephalic plate bolus toward retrolateral in chaimona males and females, as in the arising side, and with surface male huachuca; the females have been blade-shaped exposed concave, with retrolateral found in central Arizona, as was the male. ridge extending into distal point. Tibial apophysis almost Distribution (Map 24). Southcentral and hidden behind wide beneath it. southeastern Arizona. flange Markings (Figs. 160, 420): Cheek bands face variable. Records. UNITED STATES: ARIZONA; Cochise weak. Markings on quite Co.; Huachuca Mtns., Garden Canyon Road, base of Clypeus brown, sometimes with a few Sawmill Canyon, 19 March 1989 (36, MCZ), Hu- white hairs, hairs overhanging chelicerae achuca Mtns., Carr Canyon, 3 June 1952, 2,400 m el. white to brown. White forehead band con- (is, AMNH), Chiricahua Mtns., upper Cave Creek, tacts AMEs dorsally 10:30-12:00. Chelic- 10 May 1969, 1,800 m el. (1$, AMNH), Chiricahua erae scales with Mtns., Onion Saddle, 2,370 m el, 20 March 1989; lacking pale Cymbium Pima Co.; Santa CataHna Mtns. (12, AMNH); Santa none to a few white scales. Abdomen show- Co.: Santa Rita Madera Cruz Mtns., upper Canyon, ing lineate markings of females, with two 1,700 m el, 13 August 1983 (19, MCZ). medial longitudinal stripes in addition to the side bands. Measurements: Body Natural History. Collected from oaks (4 4.0(4.3)4.3 mm; carapace length records). length 2.0(2.0)2.0 mm, width/length 0.80(0.81) 0.83; n = 5 5 from Minnesota. 28. Pelegrina arizonensis Female. Epigynum (Figs. 251, 423, 424): (G. & E. Peckham, 1901) Flaps rotated 180° so that ancestrally pos- new combination terior end is anterior. Surface flat except Figures 160, 161, 217, 251, 420-425; for concavity in front of flaps. First curve IVlap25 of duct broad, on medial side of opening because of flap rotation; second curve pro- Dendryphantes arizonensis G. & E. Peckham, 1901b; 1 ceeds laterally. Markings (Figs. 161, 425): 326, pi 28, fig. 2, S. Holotype in MCZ IS with imm. with labels "Dendryphantes arizonensis Pkm, Carapace with white scales dorsally. Clyp- 1901. Arizona. Type. 3." (label is original; hand- eus densely covered with white scales. Ab- written, Elizabeth Peckham) and "G. probably by dominal markings strikingly lineate, with W. Peckham examined. G. & E. Peckham, Coll.", two central bands flanked two thin 1206. pale by 1909; 463, pi 36, fig. 7, S. Roewer, 1954; rows of dark flanked brown bands Dendryphantes glacialis Scheffer, 1905, figs. 3, 4, 8, spots, by $9. Type material lost (Cutler and Jennings, 1985), and pale side bands. Measurements: Body the Peckham collection has some material though length 4.7(4.9)5.9 mm; carapace length labeled Dendryphantes glacialis from Manhattan, 2.0(2.0)2.2 mm, width/length 0.77(0.80) Kansas, possibly sent by Scheffer, examined. G. & 0.84; n = 52 from Minnesota. E. Peckham, 1909; 463, pi 37, figs. 7, 7a, b, 69. Roewer, 1954; 1210. Courtship (13 observed from Anoka Co., 298 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

and "Coll and don Helen Van Du- Minnesota): The one male observed showed IV-7-'18," by = zee, examined. Roewer, 1954: 1211. no crouch Raise (n 16): display. spread Dendryphantes sausalitanus Chamberlin, 1925b: 137, = First Abdomen twitched on pause (n 3). 5. in MCZ 15 with label "Den- = figs. 57, 58, Type legs waved irregularly on series (n 9); as dryphantes sausalitanus Ch, 5 holotype, Cal.: Sau- salito 1909, R. V. Chamberlin Coll., 1045," ex- he got closer legs lowered, put forward and amined. more parallel but no discrete crouch dis- — = Metaphidippus helenae: Gertsch, 1934: 18. Bonnet, play was seen (n 7). Palpi wave irreg- Cutler and 1985: 12- = 1957: 2814. Jennings, 5, figs. ularly on series (n 4). 17, 59. Distribution (Map 25). Minnesota and Differs from arizonensis in Alberta south to Zacatecas and Tlaxcala. Diagnosis. having blunt embolus, tibial apophysis on Records. CANADA: ALBERTA: Medicine Hat {U an elongate projection, and epigynal laps 19, AMNH). UNITED STATES: MINNESOTA: rotated very far, to 270°. Co.: 5 km E of Bethel (15 39, MCZ; IS 39, Anoka Male. Palpus (Figs. 218, 427, 428): Erect NORTH DAKOTA: Burleigh Co.: Menoken AMNH); of embolus is and Co.: WiUiston portion blade-shaped Indian Village (1

Records. UNITED STATES: IDAHO: Blaine Co.. from the southwest. May be confused with Co.; Salmon 19 km Carey (16, AMNH); Custer River, the sympatric orestes but smaller, more N Challis (19, AMNH); Gem Co.: 11 km W of Horse- than orange, lacking the lateral chel- shoe Bend, 116°18'W, 43°57'N (56, AMNH); 13 km gray iceral and the short embolus \V of Horseshoe Bend (19, AMNH); Jerome Co.; Twin ridge, having into base Falls (1(3, AMNH); Owyhee Co.; Bruneau Canyon Hot broadening gradually prolater- Falls WYOMING; Co.; Creek (19, AMNH); Bighorn ally. 10 km E of Shell (13 29, WPM); UTAH; Little Cot- Male. Palpus (Figs. 219, 433): Embolus tonwood Campground, near Salt Lake City (S6, short, truncate, broadens grad- AMNH); Sevier Co.; Richfield (16 29, AMNH); NE- obliquely rami indis- VADA; Humboldt Co.; 48 km N of Winnemucca (19, ually into base prolaterally; AMNH); Washoe Co.; N of Reno (36 19, MCZ); Reno tinct. Markings (Figs. 162, 432): Carapace Franklin Co.: of (26, MCZ); WASHINGTON; just W with scattered white scales; side bands Palouse 46.66°N, 118.23''W, (39, MCZ); Grant Falls, weak. Cheek band weak. Clypeus brown; Co.; Wahluke Wildlife Recreation Area, 46.705°N, thin white to brown hairs 119.42rW (29, MCZ); OREGON; Baker Co.; Baker overhanging contacts (19, AMNH); Crook Co.; 8 km W of Prineville (23 chelicerae. White forehead band Co.; Redmond 79, AMNH); Deschutes (49, AMNH); AMEs dorsally 10:30-12:30. Chelicerae Co.; Tencent Lake (16 39, AMNH), Manns Harney with some pale scales, scattered but es- Lake (16, AMNH); Klamath Co.; above Algoma (16, pecially medially. Palpus entirely brown, AMNH), Bly Mountain (19, AMNH); Malheur Co.; with femur not than Succor Creek Canyon (26 19, AMNH); E of Ontario, distinctly paler cym- 116°57'W, 44°2'N (19, AMNH); Umatilla Co.; 19 km bium. Cymbium lacking white scales. Ab- Mosier SW of Echo (19, AMNH); Wasco Co.; (19, domen side bands indistinct, often with AMNH); CALIFORNIA; Marin Co.; Sausalito (6, white markings centrally. Measurements: MCZ); Mono Co.; Benton (29, AMNH); Riverside Co.: Lake Hemet 116°59'W, 33''43'N; San Diego Co.; 3 Body length 2.7(3.1)3.6 mm; carapace km E of Pine Springs (16, AMNH); Pine Valley (19, length 1.3(1.5)1.7 mm, width/length 29 = AMNH); San Francisco Co.: San Francisco (1<3 0.77(0.79)0.80; n 53 from Yavapai Co., MCZ); Sierra Co.; Peavine (39, AMNH). Additional Arizona. records are given by Cutler and Jennings (1985). Female. Epigynum (Figs. 252, 434, 435): Natural History. Collected from sage- Flaps dark, parallel, slightly convex, pos- brush (Artemisia tridentata; 7 records). teriorly flush with surface. Surface flat. Cutler and Jennings (1985) give additional First curve of duct relatively narrow, dark; habitat information. second curve proceeds slightly anteriorly. Markings (Figs. 163, 436): Carapace cov- 30. Pelegrina verecunda ered with white scales. Clypeus densely (Chamberlin & Gertsch, 1930) covered with white. Legs pale yellowish. new combination Abdomen pale, with many small dark Figures 162, 163, 219, 252, 432-436; speckles sometimes coalescing into gala- Map 26 thea-hke pattern. Measurements: Body 3.7(4.0)4.5 mm; Sassacus uteanus: —Chamberlin and Gertsch, 1929, length carapace length in 54 have been mis- 1.5(1.6)1.8 mm, width/length 0.77(0.78) part; fig. (this figure may = placed, given that it is unlikely the authors would 0.80; n 59 from Yavapai Co., Arizona. and D. ver- have confused their species S. uteanus Male/Female Matching. This associa- ecundus). tion is indicated by co-collecting, distri- Dendryphantes verecundus Chamberlin and Gertsch, of 1930: 144. Holotype 16 in AMNH with labels "Den- bution, similar size, and indistinctness Ho- dryphantes verecundus 6 / Utah; Dry Canyon markings. 6-14-29 Gertsch" and "29Bb. N40 ; Will," lotype/ (23 observed from Yavapai examined. Courtship Co., First held in a trian- Dendryphantes verecundus: — Roewer, 1954; 1216. Arizona). legs Bonnet, 1956: 1402. gular bowed position during pause of — and 1974; Metaphidippus verecundus: Jung Roth, crouch display, as in P. aeneola. Crouch 33. = = (n 15, 23): Body horizontal (n 15, 23) = and low = to (n 1). First Diagnosis. A small indistinctly marked (n 13, 23) high held low and forward, bowed, with species with dark males and pale females legs 300 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

= twists tips almost touching (n 15, 26); on series toward tip. Markings (Figs. 164, with forehead band. legs extended forward and waved vigor- 437): Carapace large = ously (n 15, 25) though somewhat irreg- Cheek band dense and distinct from side = band. brown in two ularly (n 8, 1(3); on pause legs retracted Clypeus specimens, = = with white scales between in (n 12, 2(5) or not (n 3, 15). Palpi held brown AMEs = down (n 11, 25). On series palpi held in another specimen; hairs overhanging che- = and under and motionless (n 3, 15); at licerae brown to tan. White forehead band setae end of series/start of pause palpi hang contacts AMEs dorsally; ringing = down and wave a few times (n 3, 15). AMEs white except 12:00-1:30. Chelic- of white scales Repertoires: 25 crouch only. erae with dense patch from to and wider than Distribution (Map 26). Utah south into base % length, longer northern Mexico. in variegata. Cymbium lacking white scales. Measurements: Body length 4.1, 4.2, Records. UNITED STATES (county records); 4.3, 4.4 mm; carapace length 2.1, 2.1, 2.1, UTAH: Kane (19, AMNH), Salt Lake (173 19, AMNH), 2.2 0.76, 0.77, 0.77, 0.80; Sevier Utah (3<5 79, AMNH), Wasatch mm, width/length (33. AMNH), = (113 39, AMNH), Washington (93 19, AMNH, MCZ), n 45 from Nuevo Leon. ARIZONA: Cochise 129, Weber (23, AMNH); (53 Female. Epigynum (Figs. 439, 440): MCZ), Coconino (13, AMNH), Santa Cruz AMNH, Flaps flat and convergent, medial edge at (19, MCZ), Yavapai (83 119, AMNH, MCZ); NEW level of central and lateral MEXICO: Bernalillo (23, AMNH), Catron (19, high plateau in so that down AMNH), Grant (39, AMNH), Sandoval (13, AMNH); edge concavity flap slopes CALIFORNIA: Mono (13, AMNH). MEXICO: CHI- laterally. Epigynal surface high between HUAHUA: Las Delicias (13 49, AMNH); PPrimavera flaps as noted. First curve of duct broad (19, AMNH); 21 km N of Ciudad Camargo on Hwy but does not extend so far posterior as in 45, 105°13'W, 27°52'N (29, MCZ); ?40 km SW of the neoleonis, so that second Camargo (19, AMNH); PDURANGO: Ojo de los En- sympatric cinos (19, AMNH). curve begins at or only a bit posterior to posteriormost portion of flap; second curve Natural History. Specimens collected at proceeds obliquely medial-anterior. Flow- elevations from 1,100 to 1,800 m (12 re- erlike gland opening on anterior face of In Arizona, beaten from cords). Quercus, second curve. Inner surface of third curve Cercocarpus, Alnus, Salix, Chnjsotham- fairly smooth. Markings (Fig. 441): Car- nus, and spruce (6 records). Jung and pine, apace covered above with white scales. Roth (1974) found this species in their zone Clypeus covered densely with white scales. 2 of the Chiricahua Mountains. Abdomen marked somewhat like P. gal- athea. Measurements: Body length 4.0, 4.2 31 . clavator new Pelegrina species mm; carapace length 2.0, 2.0 mm; width/ 1 1 1 64, 65, 220, 437-441 ; 5 = Figures Map length 0.76, 0.77; n 29 from Nuevo Leon. Males and fe- Holotype male and paratype female with label Male/Female Matching. "MEXICO: NUEVO LEON: Chipinque Mesa just males were matched by co-collecting at S of Monterrey, ca. 4500 ft. [1,370 m]; ca. 100.4°W two localities in Nuevo Leon, by common 25.6°N, 2 1983, W. Maddison & R. S. Anderson Jun distribution; and by the similar robust car- 83-034, beating and sweeping forest understory.' apace and abdominal markings. Etymology. A Latin noun in apposition, Geographical Variation. Females from "club-bearer," referring to the large blunt Tamaulipas and Veracruz have smaller embolus. flaps less deeply set into epigynum, with Diagnosis. A Mexican species having a less flaring ducts, and may represent an- distinctive broad, truncate embolus and other species. flaps flanking a central mound on Courtship (25 observed from Chipinque angled = the epigynal surface. Mesa, Nuevo Leon). Raisedspread (n 3, = Male. Palpus (Figs. 220, 438): Embolus 15). Crouch (n 3, 25): First legs foward, = broad, truncate, with rami small. Embolus slightly spread, horizontal (n 1) or raised Pelegrina Jumping Spiders • Maddison 301

= fairly high to ca. 40° (n 2, IS), lowered daracina, yucatecana, verecunda) the fe- = when close (n 2, 1<3); not noticeably males described below and the type of pal- = = waved (n 3, 2S). Palpi down (n 3, 26), lidata are unique in having (1) an unusually = waved on series (n 3, 25) and when very broad dark band along the margin of the = close and reaching (n 2, 13). Abdomen opening (Fig. 445), (2) first curve of duct twitched occasionally, possibly at pause (n wide and long, (3) the flowerlike gland = 2, is). Repertoires: IS crouch only; IS openings placed on the anterior surface of raisedspread and crouch. the second curve and more medially (clos- Distribution (Map 15). Nuevo Leon er to junction with third curve than first south to Veracruz. curve), and (4) fertilization ducts arising anterior to the center of the lumen of the Records. MEXICO: TAM.AULIPAS: ca. 1.5 km E spermatheca. of Tula, 99.5°W, 22.9°N, 8 June 1983 (12, MCZ); Male Associated with Fe- Sierra de Tamaulipas, 4-7 August 1945 (15, AMNH); (Tentatively SAN LUIS POTOSI: 32 km E of Ciudad del Mais, male). Palpus (Figs. 221, 443): Embolus 23 March 1940 NUEVO LEON: (12, AMNH); Chip- twists apically. Retrolateral ramus rela- inque Mesa, just S of Monterrey, 100.4°\V, 25.6°N, 2 tively long and curved, as in P. pervaga June 1983 and 7 April 1946 (33 12, MCZ; 12 AMNH); and tristis, but embolus much smaller than \'illa de Santiago, Hacienda Vista Hermosa, 19 June 1940 {IS, MCZ); VERACRUZ: 2 km SE of Naolinco in those species. Markings (Fig. 442): Car- on Hvvy 127, 96.9°W, 19.6°N, 20 June 1983 (12, MCZ). apace medium to pale brown, with well- developed side and cheek bands. Clypeus Natural History. Collected from un- brown except for patch of white scales be- derstory shrubs of broadlead forest at ca. tween AMEs that overhangs chelicerae. 1,400 m elevation (2 records); also known White forehead band contacts AMEs dor- from 600 to 800 m elevation (3 records). sally 10:30-12:00. Chelicerae with white scales medially. Cymbium with some white 32. Pelegrina pallidata scales. Legs beige with brown annulae. Ab- (F. P.-Cambridge, 1901) domen shows white spots of female. Mea- new combination surements: Body length 3.4 mm; carapace 29 = Figures 221, 442-446; Map length 1.7 mm, width/length 0.77, n 15. Female. 444, 445): Metaphidippus pallidatus F. P.-Cambridge, 1901: Epigynum (Figs. not convex. 270, pi. 24, figs. 17, 17a, 2. Holotype in BMNH 12 Flaps convergent, very Epi- with label "Dendryphantes pallidatus, sp. nov. Guat. gynal surface flat. First curve of duct fairly examined. 1957: 2816. Sarg Type 2," Bonnet, broad and long; second curve goes anter- Dendryphantes pallidatus: —Roewer, 1954: 1198. iomedially. As already noted, there is an Notes on Synonymy. The specimens de- unusually broad band along the margin of scribed here are identified with Cam- the opening (Fig. 445, arrow), the flow- are on the bridge's pallidata primarily on the basis erlike gland openings placed of similarities in the details of the epigyn- anterior surface of the second curve close um. This identification is made with some to junction with third curve, and the fer- anterior to the center hesitation, for the type material of palli- tilization ducts arise of the Mark- data consists of a single poorly marked of the lumen spermatheca. female whose epigynum was lost by me in ings (Fig. 446): Cambridge's holotype is the course of examination, but my notes now uniformly pale, though may be partly available and the figures of C. L. Scioscia (personal faded. The other specimens have communication) regarding the epigynum the carapace covered with yellowish white not dense- are fairly detailed and provide evidence scales, though densely. Clypeus for the identification. Compared to the ly covered with white scales; AMEs en- other Mexican and Central American spe- tirely ringed by white scales. Legs uniform cies with small, convergent flaps and a fair- orange-brown except Nicaragua 9, which Sternum ly flat epigynal surface (variegata, san- has some brown spots. distinctly 302 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

darker than coxae. Abdomen with paired base on retrolateral side, so as to make white spots on brown dorsum; each spot distinct angle. Markings (Figs. 166, 447): in first pair fused with spot in second pair. Carapace with extensive white markings. Venter dark between epigynum and spin- Cheek band broad and dense, fused with nerets. Measurements: Body length 4.0, side band. Clypeus brown, with setae over- 4.3 mm; carapace length 1.6, 1.7 mm, hanging chelicerae white medially, some = width/length 0.76, 0.78; n 22, female brown hairs laterally. White forehead band holotype and 19 from Nicaragua. contacts AMEs dorsally; setae ringing Male/Female Matching. This is indi- AMEs white except from 12:30 to 2:00. cated by co-collecting in Nicaragua and Chelicerae robust, though not elongate, similarity of markings. with white patch on medial surface from Distribution (Map 29). Southern Mexico base to about V2 length. Cymbium with to Nicaragua. central patch of white scales. Legs dis- tinctly annulate. Abdomen not striped as Records. MEXICO: CHIAPAS: 5 km W of San in most Pelegrina males but rather with Cristobal de Las Casas on Hwy 190, ca. 92°41"W, white almost as in 9. Mea- 16°44'N, 27-28 July 1983, W, Maddison & R. S. An- paired spots derson (19, MCZ). GUATEMALA (19, BMNH); Chi- surements: Body length 3.4(3.9-4.2)4.4 chicastenango, 6-7 August 1947, C. & P. Vaurie (29, mm; carapace length 1.7(1.9-2.1)2.2 mm, AMNH). NICARAGUA: 4 October 1952, Matagalpa, width/length 0.79(0.80-0.81)0.84; n = 63 R. B. Swain (1

ward, horizontal to 10° raised, bowed, tips desert scrub at 1,500 m elevation (Oaxaca); slightly convergent, parallel or slightly beating shrubs and trees in fairly dry bot- = spread, not touching (n 19, 46). On each tom of river valley at 600 m elevation = series legs flickered (n 9, 13) noticeably (Nuevo Leon); and from a pine forest = = (n 1) or with fairly low amplitude (n (Chiapas). Known from 220 to 1,700 m = 7, 36) or perhaps not at all (n 3, 13). Palpi elevation throughout Mexico (8 records). = held down (n 12, 43), pointing inward = and resting over chelicerae (n 7, 23), on 34. Pelegrina yucatecana new species each series flickered with fairly low am- Figures 169, 223, 452-456; Map 29 = = (n 12, 13) outward (n 1) or up plitude male and female in MCZ with and = Abdomen twitches Holotvpe paratype down (n 7, 23). labels "MEXICO: YUCATAN: 3 km E of Chichen = = (n 6, 13) at end of each series (n 3, Itza ruins on Hwy 180, ca. 88°34"W 20''40'N, 19- = 13) or in pause (n 4, 23). 20 July 1983 W. Maddison & R. S. Anderson, 8.3- 115 seasonal and trailside Distribution (Map 28). Nuevo Leon forest, beating understory shrubs and small trees." south to Panama. Etymology. An adjective, formed after Records. Most in some in from: AMNH; MCZ, yucateco (Spanish) or yucatecan (English), MEXICO: TAMAULIPAS: 11 km E of Ocampo, referring to the Yucatan Peninsula. 99°16'W, 22°49'N (19); 35 km SSW of Mante (3<5); An species with Paso del Abra 99.0rW, 22.45°N (1<5); Mante (15 52); Diagnosis. interesting in 23 km S of Villa Juarez (19); Hidalgo (U); 19 km SE unusual transverse abdominal markings; of Ciudad Victoria (19); Rio Guajolotes, 64 km S of genitalia resembling variegata and san- Victoria 24 km S of Victoria {2$ 29); Sisal, (23 29); daracina but differing from both in details. Ciudad Victoria (16 39); 18 km N of Victoria (13); Male. Palpus (Figs. 223, 453): Embolus SAN LUIS POTOSi: Covadonga, WSW of Valles with rami small. Erect 99.05°W, 21.57''N (19); Valles (29); Taninul, Valles (13 short, very portion 19); El Salto (13); 19 km E Ciudad del Maiz (19); Pujal of embolus with sides parallel; widens (19); NUEVO LEON: Santa Rosa Canyon 29 km W abruptly at base so that a distinct angle is of Linares Montemorelos CHIHUA- (23 29); (19); made between the erect portion and base HUA: 8 km S of Chihuahua (13); Catarinas (13); SIN- the ALOA: 64 km S of Culiacan (13); 48 km N of Mazatlan along prolateral margin. Markings (Fig. is its (13 19); 10 km E of Villa Union (13); Culiacancito 452): As only known 3 teneral, proper 107.32°W, 24.50"'N (23); DISTRITO FEDERAL: colors are not exactly known, though ap- Xochimilco MORELOS: Cuernavaca NAY- (13); (19); pears brown with white markings. Mar- ARIT: Tepic (143 115); 43 km S of Tepic (19); 56 km ginal band well developed. Carapace with S of Tepic (13 19); La Mesa de Nayarit (29); San Bias forehead band an acute (13); Jahsco (19); Jesus Maria (23); JALISCO: Zapot- V, proceeding lanejo (13); Zapotlanejo (13); COLIMA: 32 km N of more posteriorly from AMEs than later- Cohma GUERRERO: Chil- (13 19); Iguala (13 19); ally. Cheek band dense, and distinct from VERACRUZ: Plan del pancingo (13); Teloloapan (79); side band. Clypeus brown, with setae over- Rio (23 19); Tierra Colorado OAXACA: 2 km (23 19); chelicerae dark. White forehead S of El Tule (23 89); 3 km W of Tapanatepec (13); hanging contacts 10:30-12:00. Oaxaca (13); San Felipe, N of Oaxaca City (33 49); band AMEs dorsally Paso Real, Rio Tonto (19); Tehuantepec (43); Monte Chelicerae with dense medial patch of Alban 3 km SE of 16.32°N (13); Niltepec, 94.33°W, white scales from base to V2 length. Cym- (13); Soladad (23); CAMPECHE: Campeche (23 59); bium dark basally, paler at tip; lacking YUCATAN: Progresso (13); Motul (13); Chichen Itza white scales; and tibia dark. (23); CHIAPAS: Arriaga, N of Arriaga Mtns. (19); 24 patella Legs km NW of Arriaga 94.01''W, 16.25''N; Cintalapa (83 strongly annulate, differing from sandar- 99); Ocozucuantla (33 49); Rio de las Flores, 30 km acina in having the back 2 annulate in- NE of (73 89); Tuxtla Gutierrez (43 19); Las Cintalapa stead of longitudinally striped. Abdomen Cruzes (33 19). HONDURAS: Zamorano. NICARA- shows transverse pattern similar to 9. Mea- GUA: San Marcos (13 49). COSTA RICA: San Jose 3.4 (19). PANAMA: 8 km S of El Valle (23). surements: Body length mm; carapace length 1.7 mm, width/length 0.82. Natural History. Collected beating Aca- Female. Epigynum (Figs. 454, 455): cia, composites, and other vegetation in Flaps very pale and slightly convergent. 304 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

long, about half as long as epigynum. Sur- Diagnosis. This Mexican and Central face flat. First curve of duct pale, fairly American species shares with the sympat- narrow; second curve proceeds slightly an- ric yucatecana and variegatus prominent chelicerae of the teriorly. Markings (Figs. 169, 456): Car- pale patches on the male apace brown dorsally except three white and a relatively small embolus but differs transverse bands: between small eyes, just from both in having a patch on the clypeus in front of fovea, and just behind fovea; between the AMEs of distinctly yellow the first two are sometimes connected by scales and in lacking prominent pale two small white longitudinal bands. Face patches on the abdominal dorsum. The dark, with only scattered pale scales. In erect portion of the embolus broadens more particular, the clypeus lacks white beneath gradually into the base than in yucatecana. the AMEs except for setae overhanging The female is orange, superficially bearing chelicerae and ringing AMEs. Legs strong- close resemblance to Nagaina incunda but ly annulate. Abdominal markings unusual differs in having yellow scales on the face transverse dark spots. Measurements: Body even under the AMEs, and in having stron- length 3.4(3.8-4.0)4.9 mm; carapace length ger epigynal flaps. See also comments un- 1.7(1.8)1.9 mm, width/length 0.77(0.81) der P. pallidata. 0.83; n = 49 from Yucatan and Campeche. Male. Palpus (Figs. 224, 458): Embolus Male /Female Matching. Males and fe- small, wider at base and tapering to tip, males have similar markings on abdomen broadens gradually into embolar base so and similarly annulate legs; the thin em- that along prolateral margin there is no bolus would be expected matched to a fe- angle distinctly marking embolus from its male with weak flaps; and they are mi- base. Markings (Figs. 168, 457): Carapace crosympatric at Chichen Itza. well marked with discrete bands of yellow Distribution (Map 29). Yucatan Pen- scales. Marginal band weak or absent. insula. Cheek band broad and dense though dis- tinct from side bands, unlike variegatus. Records. MEXICO: YUCATAN; 3 km E of Chi- Clypeus with prominent patch of yellow chen 19-20 1983 Itza, 88°34'W, 20°40'N, July (16 19, scales between AMEs and overhanging the MCZ); 4 km N of Xocenpich, 88°34'W, 20°47'N, 20 chelicerae, otherwise brown. Yellow fore- July 1983 (19, MCZ); 12 km S of Muna on Hwy 261, contacts 1 LOO- 89°46'W, 20°24'N, 21 July 1983 (19, MCZ); CAM- head band AMEs dorsally PECHE: Chicanna ruins 8 km W of Xpujil, 89°31"W, LOO. Chelicerae with long dense patch of 12-14 1983 18°32'N, July (19, MCZ). yellow scales on medial edge from base to % length. Cymbium dark, lacking white Natural History. One of the few low- scales; tibia and patella paler. Legs orange land tropical species of Pelegrina. All with strongly contrasting markings of dark known specimens were collected beating brown. On posterior lateral face of second shrubs and small trees in understory and leg tibia is a longitudinal dark band. Dark along trails through short tropical forest. on femur 3 restricted to subterminal spot on front and back. Abdomen in some males 35. sandaracina new Pelegrina species with distinct paired dark spots. Measure- 29 Figures 168, 224, 457-463; Map ments: Body length 3.0(3.1)3.6 mm; car-

male in MCZ with label CAM- apace length 1.4(1.5)1.9 mm, width/length Holotype "MEXICO: = PECHE: 6 km W of Francisco Escarcega, "El Tor- 0.78(0.80-0.81)0.81; n 43 from Cam- ca. 11-12 mento" forest station, 90°48'W, 18°37'N. peche, Oaxaca, Jalisco, and "Managua," July 1983 W. Maddison 83-107, beating understory Mexico. shrubs of forest of small trees." Female. Epigynum (Figs. 459, 460, 462, Etymology. Latinized from the Greek 463): Flaps convergent, shorter than those sandaracinos, orange-colored (Woods, of sympatric yucatecana, less than half 1966). length of epigynum. Surface flat. First Pelegrina Jumping Spiders • Maddison 305

Records. MEXICO: YUCATAN: Grutas de curve of duct pale in Yucatan $, dark in Loltun, 7 km S of Oxkutzcab, 89°27'W, 20°15'N, 22 1983 others; second curve proceeds a bit ante- July (1$, MCZ); CAMPECHE: 6 km W of Francisco Es- unlike in which riorly, Nagaina incunda, carcega, "El Tormento" forest station, 90°48'W, second curve proceeds more posteriorly. 18°37'N, 11-12 July 1983 (1

4.2 mm; carapace length 1.7, 1.7, 1.8, 1.9 37. Peleghna bunites new species mm, width/length 0.74, 0.74, 0.75, 0.77; Figures 170, 171, 226, 255, 478-482; = n 4(5 from Cooper, South Carohna. Flor- Map 30 ida: body length 3.1, 3.2, 3.2 mm; carapace Holotype male and paratype female in MCZ with length 1.4, 1.4, 1.5 mm, width/length 0.74, label "ARIZONA: Santa Cruz Co., Santa Rita Mts., 0.75, 0.77; n = 3(5 from Florida. gate at 26 km of Whipple Obs[ervatory]. Rd. on Female. Epigynum (Figs. 254, 475, 476): Mt. Hopkins 7100 ft [2,170 m] el. 17 June 1985 W. Maddison montanus." Flaps pale, only slightly convex. Surface 85-059, beating Cerocarpus flat. First curve of duct pale; second curve Latinized from the proceeds medially and slightly anteriorly. Etymology. Greek hill-dweller. Markings (Fig. 477): Carapace covered bounites, In with white and some tan scales dorsally, Diagnosis. general appearance, resembles side bands distinct. Clypeus densely cov- strongly other Pelegrina species ered with white scales. Abdomen with cen- but lacks the characteristic Pelegrina em- bolus with subterminal tral longitudinal pale stripe flanked by dark opening and two rami. The most distinctive features are the stripes flanked by pale stripes. Measure- embolus whose erect ments: South Carolina: body length 4.3, portion twists and ta- toward and the distinct 4.4, 4.4, 4.6 mm; carapace length 1.9, 1.9, pers tip bend on the This is 2.0, 2.0 mm, width/length 0.76, 0.76, 0.78, epigynal flaps. species only ten- = 0.78; n 49 from Cooper, South Carolina. tatively placed in Pelegrina, for the em- bolus has its Florida: body length 3.6(4.0)4.1 mm; car- opening terminal and lacks two distinct rami. apace length 1.7(1.7)1.7 mm, width/length Male. 0.75(0.76)0.79; n = 52 from Lake Placid, Palpus (Figs. 226, 479): Erect por- Florida. tion of embolus twists and tapers toward Geographical Variation. Specimens tip. Embolus with only one ramus near the which is almost terminal. from central Florida are distinctly smaller opening, Mark- and paler than more northerly specimens ings (Figs. 170, 478): Carapace side bands and appear more yellow than brown. Males and forehead band well developed. Cheek from central Florida have the cymbium band broad, dense and distinct from side bands. yellow with a discrete brown spot at the Clypeus brown, with brown to white hairs tip and an embolus that is apparently overhanging chelicerae. Forehead band contacts slightly wider than in northern males. AMEs dorsally 10:30-12:30. Chelicerae with medial of Distribution (Map 27). North Carolina long patch white south to Florida, west to Texas. scales in Arizona males; Oaxaca male with shorter patch. Cymbium brown, lacking Records. UNITED STATES: NORTH CAROLI- pale scales. Legs mostly beige except for NA: 24 Polluckville, October 1926 {86 59, AMNH); mostly dark brown first pair and brown SOUTH CAROLINA: Cooper, 25 December 1928 marking on more posterior pairs. Abdo- (155 139, AMNH); FLORIDA: Lake Placid, Archbold men brown dorsally with distinct white Biological Station, 26 March 1968 (79, MCZ) and 1 October 1962 side bands. Mesurements: 3.4, {13, AMNH); Mariana, Blue Springs, Body length 12 March 1936 (19, AMNH); Ortega {IS, AMNH); 4.1, 4.4, 4.6 mm; carapace length Glades Co.: Fish Eating Creek, 23 February 1951 {16 1.5(2.1)2.2 mm; width/length 0.77(0.79) AMNH); MISSISSIPPI: Vancleave, = Pascagoula River! 0.81; n 5(5 from Mount Hopkins, Arizona. Wards Bayou {16, AMNH); LOUISIANA: Baton Female. (Figs. 255, 480, 481): Rouge (1<5 19, MCZ); Tallulah, 9 March 1925 (1

(Figs. 171, 482): Yellow to light brown. HUAHUA: Pelayo, 101 km W of Santa Barbara, 20 1947 (19, AMNH); Santa Barbara, 18 1947 Oaxaca females are generally darker than July July (19, AMNH); DURANGO: 16 km E of El Salto, 8 Arizona females. Carapace covered thinly August 1947 (13, AMNH); OAXACA: 50 km NW of with scales. covered with beige Clypeus Oaxaca on Hwy 190, ca. 97°00"W, 17°14'N, ca. 2,000 white to yellowish scales. Legs more or less m, 6 August 1983 (1<5 69, MCZ). uniformly beige to light yellow brown in Natural History. Beating Cercocarpus Arizona females, orange-brown in Oaxaca montanus on Mount Hopkins, Arizona; females. Abdomen often with paired dark beating pine trees in clearing in oak-pine spots posteriorly, similar to insignis. Mea- forest in Oaxaca. At elevations from 1,800 surements: Body length 3.6(4.5)4.7 mm; to 2,200 m in Arizona and Oaxaca (4 re- carapace length 1.9(2.0)2.0 mm; width/ = cords). length 0.76(0.80)0.81; n 59 from Mount Hopkins and Kitt Peak, Arizona. 38. Chromosomes. 2nS = 26 acrocentrics + Pelegrina orestes new species Figures 172, 173, 227, 483-487; 30 XXO (1(5 from Mount Hopkins, Arizona). Map observed Courtship (3<5 from Santa Rita Holotype male and paratype female in MCZ with Mtns., Arizona, and near Oaxaca City, Oa- label "ARIZONA: Santa Cruz Co., upper Madera Santa Rita ca. 5500 ft. xaca). Has crouch display with exagger- Canyon, Mts., [1,680 m] 13 Aug 1983. W, Maddison 83-158 oak woodland, ated leg waving during pauses. Raised- = = beating oaks, especially Q. hypoleucoides." spread (n 3, 2(5). Crouch (n 7, 33): = Body held normal to low (n 2, 26) or Etymology. Greek, mountaineer. = high (n 1). First legs forward, spread Diagnosis. Resembling the sympatric = slightly, horizontal (n 2, IS), or slightly verecundus but larger and more orange; = = also in raised (n 1), or slightly lowered (n 2, differing the more abrupt angle 2$) flickered rapidly with low amplitude between the erect portion of the embolus = = the base. (ca. 5-10°?, [n 1]) on series (n 5, 33), and The lack of a second ramus but waved up and down with higher am- near the embolic opening makes the place- = of this in plitude (ca. 30°? [n 1]) ca. 3-7 times (n ment species Pelegrina tentative. = = Male. 3, is) or a few times (n 1) during Palpus (Figs. 227, 484): Embolus = = widens into pause (n 3, 1<3) or at end of series (n abruptly base on prolateral side to a 3, 2S). During series legs spread slightly yield sharp discontinuity between erect but distal segments parallel; during pause portion and base. Embolus with only one = legs held wider then parallel (n 3, 13); ramus retrolateral to opening. Chelicerae: in as he got closer he reached legs to parallel Outer edge some males bears a slight = = (n 2, 23). Palpi down (n 2, 23) and ridge similar to that seen in the mannii = curled beside chelicerae (n 1), flickered group. Markings (Figs. 172, 483): Indis- = tinct with low amplitude on series (n 3, 23). beige marks on brown to orange = side Abdomen bobs very little if at all (n 1). background. Carapace bands with ex- tension Repertoires: 13 crouch only, 23 raised- toward fovea. Cheek band distinct spread and crouch. from side band. Clypeus brown, hairs chelicerae tan Distribution (Map 30). Southern Ari- overhanging to brown. zona south to Oaxaca. Forehead band does not reach AMEs, so that setae surrounding AMEs are brown Records. UNITED STATES: ARIZONA: Santa Rita above. Chelicerae with small medial Mtns., Sweetwater, 1,800 m, 25 June-2 July 1951 (19, patch of scales. AMNH); Cochise Co.: Huachuca Mtns., 18 July 1936 pale Cymbium brown, lacking (19, AMNH); Pima Co.: Quinlan Mtns., picnic area pale scales. Legs beige and brown, with near Kitt Peak m 20 Observatory, 1,950 elevation, annulate markings. Abdominal dorsum June 1985 (45 39, MCZ); Madera 8 Canyon, Septem- darker than side bands but not ber 1978 (19, MCZ); Santa Cruz Co.: Santa Rita Mtns., distinctly dusted with scales. 2,150 m el. on Whipple Observatory Road, Mt. Hop- so, pale Measurements: kins, 17 June 1985 (83 59, MCZ). MEXICO: CHI- Body length 3.8(4.2)4.8 mm; carapace 308 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

AMNH); OAXACA: 39 km NW of Oaxaca on Hwy length 1.8(2.0)2.4 mm; width/length 1983 = 190, ca. 96°57'W, 17°17'N, 6 August (1<5, MCZ). 0.78(0.81)0.84; n 56 from Santa Cruz Co., Arizona. Natural History. At Madera Canyon in Female. Epigynum (Figs. 485, 486): Arizona, beating oaks, especially Quercus Flaps depigmented, convergent; at their hypoleucoides, in oak woodland. Also col- posterior end the flaps lie beneath well- lected from oaks at other localities (2 re- pigmented medial rim of opening. Epi- cords, Arizona and Oaxaca). Collected from gynal surface more or less flat. Markings 1,200 to 1,900 m elevation in Arizona and (Figs. 173, 487): Pale, yellow-orange, with Oaxaca (5 records). At Madera Canyon, little hint of markings. Carapace thinly this species was common in August (5(5 39) covered with yellow-white scales. Clypeus but rare in June (1(5). covered with white scales. more or Legs THE GENUS NAGAINA less uniform beige to light yellow-brown. G. & E. PEGKHAM, 1896 Abdomen with small speckles somewhat as in verecundus, otherwise pale. Mea- This genus has received little attention, surements: Body length 5.0(5.2)5.7 mm; but its type species (by monotypy), N. in- carapace length 2.1, 2.1, 2.2, 2.2 mm; cunda, is a common Central American = width/length 0.79, 0.79, 0.80, 0.81; n 4$ species that has usually gone by different from Santa Cruz and Cochise Co., Arizona. names Metaphidippus flavolinea- = (e.g., Chromosomes. 2n3 26 acrocentrics + tus). It is described here to resolve the

XXO (2(5 from Madera Canyon, Arizona). taxonomic confusion surrounding it and for Courtship (3<5 observed from Santa Rita because it may be confused sympatric Mtns., Arizona, and near Oaxaca City, Oa- Pelegrina species. The status of the genus xaca). Has crouch display with unusual Nagaina awaits further study. As noted in ^ walking motion. Raisedspread (n 3, 16). the discussion of the mannii group, N. in- = of the Crouch (n 11, 3(5): Body normal height cunda resembles species both man- = (n 8, 26). First legs bowed and forward nii group and the genus Eris, but the shared = (n 10, 26). At distance: legs below hor- characteristics may be plesiomorphies. It = izontal with tips on ground (n 8, 26); is also not clear whether or not the other flickered while walking to yield strange species described in the genus (N. diade- = combined motion (n 6, 26). Within 1-2 mata Simon, N. tricincta Simon, N. mo- body lengths: first legs off ground to hor- desta di Caporiacco, N. herlandi Soares & izontal and no longer involved in walking, Camargo, N. olivacea Franganillo) belong flickered with low amplitude during series, with N. incunda. = still during pause (n 8, 3(5). Palpi down = (n 5, 33), and curled to side of chelicerae 39. incunda = Nagaina (n 1); flickered during series, still during G. & E. Peckham, 1896 = 26 crouch pause (n 6, 26). Repertoires: Figures 174, 175, 228, 488-492; only, 16 raisedspread and crouch. Map 37 Distribution (Map 30). Southern Ari- incunda G. & E. Peckham, 1896: 55, pi. 4, zona to Oaxaca. Nagaina in 19 with label figs. 10, lOa-c, 9. Holotype MCZ '883 Nagaina incunda Peck, Guatemala 9 4312 Records. UNITED STATES: ARIZONA: Cochise Type, G. W. & E. G. Peckham Coll." (in Bryant's Co.: Cave Creek Canyon, above Portal, 9 June 1977 handwriting), from the east coast to Guatemala (G. examined. Roewer, 1954: (1(3, MCZ); Chiricahua Mtns., South Fork Cave Creek, & E. Peckham, 1896), 13 June 1958 (23, AMNH); Chiricahua Mtns., July 1022. Bonnet, 1958: 3027. E. 1901b: 1985 (IS, AMNH); Santa Cruz Co.: Santa Rita Mtns., Dendryphantes vegetiis G. & Peckham, 9. in MCZ 249 5 im. Madera Canyon nr. Bog Springs Cmpgd., 13 August 323, pi. 28, figs. 7, 7a, Types Peck. 1983 and 17 June 1985 {66 39, MCZ); Madera Canyon, with labels "476 Dendryphantes vegetus Type, 9 G. W. & E. G. Peckham 16-24 July 1951 (13, AMNH). MEXICO: CHIHUA- Mexico; San Rafael 4132, Roew- HUA; Canon Prieta nr. Primavera, 30 June 1947 {IS, Coll." (in Bryant's handwriting), examined. Pelegrina Jumping Spiders • Maddison 309

er, 1954: 1201. Bonnet, 1957: 2818. NEW SYN- scales. dark below AMEs, ONYMY Clypeus lacking scales, just above vertical dark line on each Metaphidippus flavolineatus F. P.-Cambridge, 1901: chelicera, but between AMEs a triangular 268, pi. 24, figs. 9, 9a-c, $. Types in BMNH 3

ROO: 31 km NE of Felipe Carrillo Puerto, 87°52'W, examined. As already noted, there are two 19°48'N Kohunlich 18°26'N (19, MCZ); ruins, 88°48'W, species placed in Pelegrina, P. bunites and (23 39, MCZ); CHIAPAS: Palenque ruins, 92°01'W, P. orestes that may rather belong to the 17°29'N (39, MCZ); 77 km SE of Palenque on road mannii the mannii to Bonampak OLS^W, 17. FN {46, MCZ); 105 km SE group. Tentatively, of Palenque on road to Bonampak, 91.3°W, 17.0°N group is considered to exclude these. De- 76 km S of on road to Oco- {IS 29, MCZ); Palenque scribed here are the six mannii group spe- 92.2°W, 17.1°N (4

and Peckham Coll. Peckham) "G.W. ", examined. and tricolor, usually darker than carme- Both 6 lack palpi; 1 is a 3 mannii, other is diplacis; nensis and chera. the Peckhams' description indicates mannii. (Ju- Male. Palpus (Figs. 23, 229, 230, 494, nior primary homonym of Attus imperialis Rossi.) Dendryphantes tnanii G. & E. Peckham, 1901b: 326, 498, 499): Embolus more or less straight; $. in 1<5 with labels pi. 28, figs. 1, la, Holotype MCZ blade-shaped, fairly thin and triangular Mannii Pkm 1901. Arizona. "Dendryphantes Type. viewed ventrally but wide when viewed $." (label is original; handwritten, probably by Eliz- laterally. Base of embolus sclerotized abeth Peckham) and "G.W. Peckham Coll.", ex- along amined. retrolateral margin and, especially in Ar- Dendryphantes manni: — Roewer, 1954: 1212. izonan males (Fig. 230), extended into —G. & E. 1909: Dendryphantes imperialis: Peckham, prong. Markings (Figs. 178, 180, 493): 459, 37, 2b-d and 2a, 6. pi. figs. possibly Carapace dark, side bands generally ab- Dendryphantes versicolor G. & E. Peckham, 1909: sent or much reduced (Figs. 178, 493) ex- 475, pi. 36, figs. 6, 6a, 2. Types in MCZ 595 with in labels "Dendryphantes versicolor P. 9 Salem Ore- cept Arizona (Fig. 180). Cheek band gon Type" (label is original; handwritten, probably dense and white, makes striking contrast Elizabeth and "G. W. by Peckham) Peckham Coll.", against dark body. Clypeus brown. Fore- examined. 1954: 1216. Roewer, NEW SYNONY- head band absent. Setae MY. surrounding AMEs dark white Chelicerae Dendryphantes diplacis: —Chamberlin, 1924, in part: except laterally. 686 (Arizona paratype). with dense patch of white scales. Palpus Metaphidippus imperialis: —Gertsch, 1935: 29. Bon- medium to dark brown with discrete white net, 1957: 2814. band across the distal end of the femur. Metaphidippus versicolor: —Bonnet, 1957: 2818. Cymbium brown, lacking white scales. Legs light to medium brown with darker Notes on Synonymy. G. E. Peckham but indistinct annulae. Abdomen side bands (1901b) described mannii as having yel- often incomplete posteriorly. Measure- low legs and palpi with restricted brown ments: Body length 3.5(4.2)4.8 mm; car- markings and extensive white on the side apace length 1.7(1.9)2.2 mm; = width/length of the carapace, and they figured a narrow 0.78(0.81)0.85; n 5<5 from California. embolus with the embolic base rounded Female. Epigynum (Figs. 256, 495, 496, retrolateral to the erect portion of the em- 500, 501): Flaps dark, narrow and flat. Epi- bolus; in these respects, the description gynal surface more or less flat. Markings seems to match chera better than the spe- (Figs. 179, 181, 497, 502): Except in Ari- cies here considered mannii, but the spec- zona, carapace shiny brown, because in- imen labeled as type is clearly of the spe- tegument smooth and transparent bronze cies described here as mannii. Though in scales usually dominate cephalic area. 1901 the Peckhams spelled the name man- Clypeus covered with white scales, but at ii, the collector's name (Mann) and their least in coastal females the area between subsequent spelling (1909) indicate their the AMEs is covered with orange scales. intention to spell the name mannii. Legs with light to dark brown markings Diagnosis. The common species of oak in coastal females, not distinctly annulate. woodland of the Pacific coastal United Abdomen in coastal females brown with States. Dense white patches on chelicerae prominent paired dark spots; Arizona fe- and cheek bands that contrast against a males may have the abdomen partly cov- dark, shiny body distinguish males im- ered with yellow scales. Measurements: mediately. The smooth carapace, weak Body length 4.3(4.5)4.9 mm; carapace epigynal flaps, and orange scales between length 1.9(1.9)2.0 mm; width/length = the AMEs distinguish females from Pacific 077(0.79)0.80; n 5$ from California. Coast Pelegrina. Epigynal flaps shorter Geographical Variation. Two distinct than in diplacis, more robust than in chera forms might be recognized, an inland form and carmenensis. Female markings less (mannii s.s., in Arizona; Figs. 180, 181, longitudinally arranged than in diplacis 230, 498-502) and a coastal form (versi- 312 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

color, from California to British Columbia; Distribution (Map 31.) British Colum- bia south to California and east to Figs. 178, 179, 229, 493-497). In the coast- Baja al form, males are dark brown and gen- central Arizona. erally lack white side bands on the cara- Records. Many specimens, especially in CAS, pace, and the retrolateral side of the base AMNH, and MCZ: Form mannii: UNITED STATES: of embolus is little if at all prolonged into UTAH; Zion National Park (1<3); ARIZONA: Cochise Co.: Chiricahua Mtns. Coconino Co.: Mormon a spike. Females are medium to dark brown (19); Lake (1$); Pima Co.: Santa Catalina Mtns., 12.7 km and have more or less parallel epigynal from Tuscon on Catalina highway toward Mt. Lem- flaps. In the inland form, males have dense mon (82); Tuscon (19); Santa Cruz Co.: Sycamore the re- white side bands on the carapace, Canyon, 14 km W Pena Blanca Lake (56 89); 1.6 km trolateral portion of embolar base pro- S of Pena Blanca Lake {IS). Form versicolor: CAN- ADA: BRITISH COLUMBIA: Vancouver Island: longed into a pronounced spike, almost as Wellington, Mt. Benson. UNITED STATES (county in diplacis though projecting more parallel records): WASHINGTON: Asotin, Chelan, King, to axis of Arizona females are often palpus. Klickitat, Thurston, Whatcom, Whitman; IDAHO: covered with yellow scales (and could be Adams, Lemhi; OREGON: Benton, Douglas, Hood confused with chera except for their stron- River, Jackson, Josephine, Klamath, Lane, Malheur, Marion, Multnomah, Polk; CALIFORNIA: Alameda, ger epigynal flaps) and have more robust Amador, El Dorado, Fresno, Kern, Humboldt, Lake, and flaps. A variable divergent epigynal Los Angeles, Marin, Mariposa, Mendocino, Monterey, population including pale and dark 99 has Placer, Plumas, Riverside, San Benito, San Bernar- been found near Tuscon, Arizona. Until dino, San Diego, San Luis Obispo, San Mateo, Santa Santa Cruz, Shasta, better evidence is found to distinguish Clara, Siskiyou, Solano, Stanislaus, Tulare, Ventura, Yuba; MEXICO: the two forms will be considered as Trinity, BAJA them, CALIFORNIA: 12 km S Santo Tomas. one species. Chromosomes. 2n(5 = 26 acrocentrics + Natural History. Form versicolor col- XXO (25 from Apple Canyon, Riverside lected from oaks (9 records), including Co., California). Quercus agrifolia, Q. douglasi, Q. kellog- Courtship (53 observed from near No- gii, and Q. wislezenii, Arctostaphylos (4 record gales, Arizona, and Riverside Co., Califor- records), pine (2 records), and one nia). Males of both the inland form (Ari- each from Ribes, willows, Adenostema, zona) and the coastal form (California) holly, and raspberry, at elevations from 15 have a typical crouch display. Raised- to 100 m (8 records), 100 to 1,000 m (8 = = to spread (n 7, 3(3). Crouch (Arizona; n records), and 1,000 1,500 m (5 records). = 9, 33; California: n 13, 25): Body low (n Form mannii collected from oaks (5 re- = 7, 35). First legs forward and bowed, cords) and Cerocarpus (1 record), at ele- = = horizontal (n 18, 45) or raised (n 1), vations from 1,200 to 1,700 m (4 records). on series flickered legs with high frequen- = = 41 . Metaphidippus diplacis cy (n 13, 25) low amplitude (n 19, 45) (Chamberlin, 1924) while legs are pushed a bit closer together ^ = Figures 182, 183, 231, 503-508; Map 33 (n 6, 25) or not (n 3, 15). First legs = moved closer as he got closer (n 13, 25) Dendryphantes diplacis Chamberlin, 1924: 686, figs. = S. in with label "Den- until tips almost touching (n 10, 15), legs 130-132, Holotype MCZ IS = dryphantes diplacis Chamb., S holotype [in faded motionless on pause (n 3, 15). Palpi hang- red ink], Cal.: near San Diego, R. V. Chamberlin ing extended forward, down and to side = Coll. 1049," examined. Roewer, 1954: 1193. Bon- (n 1, 35), or just hanging down to side net, 1956: 1393. — 1951: (n 3, 15); on series flickering slightly (n Metaphidippus franciscanus Schenkel, 39, figs. = 9. material in Naturhistorisches Mu- 22, 55) at high frequency low amplitude 42a, b, Type = seum, Basel, from Mission Bay near San Diego, (n 13, 25), or only slightly as palpi pushed = California. NEW SYNONYMY. forward (n 3, 15), palpi motionless on — 1954: 1210. = Dendryphantes franciscanus: Roewer, pause (n 6, 25). Repertoires: 25 crouch Metaphidippus diplacis: —Richman and Cutler, 1978: only; 35 raisedspread and crouch. 89. Pelegrina Jumping Spiders • Maddison 313

Notes on Synonymy. (1) The type ma- Flaps usually at least half as long as epi- terial of M. franciscanns Schenkel remains gynum, generally longer than other man- to be examined; however, the synonymy nii group 29. Epigynal surface more or less is clear on the basis of his description and flat. Notch usually narrow. Markings (Figs. subsequent collecting at Mission Bay, 183, 507, 508): Carapace orange-brown where M. diplacis is very common. (2) The with bronze scales, covered with yellowish Peckhams' (1909) figures of the female of white scales densest on upper sides, giving covered Dendryphantes imperialis (pi. 37, figs. 2, hint of side bands as in S. Clypeus 2a) may actually be of M. diplacis. densely with white scales; between ante- Diagnosis. Among specimens of the rior eyes are usually orange-brown setae. mannii group collected along the Pacific Abdomen with somewhat lineate mark- Coast, only M. diplacis has the retrolateral ings, with pale medial band flanked by dark basal edge of the embolus so prolonged light brown with paired elongate (though inland M. mannii are similar in spots. Measurements: Body length this). males differ 5.0(5.3)6.2 mm; carapace length 2.0(2.1)2.3 Metaphidippus diplacis = from mannii in having more extensive side mm; width/length 0.78(0.79)0.80; n 59 bands and much weaker white patches on from California. = the chelicerae; females by the more lineate Chromosomes. 2n(5 26 acrocentrics + abdominal markings and the longer epi- XXO (26 from San Diego, California). from San gynal flaps. Metaphidippis diplacis can be Courtship (2$ observed Diego separated from the more northerly but and Santa Barbara Cos., California). No similar tricolor by the wider embolus, more apparent crouch display seen. Raised- = robust tibial apophysis, the shinier body in spread (n 24, 23): First legs waving ir- = = both and the more extensive white (n 8, 1(3) up and down (n 8, sexes, regularly = markings in males, and darker epigynal 23) with high amplitude (n 5, 13). Palpi and flaps. waving at low amplitude irregularly = horizontal = Male. Palpus (Figs. 231, 504, 505): Em- (n 5, 13). Abdomen (n 5, = he bolus blade-shaped, thin in ventral view 13) or down and trailing (n 3, 13). As and wide in lateral. Embolic base with closer lowered into reach got legs gradually = sclerotized retrolateral projection. Tibial with no discrete crouch display (n 21, ^ with apophysis robust. Markings (Figs. 182, 23). Reach (n 8, 23): Short stage 503): Body brown, with bronze sheen. gradual transition from raisedspread (not = White carapace side bands extending discrete crouch) (n 8, 23). First legs for- backward almost to margin. ward and parallel, waving alternately but posterior = = Cheek band broad and short, mostly fused irregularly (n 5, 13). Palpi forward (n with side band. Clypeus brown. Forehead 5, 13). Repertoires: 26 raisedspread only. Pacific Coast of band lacking or rudimentary. Setae sur- Distribution (Map 33). rounding AMEs dark except for a few southern California and Baja California white scales laterally. Chelicerae with Norte. patch of pale scales restricted to basal half, CALIFORNIA: Los as in tricolor, but generally white. Palpus Records. UNITED STATES: Beach brown with scattered white setae near end Angeles Co.: South Huntington {16, AMNH); Orange Co.: Laguna Beach 117.47/33.33 (16, AMNH); of femur. Abdomen brown dorsally with San Diego Co.: E of Lake Hodges, Escondido (12, two lon- paired dark brown spots forming MCZ); Oceanside (29, MCZ); San Diego, Mission Bay, near San gitudinal lines; side bands complete. Mea- Fiesta Island (43 219, MCZ; 2<5 29, UCB); San Luis Co.: Pismo Beach surements: length 3.5(4.4)4.6 mm; Diego (6(3, MCZ); Obispo Body El Estero {1$, UCB); Santa Barbara Co.: NW edge of carapace length 1.9(2.1)2.2 mm; width/ marsh just E of Carpinteria (13 5$, MCZ); Gaviota n = 56 from Cali- length 0.77(0.80)0.81; (29, AMNH); Goleta (IS, AMNH). MEXICO: BAJA fornia. CALIFORNIA NORTE: Arroyo Soccorro dunes, S of En- Female. Epigynum (Figs. 506, 507): San Quintin (19, UCB); El Rosario (29, AMNH); 314 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

senada (1<5, AMNH); Rancho Las Parritas, 16 km S tween diplacis and chera. Whether or not of San Quintin (4<5 39, UCB); Santa Maria, 37 km S tricolor grades into diplacis in the south is of Colonia Guerrero (19, AMNH); San Telmo de Ar- not now clear. Females are notable for their riba (19, AMNH); Santo Tomas (19, AMNH). lineate markings; males for the dark face. Natural Baccharis re- History. On (3 Male. Palpus (Figs. 232, 510): Embolus Fiesta San and dunes cords. Island, Diego) narrow, though slightly wider than in (2 records, BCN). Appears to be restricted chera, and sclerotized retrolateral projec- to localities near the seashore; not found tion on embolar base better developed. Te- so far inland as mannii. gulum bulbous prolaterally, in which re- spects it approaches diplacis. Markings dark brown with 42. Metaphidippus tricolor (Figs. 184, 509): Body white side bands often Chamberlin & Ivie, 1941 poorly developed. Cheek band broad but weak, fused to side Figures 184, 185, 232, 509-513; Map 32 band. Clypeus brown. Forehead band tricolor Chamberlin and 1941: Metaphidippus Ivie, lacking. Setae surrounding AMEs dark ex- 29, figs. 30-32, S. Type in AMNH 16 from 122°5'W, cept for a few white scales Che- 37°5'N, Ben Lomond, California. laterally. licerae with Dendryphantes iviei Roewer, 1951: 453 (n. nov. for inconspicuous orange-brown tricolor Chamberlin and Ivie, junior secondary patch of pale scales restricted to basal half. homonym of Plexippus tricolor C. L. Koch, 1846, Palpus dark, with few white scales and both in Roew- placed Dendryphantes by Roewer). none on cymbium. Legs dark, with indis- er, 1954: 1212. tinct annulations. Abdomen brown dor- Notes on Synonymy. The name tricolor sally with two longitudinal dark bands. is maintained despite the ICZN's rule (1985 Measurements: Body length 3.5(4.1)4.6 code Art. 59(b)) that junior secondary mm; carapace length 1.7(1.9)2.3 mm; = homonyms rejected before 1961 must re- width/length 0.80(0.80)0.82; n 5<5 from main rejected. The placement of almost Monterey Co., California. all New World dendryphantines into Den- Female. Epigynum (Figs. 511, 512): dryphantes was a practice mostly of cat- Flaps long, lightly pigmented. Epigynal aloguers (Petrunkevitch, 1911; Roewer, surface more or less flat. First curve of 1954) and not of practicing North Amer- ducts long, pale. Notch broad. Markings ican systematists. Roewer's new name and (Figs. 185, 513): Body scales dull, not shiny the placement of the two tricolors together as in diplacis. Carapace surface not as shiny in Dendryphantes are better considered as mannii, with brown or gray scales above, temporary anomalies rather than long-ac- darker than chera. Clypeus densely cov- cepted changes that need to be protected ered with white scales; between anterior by the code, for neither his new name iviei eyes are orange-brown setae. Abdominal nor its placement in Dendryphantes have markings strikingly linear, central pale been since accepted (e.g., Richman and stripe flanked by black stripes flanked by Cutler, 1978). Plexippus tricolor C. L. lateral pale stripe. Measurements: Body Koch, at least by Koch's figures, appears length 4.3(5.2)5.4 mm; carapace length to be near Eris aurantia and, thus, not now 1.9(1.9)2.0 mm; width/length 0.78(0.79) = considered congeneric with the marinii 0.82; n 59 from Monterey Co., Califor- group. Until a generally accepted second- nia. = ary homonymy occurs, it serves little pur- Chromosomes. 2n(3 ? -I- XXO (2(3 from pose to allow Roewer's changes to return Lucia, California). and haunt us, and so tricolor Chamberlain Courtship (23 observed from Monterey & Ivie will be maintained. Co., California). With typical crouch dis- = Diagnosis. A dark species restricted to play. Raisedspread (n 7, 23). Crouch (n = = the coast of central and northern Califor- 6, 3(5): Body low-normal (n 5, 23). First = nia, in some respects intermediate be- legs forward and horizontal (n 6, 33), Pelegrina Jumping Spiders • Maddison 315

= = spread slightly (n 1), or bowed (n 5, ings, annulate in males, a narrower = legs 2(3) though may sometimes be raised (n embolus, narrower tegulum, thinner tibial = 4, 1(5); waved little if at all (n 3, 1(5) or apophysis, and weaker, shorter, and de- tips of legs flickered at high frequency low pigmented epigynal flaps. The female ab- = = amplitude (n 2, 16). Palpi down (n 3, dominal markings are never so lineate as series = 1(5) waved/flickered on each (n 6, in tricolor. The scales covering the female = 3(5) rapidly (n 2, 1(5). Repertoires: IS carapace are not shiny as in mannii or raisedspread only, 2(5 crouch only, 1(5 ra- diplacis, nor as dark as tricolor. Metaphi- isedspread and crouch. dippus chera is perhaps most likely con- Distribution (Map 32). Pacific Coast of fused with carmenensis but differs in hav- central and northern California. ing the embolus straighter in retrolateral view, and the left and ducts Records. UNITED STATES: CALIFORNIA: Mar- right epigynal at midline before in Co.: Pt. Reyes {IS 22, UCB); North Beach, Pt. Reyes meeting going posteri- National Seashore (23 42, MSUW, UCB); Monterey orly. Though the male is easily distin- Co.: Hastings Natural History Reserve (1<5 12, AMNH); guished from that of emmiltus by mark- 12.2 km N of Lucia on 1 (73 92, MCZ); on Hwy ings and cheliceral size, the female is much Nacimiento-Fergusson Road 0.3-1.4 km from Hwy like that of emmiltus but the epigynal ducts 1 (1(5 22, MCZ); ocean-facing slopes of Santa Lucia are wider and meet at midline before Mtns., 5 km NW of San Luis Obispo State Border on go- Hwv 1 (13 22, MCZ); Pacific Grove, 121.55°W, 36.38°N ing posteriorly. Pebble Beach (12, AMNH); 8 km N of (12,'aMNH); Male. Palpus (Figs. 33, 233, 515-523): Point Sur (13, AMNH); Santa Cruz Co.: Ben Lomond, Embolus thin and straight; usually lacking 122.05''W, 37.05°N (2 imm, AMNH); Trinity Co.: 72 sclerotized on retrolateral side km W of Redding (13, AMNH), projection of base embolar base, though this varies Natural On Baccharis and oth- History. considerably (Figs. 518-523). Tegulum er shrubs in coastal scrub records, Mon- (4 fairly narrow, not bulbous prolaterally. Co.) and from on beach (2 terey Lupinus Markings (Figs. 186, 514): Carapace with Marin records, Co.). strong side bands often with thoracic pro- jections toward fovea. Cheek band usually 43. Metaphidippus chera distinct from side band. Clypeus with or- (Chamberlin, 1924) ange-brown scales; some white setae over- new combination hanging chelicerae. Forehead band does Figures 33, 186, 187, 233, 257, not reach AMEs; setae surrounding AMEs 35 514-528; Map orange-brown except laterally. Chelicerae with prominent white patch extending Dendryphxintes chera Chamberlin, 1924: 683; fig. 124, more than half of chelic- 2, Holotype in CAS 12 with labels "Dendryphantes usually length chera Chamb., 2 type, San Joseph Id. 6/10/21, 165 erae. Cymbium dark brown, often with a C. Chamberlin" and examined. Cham- J. "1462," few white scales. Legs strongly annulate. berlin cites the type locality as San Josef Island, Abdomen with strong side bands; dorsally Gulf of California. I interpret this as San Jose Island, variable as in either solid Baja California Sur. Roewer, 1954: 1192. Bonnet, females, light 1956: 1393. — brown or with brown spots, which are Metaphidippus manni: Carpenter, 1972: 163. sometimes fused into longitudinal dark Richman and Roth, 1976: 201. Gertsch and Riech- bands flanking central pale stripe. Mea- ert, 1976: 7. surements: Body length 3.4(4.3)4.8 mm; Diagnosis. One of the most common sal- carapace length 1.6(2.0)2.3 mm; width/ = ticids in the southwestern United States length 0.81(0.82)0.84; n 5<5 from New and northern Mexico, this species is usually Mexico. identified as mannii. Metaphidippus chera Female. Epigynum (Figs. 257, 524-526): can be easily distinguished from mannii, Flaps depigmented and short. Epigynal as well as from diplacis and tricolor, in surface flat. Ducts meet at midline at junc- having much more extensive pale mark- tion of second and third curves. Markings 316 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

= (Figs. 187, 527, 528): Very variable, in some sometimes trailed a bit on sidles (n 2, populations solid yellow, in others darker 23). Repertoires: 13 raisedspread only, 123 with annulate legs and dotted or somewhat crouch only, 13 raisedspread and crouch. lineate abdominal markings. Carapace Distribution (Map 35). Texas west to covered with mostly white to yellow-white California, Nevada south to Baja Califor- scales, not shiny. Clypeus covered densely nia del Sur and San Luis Potosi. with yellow-white scales. Legs uniformly Records. Many specimens in AMNH, MCZ, UCB, yellow to strongly annulate. Abdomen and MSU, from: UNITED STATES (county records); sometimes otherwise var- entirely yellow, OKLAHOMA: Jefferson; TEXAS: Archer, Baylor, iously marked (Figs. 187, 527, 528). Mea- Bexar, Foard, Haskell, Presidio, Reagan, Wichita, surements: Body length 3.4(4.1)5.0 mm; Winkler; UTAH: Washington; NEVADA: Churchill; NEW MEXICO: Dona Ana, Lincoln; ARIZONA: Co- carapace length 1.6(1.8)2.0 mm; width/ = chise, Coconino, Graham, Maricopa, Mohave, Pima, 0.79(0.79)0.84; n 59 from New length Santa Cruz, Yavapai, Yuma; CALIFORNIA; Fresno, Mexico. Imperial, Inyo (nr. Bishop), Kern, Los Angeles, Mono, ChroTTiOSomes. 2n6 = 26 acrocentrics + Riverside, Santa Barbara, San Benito, San Bernardino, San San Luis Ventura. XXO (16 from Imperial Dam, California). Diego, Obispo, Stanislaus, MEXICO: TAMAULIPAS; Victoria; 16 km S of observed from Texas, Rey- Courtship (146 SAN LUIS POTOSI: border on New Nuevo nosa; Guanajuato Hwy Mexico, Arizona, California, 57 (100°45'W, 23''19'N); NUEVO LEON; 41 km NE Leon, San Luis Potosi, and Zacatecas). With of (98°54'W, 25°51'N); COAHUILA: 16 km E 20 strong crouch display. A brief description of Cuatro Cienega; San Pedro; ZACATECAS: km N of Fresnillo 23°19'N); 15 km NE Con- of the courtship was given by Richman (102°57'W, cepcion de Ora; CHIHUAHUA; Las Delicias; 21 km (1982: 38, 1-3), under the name Me- figs. = N of Ciudad Camargo (105°13'W, 27°52'N); 40 km taphidippus manni. Raisedspread (n 6, W of SONORA; 25 km S of Hermosillo; = Camargo; 2(5). Crouch (n 28, 133). Body raised Sonoyta; 1.6 km W of San Carlos Bay; 10 km S of = = CALIFORNIA NORTE; 11 slightly (n 6, 33) or low (n 2, 13) or at Presa, Obregon; BAJA - km SE of Mexicali; Rancho Santa Cecelia nr. El Pro- normal (n 3, 13). First legs for- height San Ranch; San 12 km S wide = even gresso; Jose, Meling Felipe; ward, spread (n 12, 53), great- of Santo Tomas; BAJA CALIFORNIA SUR; Concep- er than 90° apart, especially when male at tion 3 km S of La Paz; 42 km S of Loreto; San = Bay; a distance (n 6, 23) to more or less par- Franciscito Bay; San Jose Island; DURANGO; Du- = allel (n 9, 33), especially when close to rango. = female (n 6, 23). First legs horizontal (n = Natural History. Common on desert 20, 103), sometimes with tips on ground = = vegetation, including mesquite, tamarisk, (n 3, 13), or slightly raised (n 7, 33). = Acacia, creosote bush, oaks, and Chilopsis. On series, legs flickered (n 17, 93) with = Elevations recorded from -70 to 1,000 m low amplitude (n 10, 53) and high fre- = (5 records), 1,000 to 1,500 m (5 records), quency (n 3, 13); on pause, legs motion- = and 1,500 to 2,100 m (4 records), though less (n 17, 93). On series, legs pushed = these may not be representative because medially together (n 7, 43), raised slight- = = elevations are probably often not recorded ly (n 3, 23), and pushed forward (n 2, = for lowland localities. Where living on the 13). Sometimes legs not raised (n 1) or same hillside with mannii in Arizona, there not pushed together, rather kept parallel on = is a clear division in habitat: M. chera (n 4, 23), espeically when close to female = mesquite and other typically desert shrubs (n 2, 13). Sometimes legs held slightly and trees and M. mannii on oaks. asymmetrically, one more extended than = = other (n 2, 13). Palpi down (n 9, 53), = 44. Metaphidippus carmenensis and over chelicerae (n 1) or curled to = (Chamberlin, 1924) side (n 1), on each series pushed forward - = new combination (n 10, 43) and waved (n 21, 103), up = = Figures 188, 189, 234, 529-533; IVlap 34 and down (n 5, 33); still on pause (n 21, 103). Abdomen bobbed occasionally (n Dendryphantes carmenensis Chamberlin, 1924: 682, = = S. in CAS 1<3 and 4, 23), specifically after series (n 1), figs. 122, 123, Type (its right Pelegrina Jumping Spiders • Maddison 317

in MCZ) with labels car- palpus "Dendryphantes (Figs. 189, 533): Carapace covered with menensis Chamb., $ holotype, Carmen Id. 6/16/ " white scales. Clypeus covered densely with 21. #177 J. C. Chamberlin and "1461." Cham- white scales. Ab- berlin reports the t> pe locality as Salinas Bay, Car- Legs uniformly yellow. men Island, Gulf of California. Roewer, 1954: 1 192. domen uniformly pale, with white scales, Bonnet, 1956: 1392. in northern females; with paired dark Dendryphantes imperialis: —Chamberlin, 1924, in brown spots in southern females. Mea- 681 de la Guarda and San part: (Isla Angel Jose surements: Island records). Body length 4.1(4.8)5.6 mm; Dendryphantes chera: —Chamberlin, 1924, in carapace length 2.0(2.0)2.4 mm; width/ part: = 683 (San Diego Island record). length 0.79(0.79)0.83; n 59 from Baja California Sur, Baja California Norte, and Diagnosis. Much like chera, but with California. more curved embolus and left and right Geographical Variation. Northern epigynal ducts failing to meet at midline specimens (California; Baja California at junction of second and third curves. Norte; Sonora) are large and pale, es- Northern specimens are further distinct by pecially 33, whose faces are covered with their extensive covering of pale scales. white. Southern form (Baja California Sur) Male. Palpus (Figs. 234, 530): Erect por- is smaller and darker, almost indistinguish- tion of embolus thin, curving strongly to- able from chera except by genitalia. ward the ventral. Markings (Figs. 188, Courtship (23 from Imperial Co., Cal- 529): Very pale with dense covering of ifornia). The five displays observed showed white and orange scales in northern males, only an apparently low-intensity raised- = though southern males darker. Carapace spread stage. Raisedspread (n 5, 23): First = in northern males covered mostly with legs waved slowly and irregularly (n 5, = white except orange around eyes and in 23) up and down (n 1); as he got closer middle of thorax; in southern males marked legs moved more parallel until he reached = more as in chera. Clypeus of northern 3 to touch her (n 1). densely covered with white scales except Distribution (Map 34). Baja California orange immediately under AMEs; south- and Sonora extending north into California ern males darker, only white setae are those and Arizona. overhanging chelicerae. Setae surrounding Records. UNITED STATES: CALIFORNIA: Riv- AMEs entirely orange in northern males; erside Co.: Desert Beach (IS, AMNH); Desert Beach some white scales in southern laterally Cmpgd (25 59, MCZ); ARIZONA: Maricopa Co.: males. of Chelicerae with patch white Wickenburg (19, AMNH). MEXICO. BAJA CALI- Isla de la scales, patch very broad in northern <5. FORNIA NORTE: Angel Guarda (SS 19); San Felipe (133 229, AMNH); BAJA CALIFORNIA Cymbium beige to light brown dorsally SUR: La Burrera, 19 air km ENE of Todos Santos and with white scales, darker brown on (4

R[oad] 219, 6.0 mi [9.7 km] N of Pastura 5500 ft. surface flat. Second curve of left and right el. 22 D. WPM#83- [1,680 m], Sept. 1983. Richman, ducts do not meet at midline; bend be- 173 on juniper." tween second and third curves abrupt; sec- Etymology. After the Greek emmiltos, ond and third curves narrow. Markings referring to the reddish scales around the (Figs. 177, 538): Carapace covered with eyes. beige to tan scales. Clypeus densely cov- Diagnosis. A beautiful species living on ered with white scales. Legs more or less juniper, bearing superficial resemblance to uniformly beige to light brown. Abdomen the pervaga group of Pelegrina. The white- tan to light brown, dorsally darker and fringed legs, dense and distinct marginal with paired white spots; fourth pair of spots and side bands on the carapace, and red- unusually large. Measurements: Body ringed anterior median eyes are distinc- length 4.2, 4.5, 4.8 mm; carapace length tive. The female differs from that of chera 1.9, 2.0, 2.1 mm; width/length 0.79, 0.81, in the failure of the second curves of the 0.82; n = 39 from New Mexico. epigynal ducts to meet at the midline, the Geographical Variation. Males from large fourth pair of white spots on the ab- New Mexico and California differ in car- domen, and in New Mexican specimens apace shape and forehead markings, as al- the swollen carapace behind the anterior ready noted. lateral eyes. The bend between the second Distribution (Map 36). New Mexico west and third curves is more distinct than in to southern California. carmenensis. Records. UNITED STATES: NEW MEXICO: Male. 235, 535): Erect Palpus (Figs. por- Guadalupe Co.: along SR 219, 9.7 km N of Pastura, thin. tion of embolus Carapace: Bulges 22 September 1983, on juniper (2<5 12, MCZ); Lincoln 21 slightly at ALEs and narrowed behind Co.: T6N R6E S24, June 1974, beating junipers T6N RIOE 24 1971 ALEs in New Mexico males. Markings (19, AMNH); S25, May {16, AMNH); Sandoval Co.: northwest of Bernalillo (15, (Figs. 176, 534): Generally yellowish. Car- AMNH); Santa Fe Co.: 16 km S of Santa Fe (13 19, with distinctive black on fore- apace stripe AMNH); CALIFORNIA: Los Angeles Co.: 1.6 km W head in New Mexico males, with V-shaped of Desert Springs, 1 June 1957, montane forest (13, white forehead band in California males. AMNH); Palmdale, 5.6 km S of Hwy 6, 26 May 1957, woodland, creosote bush scrub {IS 19, AMNH). Cheek band long and marginal, separated juniper from side band by band of dark hairs. LITERATURE CITED Clypeus orange-brown. Setae surrounding 1792. of the Insects of AMEs red; entirely red in New Mexico Abbot, J. Drawings Georgia, in America, Vol. 14, manu- males, in California males red with some Spiders. Unpublished script in the British Museum, London (copy seen). white scales and where forehead laterally Alay6n, G. 1982. Redescripcion de Dolomedes band contacts AMEs dorsally 10:30-12:30. fuscus Franganillo (Arachnida: Araneae: Pisaur- Chelicerae yellow-brown with orange idae). Poeyana, 250: 1-7. Banks, N. 1892. The fauna of the Upper scales except for small medial basal spot spider Cayuga Lake Basin. Proceedings of the Academy of white scales. Palpus pale yellowish, with of Natural Sciences, Philadelphia, 1892: 11-81, dense of white scales on femur. patch Legs 5 pis.

yellowish, with white fringe on first pair. . 1895. Some new Attidae. Canadian Ento- 27: 96-102. Abdomen brown dorsally, paler centrally mologist, 1921. New Californian spiders. Proceed- and with white side band and fourth pair ings of the California Academy of Sciences, 4th of white Measurements: spots prominent. series, 11: 99-102. Body length 3.6, 3.7, 3.7, 3.8 mm; carapace Barnes, R. D. 1955. North American jumping spi- American Museum length 1.8, 1.8, 1.9, 1.9 mm; width/length ders of the genus Maevia. = Novitates, 1746: 1-13. 0.79, 0.79, 0.80, 0.80; n 43 from New

, 1958. North American of Mexico. jumping spiders the subfamily Marpissinae (Araneae, Salticidae). Female. Epigynum (Figs. 536, 537): American Museum Novitates 1867: 1-50. B. A. Scares. 1982. Flaps weak and depigmented. Epigynal Bauab-Vianna, M. J., AND M. Pelecrina Jumping Spiders • Maddison 319

Contribuiqao ao estiido dos Salticidae (Araneae) collected by L. W. Saylor and others, mostly in do Brasil. IX. Revista Brasileira de Entomologia, California. Bulletin of the University of Utah, 26(1): 87-91. Biological Series, 31(8): 1-49.

R. D. .and -. Bhatnag.^r, S., J. G. Rempel. 1962. The 1944. Spiders of the Georgia region of North structure, function and postenibryonic develop- America. Bulletin of the University of Utah, Bi- ment of the male and female copulatory organs ological Series, 35(9): 1-267. of the black widow spider Latrodectus curaca- Chickering, a. M. 1944. The Salticidae (jumping viensis (Miiller). Canadian Journal of Zoology, spiders) of Michigan. Papers of the Michigan 40: 465-510. Academy of Sciences, Arts and Letters, 29: 139- Blest, A. D. 1983. Ultrastructure of secondary ret- 222.

inae of primitive and advanced jumping spiders . 1946. The Salticidae (spiders) of Panama. (Araneae, Salticidae). Zoomorphology, 102(2): Bulletin of the Museum of Comparative Zoology, 125-141. 97: 1-474. Blest, A. D., .and C. Sigmund. 1984. Retinal mo- Chickering, A. M., and G. Bacorn. 1933. Notes saics of the principal eyes of two primitive jump- and studies on Arachnida. V. Additions to the list ing spiders, Yaginumanis and Lyssotnanes. clues of Araneae from Michigan. Papers of the Mich- to the evolutin of Salticid vision. Proceedings of igan Academy of Sciences 17: 521-528. the of Royal Society London B, 221: 111-125. Crane, J. 1949a. Comparative biology of Salticid Bonnet, P. 1955-59. Bibliographia Araneorum, A-B spiders at Rancho Grande, Venezuela. Part III. (1955), C-F (1956), G-M (1957), N-S (1958), T-Z Systematics and behavior in representative new (1959). Analyse Methodique de toute la literature species. Zoologica, 34(2): 31-52.

araneologique jusqu'en 1939. Toulouse, Paris. . 1949b. Comparative biology of Salticid spi- Brignoli, P. M. 1983. A catalogue of the Araneae ders at Rancho Grande, Venezuela. Part IV. An described between 1940 and 1981. Manchester: analysis of display. Zoologica, 34(4): 159-214. Press. Manchester Univ. 755 pp. Curtis, J. L. 1892. A new . Zoe, 3: Bryant, E, B. 1940. Cuban spiders in the Museum 332-337. of Comparative Zoology. Bulletin of the Museum Cutler, B. E. 1979. Variation in the embolus of of Comparative Zoology, 86: 259-532. Metaphidippus insignis (Banks) (Araneae: Sal-

. 1941. Notes on the spider fauna of New ticidae). New York Entomological Society, 87: England. Psyche, 48: 129-146. 270-274.

. 1943. The salticid spiders of . . 1981a. A revision of the spider genus Par- Bulletin of the Museum of Comparative Zoology, adamoetas (Araneae, Salticidae). Bulletin of the 92: 445-522. American Museum of Natural History, 1 70: 207-

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letin of the Museum of Comparative Zoology, . 1981b. Key to late instar immatures of 103: 163-209. Metaphidippus and Eris (Salticidae) in Minne- Cambridge, F, O. P. 1901. Arachnida-Araneida, sota. Peckhamia, 2(2): 31-32. pp. 173-312. In F. D. Godman and O. Salvin -. 1987. A revision of the American species (eds), Biologia Centrali- Americana, Vol. 2. Lon- of the antlike jumping spider genus Synageles don. (Araneae, Salticidae). Journal of Arachnology, Carpenter, R. 1972. The jumping spiders (Salti- 15(3): 321-348. cidae) of Wichita County, Texas. The South- Cutler, B. E., and D. T. Jennings. 1985. A re- western Naturalist, 17(2): 161-168. vision of the Metaphidippus arizonensis group Chamberlin, R. V. 1924. The spider fauna of the (Araneae, Salticidae). Journal of Arachnology, shores and islands of the Gulf of California. Pro- 13(): 1-8. ceedings of the California Academy of Sciences, DoNDALE, C. D. 1961. Life histories of some com- 12: 678-694. mon spiders from trees and shrubs in Nova Scotia.

. 1925a. Diagnoses of new American Arach- Canadian Journal of Zoology, 39: 777-787. nida. Bulletin of the Museum of Comparative Eakin, R. M., and J. L. Brandenburger. 1971. Zoology, 67: 211-248. Fine structure of the eyes of jumping spiders.

. 1925b. New North American spiders. Pro- Journal of Ultrastructure Research, 37: 616-663. ceedings of the California Academy of Sciences, Edwards, G. B. 1977. Comments on some genus (4) 14(7): 105-142. and species problems in the Salticidae, including Chamberlin, R. v., AND W.J. Gertsch. 1929. New Walckenaerian names. Peckhamia, Gainesville, spiders from Utah and California. Journal of En- 1(2): 21-23.

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. 1930. On fifteen new North American spi- hamia, Gainesville, 2(1): 11-14.

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Franganillo Balboa, P. 1930. Aracnidos de Cuba. Jennings, D. T. 1973. Egg retreat of Metaphidip- Mas aracnidos nuevos de la Isla de Cuba. Me- pus arizonensis (Peckham) (Araneae: Salticidae) morias del Institute Nacional de Investigaciones in a hollow stem. Entomological News, 84: 317- Cientificos, Havana, 1: 47-99. 320.

. 1936. Los Aracnidos de Cuba hasta 1936. Jung, A. K. S., and V. D. Roth. 1974. Spiders of Havana: Cultural. 180 pp. the Chiracahua Mountain Area, Cochise Co., Ar- Galiano, M. E. 1963. Las especies americanas de izona. Journal of the Arizona Academy of Sci- aranas de la familia Salticidae, descriptas por ences, 9: 29-34. los in Eugene Simon. Redescripciones basadas en K ASTON, B. J. 1945. New spiders the group Dion- ejemplares tipicos. Physis (Buenos Aires), 23(66): ycha with notes on other species. American Mu- 273-470. seum Novitates, 1290: 1-25.

. 1980. Catalogo de los especimenes tipicos . 1948. Spiders of Connecticut. Bulletin of de Salticidae (Araneae) descriptos por Candido the Connecticut State Geological and Natural F. de Mello-Leitao. Primera Parte. Physis (Bue- History Survey, 70: 1-874. nos Aires), Secc. C, 39(96): 31, 40. 1973. Four new species of Metaphidippus,

Gertsch, W. J. 1934. Further notes on American with notes on related jumping spiders (Araneae: spiders. American Museum Novitates, 726: 1- Salticidae) from the eastern and central United 26. States. Transactions of the American Microscop-

. 1935. Spiders from the southwestern Unit- ical Society, 92: 106-122. ed States, with descriptions of new species. Amer- Keyserling, E. G. 1884. Neue Spinnen aus Amer- ican Museum Novitates, 792: 1-31. ika. VI. Verhandlungen. Zoologisch-botanische

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Maddison, W. P., .\nd G. E. Str.^tton. 1988. A . 1973a. Revision of the spider genus St<

American Museum Novitates, 1257: 1-14. . 1973b. Systematic studies on east palearctic OsTERLOH, A. 1922. Beitrage zur Kenntnis des Ko- Salticidae, II. Redescriptions of Japanese Salti- pulationsapparates einiger Spinnen. Zeitschrift cidae of the Zoological Museum in Berlin. An- fiir wissenschaftliche Zoologie, 119: 326-421. nales Zoologici (Warsaw; Polska Akademia Nauk, Peckham, G. W., and E. G. Peckham. 1883. De- Instytut Zoologiczny), 30(5): 97-128.

scriptions of new or little known spiders of the . 1976. Studium systematyczno-zoogeograf- family Attidae, from various parts of the United iczne nad rodzina Salticidae (Aranei) Regionow States of North America. Milwaukee, Wisconsin. Palearktycznego i Nearktycznego. Wyzsza Szko- 33 pp. la Pedagogiczna w Siedlcach Rozprawy 6: 1-260.

. 1888. Attidae of North America. Trans- 1980. Revision of the spider genus Sitticus actions of the Wisconsin Academy of Sciences, Simon, 1901 (Aranei, Salticidae), IV. Sitticus Arts and Letters, 7: 3-104. floricola (C. L. Koch) group. Annales Zoologici

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lection in spiders of the family Attidae, with some . 1984. Atlas rysunkow diagnostycznych remarks on Mr. Wallace's theory of sexual or- mnief znznych Salticidae [Diagnostic drawings namentation. Occasional Papers of the Natural of less known Salticidae (Araneae)—an atlas.] History Society of Wisconsin, 1(3): 117-151. Siedlce: WSRP. 177 pp.

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. 1901a. On spiders of the family Attidae RiCHMAN, D. B. 1981. A revision of the genus Ha- found in Jamaica. Proceedings of the Zoological hrocestum (Araneae, Salticidae) in North Amer- Society of London, 1901: 6-16. ica. Bulletin of the American Museum of Natural

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SCHENKEL, E, 1951. Spinnentiere aus dem westlich- . 1837. Histoire naturelle des Insectes. Ap- en Nordamerika, gesammelt von Dr. Hans teres. Tome 1. Paris, pp. 1-682. Schenkel-Rudin. Verhandlungen der Naturfor- Wanless, F. R. 1978. A revision of the spider gen- schenden Gesellschaft in Basel, Separatabdruck, era Belippo and Myrmarachne (Araneae: Salti- 62: 24-62. cidae) in the Ethiopian region. Bulletin of the British of Natural SCHULT, J. 1980. Die Genitalstruckturen haplogy- Museum History (Zoology), ner Araneae unter phylogenetischem Aspekt 33: 1-139. (Arachnida). Dissertation, Biol., Univ. Hamburg. . 1984. A review of the spider subfamily Simon, E. 1901. Histoire Naturelle des Araignees. Spartaeinae nom. n. (Araneae: Salticidae) with Bulletin of the Deuxieme edition. Tome 2, fasc. 3. Paris, pp. descriptions of six new genera. 381-668. British Museum of Natural History (Zoology),

. 1903. Histoire Naturelle des Araignees. 46(2): 135-205. G. Rempel. 1959. A Deuxieme edition. Tome 2, fasc. 4. Paris, pp. Whitehead, W. P., and J. 669-1080. study of the musculature of the black widow Steiner, W. W. M., and M. H. Greenstone. 1991. spider, Latrodectus mactans (Fabr.). Canadian Segregation studies of isozyme variation in Me- Journal of Zoology, 37: 831-870. taphidippus galathea (Araneae, Salticidae). Jour- Woods, R. S. 1966. An English-Classical Dictionary nal of Arachnology, 19: 157-160. for the Use of Taxonomists. Claremont, Califor- Walckenaer, C. a. 1805. Tableau des Araneides. nia: Pomona College. Paris. 88 pp.

Index

Listed are the names of genera, subfamilies, and species used in the text. For many names, only the first or most important pages are cited. Boldface indicates primary description of taxon.

Genera and Subfamilies Dendryphantes C. L. Koch, 1837, 227, 231, 232, 235, 242 Admestina G. & E. Peckham, 1888, 226, 239 Dendryphantinae, 226, 227 Admirala G. & E. Peckham, 1901, 227, 241 Descanso G. & E. Peckham, 1892, 226 Agassa Simon, 1901, 227, 243 Donaldius Chickering, 1946, 227 Agobardus Keyserling, 1884, 266 Dryphias Simon, 1901, 227, 236 Amerotritte Mello-Leitao, 1944, 227 Eris C. L. Koch, 1846, 224, 227, 230, 238, 241 Amycus C. L. Koch, 1846, 226 Euophryinae, 226 Anamosa G. & E. Peckham, 1895, 227, 236 Evarcha Simon, 1902, 226 Anicius Chamberlin, 1952, 227, 228, 231, 235, 238, Gastromicans Mello-Leitao, 1917, 227, 232, 233, 234 239, 330 Ghelna new genus, 227, 232, 239 Anoka G. & E. Peckham, 1893, 227 Habronattus F. P.-Cambridge, 1901, 217, 225, 267 Ashtabula G. & E. Peckham, 1894, 220, 227, 233, Harmochirus Simon, 1885, 226 234 Heliophaninae, 226 Avitus G. & E. Peckham, 1896, 227 Hentzia Marx, 1883, 225, 227, 230, 231, 238, 242 Bagheera G. & E. Peckham, 1896, 220, 227, 231, 232, Homalattoides F. P.-Cambridge, 1901, 227, 236 233, 234 Homalattus White, 1841, 227 Balltis C. L. Koch, 1851, 226 Hylltis C. L. Koch, 1848, 226 Beata G. & E. Peckham, 1895, 227, 232, 236, 241, Itata G. & E. Peckham, 1894, 238 242 Leptorchestes Thorell, 1870, 227 Bellota G. & E. Peckham, 1892, 227, 241, 242 Lurio Simon, 1901, 227, 233 Bianor G. & E. Peckham, 1885, 225, 226 Lyssomanes Hentz, 1884, 266 Bryantella Chickering, 1946, 227, 238 Lyssomaninae, 226 Cerionesta Simon, 1901, 227 Mabellina Chickering, 1946, 225, 227, 230 Cheliferoides F. P. -Cambridge, 1901, 227 Maeviobeata Caporiacco, 1947, 227 Chirothecia Taczanowski, 1879, 227 Marengo G. & E. Peckham, 1892, 226 Cocalodes Pockock, 1897, 226 Megatimus Thorell, 1897, 227 Colaxes Simon, 1900, 226 Menemerus Simon, 1868, 238, 239 Consingis Simon, 1900, 226 Messua G. & E. Peckham, 1896, 227, 230-232, 233, Corythalia C. L. Koch, 1851, 267, 330 234 Cydonia G. & E. Peckham, 1893, 227 Metacyrba F. P.-Cambridge, 1901, 266 Pelecrina Jumping Spiders • Maddison 323

Metaphidippus F. P.-Cambridge, 1901, 217, 224, 227, Species 233, 234, 237 Karsch, 1878, 238 Mopsus Valid names are in italics. Myrmarachne MacLeav, 1839, 226 Nagaiiw G. & E. Peckham, 1896, 227, 241, 308 Neon Simon, 1876, 239 aeneola (Curtis, 1892), Pelegrina, 222, 228, 241, 244, Nilakantha G. & E. Peckham, 1901, 266 246, 288 Osericta Simon, 1901, 227 aestivalis G. & E. Peckham, 1883, Attus, 270 Pachyballus Simon, 1900, 226 albeolus (Chamberlin & Ivie, 1941), Phanias, 229, Padilla G. & E. Peckham, 1894, 226 238, 330, 335 Paradamoetas G. & E. Peckham, 1885, 227, 229- alboimmaculata (G. & E. Peckham, 1883), Poulto- 231, 242 nella, 229 Parahentzia Bryant, 1943, 227 albopilosa (Simon, 1903), Gastromicans, 234 Paraphidippus F. P.-Cambridge, 1901, 227 algerina (Lucas, 1846), Cyrba, 328 Parnaenus G. & E. Peckham, 1896, 227, 233, 238 annectans (Chamberlin, 1929), Metaphidippus, 229 Partona Simon, 1904, 227 antillanus Peckham, Peckham, & Wheeler, 1889, Peckhamia Simon, 1901, 226 Lyssomanes, 266 Pelegrina Franganillo, 1930, 224, 225, 227, 240-308 arcuata Franganillo, 1930, Corythalia, 266 Pellenes Simon, 1876, 225, 226 arcuatus Simon, 1902, Sassacus, 237 Pelleninae, 225 arizonensis (G. & E. Peckham, 1901), Pelegrina, 225, Phanias F. P.-Cambridge, 1901, 224, 227, 229, 230, 228, 241-243, 246, 297 232, 234, 238, 240-242 atopodon Chamberlin, 1925, Dendryphantes, 270 Phiale C. L. Koch, 1846, 226 attentus Walckenaer, Attus, 263, 270 Phidippits C L. Koch, 1846, 217, 224, 227, 231, 241, audax (Hentz, 1845), Phidippus, 229, 266, 267, 333, 242 335 Phintella Strand, 1906, 330 aurantia (Lucas, 1833), Eris, 228, 230, 333 Pidegra Simon, 1876, 238 annectans (Chamberlin, 1929), Metaphidippus, 315 Platycryptus Hill, 1979, 266 balia new species, Pelegrina, 244, 246, 390 Plexippinae, 226 barrowsi (Kaston, 1973), Ghelna, 228, 239 Plexippus C. L. Koch, 1846, 226 berlandi Soares & Camargo, 1948, Nagaina, 308 Potdtonella G. & E. Peckham, 1909, 225, 227, 234 bicavatus F. P.-Cambridge, 1901, Metaphidippus, 233 Ramboia Mello-Leitao, 1943, 227, 229, 237 bicuspidata (F. P.-Cambridge, 1901), Pelegrina, 222, Rhanis C. L. Koch, 1848, 227 242, 246, 295 Rhene Thorell, 1869, 227, 236, 238 bifida Banks, 1895, Dendryphantes, 288 Rhetenor Simon, 1902, 227 bispinosus F. P.-Cambridge, 1901, Metaphidippus, Rudra G. & E. Peckham, 1885, 227, 230 310 Salticidae, 226 bivittatus (Dufour, 1831), Menemerus, 266, 267 Salticine Division, 226 bunites new species, Pelegrina, 228, 241, 253, 306, Salticus Latreille, 1804, 226 310 Sassacus G. & E, Peckham, 1895, 227, 230, 235, 237, californicus (G. & E. Peckham, 1888), Terralonus, 238, 243 239, 333 Sebastira Simon, 1901, 227, 233 canadensis (Banks, 1897), Ghelna, 239 Selimus G. & E. Peckham, 1901, 227, 229, 238, 241 capitatus Hentz, 1845, Attus, 263, 270, 284 Semora G. & E. Peckham, 1892, 227 carmenensis (Chamberlin, 1924), Metaphidippus, Sitticus Simon, 1901, 224, 226, 270 222, 316 Spartaeinae, 226 castanea (Hentz, 1846), Ghelna, 239, 333 Synageles Simon, 1876, 226 centralis (G. & E. Peckham, 1896), Messua, 233 Tacuna G. & E. Peckham, 1901, 227 cephalica F. P.-Cambridge, 1901, Beata, 237 Telamonia Thorell, 1887, 226 chaimona new species, Pelegrina, 222, 246, 286 Terralonus new genus, 225, 227, 232, 239 chalceola new species, Pelegrina, 242, 246, 262, 291 Thammaca Simon, 1901, 227, 233 chera (Chamberlin, 1924), Metaphidippus, 222, 228, Thiodina Simon, 1900, 227 315, 317, 331 Thyene Simon, 1885, 226 chuldensis Proszynski, 1982, Dendryphantes, 236 Tulpius G. & E. Peckham, 1896, 227, 240, 242 cinereonitida Simon, 1902, Beata, 237 Tutelina Simon, 1901, 224, 225, 227, 230, 235, 239, clarus (Keyserling, 1885), Phidippus, 229, 231 240, 242 clavator new species, Pelegrina, 274, 300 Uluella Chickering, 1946, 227 clemata (Levi & Levi, 1951), Pelegrina, 225, 228, Wala Keyserling, 1884, 227 244, 246, 284, 287 Zeuxippus Thorell, 1891, 227 concolor (Banks, 1895), Sitticus, 270 Zygoballus G. & E. Peckham, 1885, 225, 227, 231, concoloratus (Chamberlin & Gertsch, 1930), Phanias, 235, 242 238 324 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

crassi venter Keyserling, 1884, Icius, 263 hondurensis (G. & E. Peckham, 1896), Gastromicans cubensis (Franganillo, 1934), Agobardus, 266, 267 234, 263 F. cupreus P.-Cambridge, 1901, Metaphidippus, 234 huachuca new species, Pelegrina, 222, 246, 296 cursor Barrows, 1919, Sitticus, 270 imperialis G. & E. Peckham, 1888, Attus, 310, 317 czekanotvskii Proszynski, 1979, Dendryphantes, 243 incertus (Banks, 1929), Zygoballus, 229, 230, 235 dentata F. P.-Cambridge, 1901, Ashtabula, 228 330, 331, 333 dentiger F, P.-Cambridge, 1901, Messua, 233 inclemens (Walckenaer, 1837), Maevia, 270 desidiosa G. & E. Peckham, 1896, Messua, 233, 335 inconcinna (G. & E. Peckham, 1895), Beata, 237 diademata Simon, 1902, Nagaina, 308 incunda G. & E. Peckham, 1896, Nagaina, 308 digitatus F. P.-Cambridge, 1901, Metaphidippus, 263 inflatus F. P.-Cambridge, 1901, Metaphidippus, 234 diplacis (Chamberlin, 1924), Metaphidippus, 222, 242, insignarius C. L. Koch, 1846, Phidippus, 331 311, 312 insignis (Banks, 1892), Pelegrina, 228, 244, 246, 284 iridescens F, dithalea new species, Pelegrina, Til, 246, 262, 268 P.-Cambridge, 1901, Metaphidippus, dolius Chamberlin, 1925, Anicius, 333 234 iviei dominatus (Chamberhn & Ivie, 1941), Phanias, 238 Roewer, 1951, Dendryphantes, 314 donalda (Kraus, 1955), Messua, 233 juZ?a

nigropictus F. P. -Cambridge, 1901, Metaphidippus, sexmaculata (Banks, 1985), Ghelna, 228, 239 233 sexpunctatus (Hentz, 1845), Zygoballus, 229 nitidus & E. (G. Peckham, 1896), Metaphidippus, shaferi Gertsch & Riechert, 1976, Metaphidippus, 238 239 noxiosa 234 (Simon, 1886), Gastromicans, similis (Banks, 1895), Tutelina, 338 ijoxiosus 331 (Hentz, 1850), Synageles, siticulosus G. & E. Peckham, 1909, Pseudicius, 229, nubilis Hentz, 1846, Attus, 263 241, 333 ochracea P. (F. -Cambridge, 1901), Pelegrina, 246, smithii G. & E. Peckham, 1893, Synemosyna, 266 295 squamata Bryant, 1949, Corythalia, 266 octavus Hentz, 1846, Attus, 270, 284 squamulata Mello-Leitao, 1917, Gastromicans, 234 octonotata (F. 1901), Messua, 234 P.-Cambridge, striata Petrunkevitch, 1925, Beata, 237 octopttnctata (G. & E. Peckham, 1893), Beata, 228, taeniola (Hentz, 1846), Metacyrba, 266, 267 230, 241 taylori (G. & E. Peckham, 1901), Metaphidippus, 238 octopunctatus (G. & E. Peckham, 1883), Phidippus, tenuior (Keyserhng, 1882), Simaetha, 236 231, 243 texanus (Banks, 1904), Metaphidippus, 237 olivacea Franganillo, 1930, Nagaina, 308 fiMa/is (Bryant, 1940), Hentzia, 266 orestes new species, Pelegrina, 228, 241, 244, 307, tti^ia/is F. 231 310 P.-Cambridge, 1901, Zygoballus, tillandsiae (Kaston, 1973), Pelegrina, 222, 228, 241, ornatus Banks, 1892, Dendryphantes, 263 244, 305 ovatus F. P.-Cambridge, 1901, Metaphidippus, 234 tricincta Simon, 1902, Nagaina, 308 paetula (Keyserhng, 1882), Simaetha, 230 tricolor Chamberlin & Ivie, 1941, Metaphidippus, paiutus (Gertsch, 1934), Sassacus, 238 Til, 314 pollens F. P.-Cambridge, 1901, Metaphidippus, 238 tricolor C. L. Koch, 1846, Plexippus, 314 pallidata (F. P.-Cambridge, 1901), Pelegrina, 222, tridentata (F. 1901), Messua, 234 246, 300 P.-Cambridge, tristis new 222, 246, 274 palmarum (Hentz, 1832), Hentzia, 228, 330, 331 species, Pelegrina, tropicus G. & E. Peckham, 1901, palustris (G. & E. Peckham, 1883), Sitticus, 330 Dendryphantes, 230, 238, 333 papenhoei G. & E. Peckham, 1895, Sassacus, 229, 238, 242, 333, 335 turquinensis Bryant, 1940, Sidusa, 266 unicus (Chamberlin & Gertsch, Terralonus, paykulli (Savigny & Audouin, 1825), Plexippus, 266, 1930), 267 239 uteanus Chamberlin & peckhamorum (Kaston, 1973), Pelegrina, 228, 246, Gertsch, 1929, Dendry- 272 phantes, 222, 288 uteanus Chamberlin perfectus (G. & E. Peckham, 1901), Metaphidippus, & Gertsch, 1929, Sassacus, 299 238 validus Chickering, 1946, Paraphidippus, 229, 333 perntx (G. & E. Peckham, 1901), Beata, 237 variegata (F. P.-Cambridge, 1901), Pelegrina, 241, 302 pervaga (G. & E. Peckham, 1909), Pelegrina, 222, 244, 246, 244, 245, 276 vegetus G. & E. Peckham, 1901, Dendryphantes, 308 pikei (G. & E. Peckham, 1888), Marpissa, 267 venusta Chickering, 1946, Beata, 237 prescotti Chickering, 1946, Mabellina, 231, 333 verecunda (Chamberlin & Gertsch, 1930), Pelegrina, prosper (G. & E. Peckham, 1901), Bagheera, 228, 233 241, 244, 246, 286, 299 proterva (Walckenaer, 1837), Pelegrina, 228, 236, versicolor (C. L. Koch, 1846), Phintella, 330 244, 246, 270, 331 versicolor G. & E. Peckham, 1909, Dendryphantes, proximo (G. & E. Peckham, 1901), Pelegrina, 222, 222, 311 240, 246, 265 versicolor (G. & E. Peckham, 1909), Terralonus, 239 prudensG. & E. Peckham, 1901, Dendryphantes, 265 vigens G. & E. Peckham, 1901, Gastromicans, 234 pluripunctatus (Mello-Leitao, 1944) Metaphidippus, virginis Chamberlin, 1925, Dendryphantes, 281 229 viridis (Walckenaer, 1837), Lyssomanes, 329 pura (Bryant, 1948), Messua, 234 vitis group, Metaphidippus, 228, 230, 232, 237 purpuratus Keyserhng, 1884, Phidippus, 229 vitis (Cockerell, 1894), Metaphidippus, 237, 242, 330, quadrinotatus F. P.-Cambridge, 1901, Metaphidip- 333 pus, 234 vittatus (Banks, 1901), Terralonus, 239 recurvus Chickering, 1946, Parnaenus, 229, 333 volcano new species, Pelegrina, 222, 246, 294 rudis (Sundevall, 1832), Dendryphantes, 228, 236, ivatonus (Chamberlin & Ivie, 1941), Phanias, 229, 243, 335 238 rufipes G. & E. Peckham, 1885, Zygoballus, 229, 330, wheeleri (G. & E. Peckham, 1888), Admestina, 331 331, 333 wheeleri G. & E. Peckham, 1909, Bellota, 228, 333 rus

flower-shaped flap tegulum ducts OT tegular glands gland openings

Figure 1. Adult male, Pelegrina montana (Montana: Jefferson Co.). Scale bar 1 mm.

Figure 2. Adult female, Pelegrina proterva (Pennsylvania: Adams Co.). Scale bar 1 mm.

Scale bar 0.1 mm. Figure 3. Trypsin-cleared left palpus, ventral view, Pelegrina proterva (Massachusetts: Middlesex Co.).

Figure 4. External view of epigynum, Pelegrina edrilana (Oaxaca; El Tule). Scale bar 0.1 mm.

Figure 5. Trypsin-cleared epigynum, oblique internal view showing spermathecal ducts, Pelegrina galathea (Massachusetts: Middlesex Co.). Anterior is to left. Scale bar 0.1 mm. Pelegrina Jumping Spiders • Maddison 327

Figures 6-9. Scanning electron micrographs of palpus of Pelegrina proterva. 6. Left palp, ventral view. 7. Left palp, expanded, view. 9. Left view. ventral view. 8. Bulb of righit palp, dissected from cymbium, dorsal palp, expanded, apical

Abbreviations. bH, basal hematodocha; eb, embolic base; eH, embolic hematodocha; es, embolic suture; T, tegulum; tl, tegular ledge.

Scale bar. 0.1 mm. 328 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

poison gland serrate edge retromarg openi

13 Salticine Division

Figures 10-15. Mouthparts of salticids. 10. Left chelicera of male, posterior view, Pelegrina proterva (Massacfiusetts). 11. Fang of left chelicera of male, oblique view from the posterior, Pelegrina galathea (North Carolina). 12. Right chelicera of male, medial view, Cyrba algerina (Yugoslavia). 13. Right chelicera of female, medial view, Pelegrina galathea (Massachusetts). 14. Right endite of male, dorsal view, Lyssomanes viridls (Texas). 15. Right endite of male, dorsal view, Pelegrina proterva (Mas- sachusetts).

Scale bars. 0.1 mm. Pelegrina Jumping Spiders • Maddison 329

gnathocoxal gland openings

Salticine Division

Figures 10-15. Continued. 330 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Figures 16-27. Trypsin-cleared bulbs of left palpi of dendryphantes (Figs. 20-27) and other salticlds (Figs. 16-19). 16. Sitticus palustris (British Columbia: nr. Fernie). 17. Phintella cf. versicolor (Ch\na: E. Kwantung). 18. Phlegra fasciata (Ontario: Long Point). 19. Corythalla sp, (Quintana Roo: 31 km NE of Felipe Carrillo Puerto). 20. Phanlas albeolus (California: Monterey Co.).

21 . Aniclus sp. (Nuevo Leon: Chipinque Mesa). 22. Terralonus mylothrus (Colorado: Gunnison Co.). 23. Metaphldippus mannii (California: Riverside Co.). 24. Hentzla palmarum (F\or\6a: Collier Co.). 25. Zygo£>a//us ruf/pes (Tamaulipas: 99.1°W, 23.0°N). 26. Zygoballus Incertus (Panama: El Valle). 27. Metaphldippus vltis (Alberta: Taber).

Scale bars. 0.1 mrn. Pelegrina Jumping Spiders • Maddison 331

Euophryinae Synagelinae Ballinae Dendryphantinae

28. Figures 28-39. Expanded palpi of dendryphantes (Figs. 31-39) and other salticids (Figs. 28-30). Corythalia sp. (Chiapas: 31. Eris flava Palenque). 29. Synageles nox/osus (Florida: Alachua Co.). 30. Admestina tibialis {New Hampshire: Concord). {G. chera & E. Peckham) (Nebraska: Morrill Co.). 32. Phidippus insignarius C. L. Koch (Colorado: Logan Co.). 33. Metapliidippus Bent (Nevada: Chruchill Co.). 34. Pelegrina proterva (Massachusetts: Middlesex Co.). 35. Pelegrina galathea (Colorado: Co.). Los 36. Phanlas sp. (Chiapas: San Cristobal). 37. Hentzia palmarum (Florida: Monroe Co.). 38. Zygoballus rufipes (Veracruz: Tuxtlas). 39. Species near Zygoballus incertus (Quintana Roo: Kohunlich ruins).

Abbreviations. bH, basal hematodocha; E, embolus; eb, embolic base; eH, embolic hematodocha; es, embolic suture; T, tegulum.

Scale bars. 0.1 mm. 332 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 Pelegrina Jumping Spiders • Maddison 333

X^

n/ embolus 64 X g

64. Figure Hypothetical transformations among embolus types from the euophryine type (a) to the dendryphantine types (b- At left left h). the or lower of each of a-h is the base of the embolus; the tip of the embolus is at the top of each figure. Examples with these are types Metaphidippus chera, Ens aurantia, and Bellota wheeleri (b); Hentzia and Zygoballus (c); Eris militaris (d); Paradamoetas (e); Dendryphantes tropicus and Mabellina (f); Metaphidippus mandibulatus (g); and Messua, Bagheera, and Gastromicans (h).

Figures 40-63. Left palpi of various salticids of the subfamily Dendryphantinae. 40. Eris cf. aurantia (Chiapas: San Cristobal). 41. Phidippus audax (Ontario: Burlington). 42. Bellota wheeleri {Oaxaca: SW of Valle Nacional). 43. "Pseudicius" siticulosus (Arizona: Yavapai Co.). 44. Terralonus ca/zfom/cus (California: Santa Cruz Co.). 45. Sassacus papenhoei (Neva6a: Lander Co.). 46. Tulpius hilarus (Quintana Roo: Kohunlich ruins). 47. Phanias harfordii (California: San Mateo Co.). 48. Ghelna castanea (Virginia: Falls Church). 49. Anicius dolius Chamberlin (holotype; Jalisco: Guadalajara). 50. Hentzia mitrata (Hentz) (Minnesota: Washington Co.). 51. Zygoballus rufipes (Ontario: Essex Co.). 52. Rhene cf. flavigera (China: E. Kwantung: Yim Na San). 53. Eris militaris (Ontario: Port Elgin). 54. Rudra geniculata (Panama: Canal Zone). 55. Parnaenus recurvus (paratype; Panama: " Barro Colorado Island). 56. Paraphidippus" validus (paratype; Panama: Barro Colorado Island). 57. Species near Zygoballus /ncertus (Quintana Roo: Kohunlich ruins). 58. Paradamoetas fonfana (Ontario: Hastings Co.). 59. Metaphidippus ct wf/s(Puebla: nr. Xicotepec de Juarez). 60. Dendryphantes perfectus G. & E. Peckham (holotype; : Para). 61 . "Eris" nidicolens (France: Marseille). 62. Dendryphantes tropicusG. & E. Peckham (holotype; Brazil: Chapoda). 63. Mabe///naprescoft/Chickering (paratype; Panama: El Valle).

Scale bars. 0.1 mm. 334 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 Pelegrina Jumping Spiders • Maddison 335

71-79. Tibial of left of Figures apophyses palpi dendryphantines. 71. Bagheera kiplingi (Oaxaca: nr. Tuxtepec). 72. Messua desidiosa Rica: San 73. Gastromicans (Costa Jose). levispina (Panama: El Valle). 74. Metaphidlppus mandibulatus (holotype; Costa 75. audax Halton Rica). Phidippus (Ontario: Co.). 76. (Poland: Smogorzew). 77. Beata hispida Roo: Kohunlich 78. (Quintana ruins). Pelegrina galathea (Texas: Bexar Co.). 79. Eris militaris (North Carolina).

Scale bars. 0.1 mm.

Figures 65-70. Scanning electron micrographs of epigyna of dendryphantines, showing teardrop-shaped flaps over openings. View is mostly ventral, slightly oblique lateral. 65. Dendryphantes rudis (U.S.S.R.: Buzjatia). 66. ErIs militaris (Michigan: Emmet Co.). 67. Phidippus audax (Minnesota: Rochester). 68. Terraionus mylothrus (Colorado: Pitkin Co.). 69. Sassacus papenhoei (California: Santa Barbara Co.). 70. Phanias albeolus (Oregon: Lane Co.).

Scale bars. 0.1 mm. 336 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 Pelegrina Jumping Spiders • Maddison 337

"^ 105 \ 103

109 110 v^S

103-108. left Figures Dendryphantes species, palpi (Figs. 103, 105, 107) and emboli, oblique ventral-retrolateral view (Figs. 104, 106, 108). 103, 104. Dendryphantes hastatus (Po\an6: Smogorzew). 105, 106. Dendryphantes rudis {\Qb, France; 106, Barcelona: 107, 108. Spain: Baga). Dendryphantes nigromaculatus {: 07 , Colorado: Chaffee Co.; 108, Colorado: Gunnison Co.).

Figures 109-1 12. Beata. 109. Beata magna (one of the types; Panama: Bugaba): ventral view/ of epigynum. 110-112. Beata hispida (Quintana Roo: Kohunlich ruins). 110. Left palpus; epigynum. 111. Dorsal view. 112. Ventral view.

Scale bars. 0.1 mm.

Figures 80-85. Bagheera. 80-83. Bagheera kiplingi {Oaxaca: nr. Tuxtepec): 80. Male face. 81. Left palpus; epigynum. 82. Ventral views. 83. Dorsal view. 84, 85. Bagheera prosper {Oaxaca: Valle Nacional). 84. Left palpus. 85. Epigynum, ventral view.

Figures 86-92. Messua. 86-89. Messua desidiosa (Costa Rica: San Jose). 86. Male face. 87. Left palpus; epigynum. 88. Ventral view. 89. Dorsal view. 90. Messua llmbata (Quintana Roo: nr. Tulum ruins): left palpus. 91. Messua cf. octonotata (Chiapas: Palenque): left palpus. 92. Messua sp. cf. Metaphidippus mandibulatus {CosXa Rica: Puntarenas Province): left palpus.

Figures 93-95. Gastromlcans levispina (Panama: El Valle). 93. Left palpus; epigynum. 94. Ventral view. 95. Dorsal view.

Figures 96-98. Metaphidippus mandibulatus (holotype; Costa Rica). 96. Male face. 97. Oblique ventral-retrolateral view of left embolus. 98. Left palpus.

Figures 99-1 02. Trypsin-cleared palpi of Bagheera and similar dendryphantines. 99. Bagheera prosper{Oaxaca: Valle Nacional). 100. Messua llmbata (Arizona: Santa Cruz Co.). 101. Gastromlcans levispina, right palp, image photographically reversed (Panama: El Valle). 102. Ashtabula dentata {Panama: El Valle).

Scale bars. 0.1 mm, except for male faces 0.5 mm. 338 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

127

Figures 1 1 3-1 28. Courtship poses of various dendryphantine males, traced from photographs. 1 1 3. Phidippus audax (Ontario: Halton Co.). 114. Paradamoetas fontana (Levi) (Ontario: Richmond). 115. Zygoballus rufipes (Arizona: Santa Cruz Co.). 116. Tulpius hilarus (Tamaulipas: 99.1°W, 23.0°N). 1 1 7. Messua limbata (Arizona: Santa Cruz Co.). 1 1 8. Phanlas watonus (California: Los Angeles Co.). 119. "Pseudicius" s/f/cu/osus (Arizona: Yavapai Co.). 120. Dendryphantes nigromaculatus [Montana: Jefferson

Co.). 121 . Peleghna furcata (Arizona: Santa Rita Mtns.). 122. Hentzia mitrata (Florida: Dade Co.). 123. Tutelina similis (Banks) (Alberta: Cypress Hills). 124. £r;sm///far;s (Sasl

Pelegrina

Figure 129. Cladogram for Pelegrina species. 340 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

I

131

133

f ^^^^

134 135

Figures 130-135. Photographs of living Pelegrina. 130, 131. Pelegrina galathea (sp. 1). 130. Male (Ontario: Mississauga). 131. Female (Massachusetts: Middlesex Co.). 1 32, 1 33. Pelegrina dithalea (sp. 3). 1 32. Male (holotype; Arizona: Sycamore Canyon). 133. Female (Arizona: Kitt Peak). 134, 135. Pelegrina proterva (sp. 5). 134. Male (Manitoba: Binscarth). 135. Female (Ontaho: Sudbury District). Pelecrina Jumping Spiders • Maddison 341

Figures 136-141. Photographs of living Pelegrina. 136, 137. Peleghna peckhamorum(sp. 6); Massachusetts. 136. Male. 137. Female. 138. Peleghna neoleonis (sp. 7): male (Nuevo Leon: Cerro Potosi). 139. Pelegrina chalceola (sp. 21); male (holotype: Arizona: Madera Canyon). 140. 141. Pelegrina kastoni {sp. 11). 140. Male (holotype; Arizona: Mount Hopkins). 141. Female (Arizona: Madera Canyon). 342 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Figures 142-147. Photographs of living Peleghna. 142, 143. Pelegrina flavipedes {sp. 12); Manitoba: nr. Neepawa). 142. Male. 143. Female. 144, 145. Peleghna flaviceps (sp. 13). 144. Male (Maine; Sagadahoc Co.). 145. Female (New Hampshire: Durham). 146, 147. Pelegrina exigua (sp. 14) (dull form). 146. Male (Maryland: Montgomery Co.). 147. Female (Virginia: Shenandoah Co.). Pelegrina Jumping Spiders • Maddison 343

150 151

Figures 148-153. Photographs of living Pelegrina. 148, 149. Pelegrina exigua (sp. 14) (striped form). 148. Male (Maryland: Montgomery Co.). 149. Female (Virginia: Washington Co.). 150, 151. Pelegrina insignis {sp. 16; Minnesota: Hennepin). 150. Male. 151. Female. 152, 153. Pelegrina clemata (sp. 18; Saskatchewan: Outlook). 152. Male. 153. Female. 344 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

of Figures 154-159. Photographs living Pelegrina. 154. Pelegrina montana (sp. 15); male (Montana: Jefferson Co.). 155. Pelegrina helenae (sp. 29): female (Washington: Franklin Co.). 156, 157. Pelegrina aeneola (sp. 19; California: Riverside Co.). 1 56. Male. 1 57. Female. 1 58, 1 59. Pelegrina furcata (sp. 22). 1 58. Male (Arizona: Mount Hopkins), 1 59. Female (Arizona: Yavapai Co.). Pelegrina Jumping Spiders • Maddison 345

161

'"'•iN

163

165

Figures 1 60-1 65. Photographs of living Pelegrina. 1 60, 1 61 . Pelegrina arizonensis (sp. 28; Minnesota: Anoka Co.). 1 60. Male. 161. Female. 162, 163. Pelegrina verecunda (sp. 30; Arizona: Yavapai Co.). 162. Male. 163. Female. 164, 165. Pelegrina clavator (sp. 31). 164. Male (Nuevo Leon: Chipinque Mesa). 165. Female (Veracruz: Naolinco). 346 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Figures 166-171. Photographs of living Peleghna. 166, 167. Peleghna variegata (sp. 33; Oaxaca: nr. El Tule). 166. Male. 167. Female. 168. Peleghna sandaracina (sp. 35): male (holotype; Campeche: nr. Francisco Escarcega). 169. Peleghna yucatecana (sp. 34): female (Campeche: Xpujil). 170, 171. Peleghna bunites (sp. 37; Arizona: Mount Hopkins). 170. Male (holotype). 171. Female. Pelegrina Jumping Spiders • Maddison 347

!

174 175

^ /f-\

176

mannii 173. Figures 172-177. Photographs of living Pelegrina, Nagaina, and Metaphldippus group species. 172, Pelegrina Orestes (sp. 38; Arizona: Madera Canyon). 172. Male. 173. Female. 174, 175. Nagaina Incunda (sp. 39). 174. Male (San Luis Potosi: nr. Las Abritas). 175. Female (Tamaulipas: 99°04W, 23°00'N). 176, 177. Metaphldippus emmiltus (sp. 45; New Mexico: Guadalupe Co.). 176. Male (holotype). 177. Female (paratype). 348 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

i

178 179

Figures 1 78-1 83. Photographs of living Metaphidippus mannii group species. 1 78, 1 79. Metaphldippus mannii {form versicolor) (sp. 40). 178. Male (California: Riverside Co.). 179. Female (Washington: Seattle). 180, 181. Metaphidippus mannii (form mannif)

(sp. 40). 1 80. Male (Arizona: Sycamore Canyon). 1 81 . Female (Arizona: Santa Catalina Mtns.). 1 82, 1 83. Metaphidippus diplacis (sp. 45). 182. Male (California: San Diego Co.). 183. Female (California; Santa Barbara Co.). Pelegrina Jumping Spiders • Maddison 349

Figures 184-189. Photographs of living Metaphidippus mannii group species. 184, 185. Metaphidippus tricolor {sp. 42; Cali- fornia: Monterey Co.). 184. Male. 185. Female. 186, 187. Metaphidippus chera (sp. 43). 186. Male (California; San Luis Obispo Co.). 1 87. Female (California; Santa Barbara Co.). 1 88, 1 89. Metaphidippus carmenensis (sp. 44; California; Riverside Co.). 1 88. Male. 189. Female. 350 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

220 Pelegrina Jumping Spiders • Maddison 351

226 228 227

229

231 232 233 235

Figures 226-235. Embolus of left palpus, oblique ventral-retrolateral view, of Pelegrina, Nagalna, and Metaphidippus mannii = group species. All figures to same scale; scale bar 0.1 mm. 226. Pelegrina bunites (sp. 37; Arizona: Quinlan Mtns.). 227. Pelegrina orestes (sp. 38; Arizona: Madera Canyon). 228. Nagaina incunda (sp. 39; Quintana Roo: Kohunlich ruins). 229. Metaphidippus mannii (form versicolor) (sp. 40; California: Riverside Co.). 230. Metaphidippus mannii (form mannii) (sp. 40; Arizona: Santa Cruz Co.). 231. Metaphidippus diplacis (sp. 41; California: San Diego). 232. Metaphidippus tricolor (sp. 42; California: Monterey Co.). 233. Metaphidippus chera (sp. 43; Arizona: Pima Co.). 234. Metaphidippus carmenensis {sp. 44; Baja California Norte: Isia Angel de al Guardia). 235. metaphidippus emmiltus (sp. 45; holotype).

Figures 190-225. Embolus of left palpus, oblique ventral-retrolateral view, of Pelegrina species. The opening to the sperm duct, the prolateral ramus, and retrolateral ramus are labeled with "o," "p," and "r," respectively. All figures to same scale; scale bar 0.1 mm. 190. P. galathea (sp. 1; Chihuahua: 105.2°W, 27.9°N). 191. P. proxima (sp. 2 Cuba: Holquin). 192. P. dithalea (sp. 3; Arizona: Santa Cruz Co.). 193. P. edrilana (sp. 4; Distrito Federal: TIalpam). 194. P. protervaw (sp. 5; Ontario: Kenora District). 195. P. peckhamorum (sp. 6; Massachusetts: Barnstable Co.). 196. P. neoleonis (sp. 7; Nuevo Leon: Chipinque Mesa). 197. P. tristis (sp. 8; Arizona: Chiricahua Mtns.). 198. P. sabinema (sp. 9; Arizona: Coconino Co.). 199. P. pervaga (sp. 10; Texas: Erath Co.). 200. P. kastoni (sp. 11; Arizona: Chiricahua Mtns.). 201. P. flavipedes (sp. 12; Manitoba: 19 km E of Barnstable P. Neepawa). 202. P. flaviceps (sp. 13; Maine: Sagadahoc Co.). 203. P. exigua (sp. 14; Massachusetts: Co.). 204. 206. P. montana (sp. 15; New Hampshire: Jeffrey). 205. P. insignis (sp. 16; Saskatchewan: North Battleford). chaimona (sp. 17; Arizona: Chiricahua Mtns.). 207. P. clemata (sp. 18; Alberta: Morrin Recreational Area). 208. P. aeneola (aeneola) (sp. 19; Oregon: Lane Co.). 209. P. aeneola (uteanus) (sp. 19; South Dakota: Custer Co.). 210. P. balia (sp. 20; Oregon: Deschutes 211, P. 21 Chiricahua 212. P. furcata 22; Arizona: Santa Rita 213. P. volcana Co.). chalceola (sp. ; Ahzona: Mtns.). (sp. Mtns.). (sp. 23; Panama: El Volcan). 214. P. bicuspidata (sp. 24; right palp, image photographically reversed) (holotype; Guatemala). P. Ahzona: 215. P. morelos (sp. 26; holotype; Morelos: nr. Cernavaca). 216. huachuca (sp. 27; holotype; Huachuca Mtns.). P. nr. P. 217. P. arizonensis (sp. 28; New Mexico: Bernalillo Co.). 218. helenae(sp. 29; Oregon: Phneville). 219. verecunda (sp. 221. P. 30; Arizona: Yavapai Co.). 220. P. clavator (sp. 31; Nuevo Leon: Chipinque Mesa). pallidata (sp. 32; Nicaragua: Matagalpa). 222. P. variegata (sp. 33; Oaxaca: El Tule). 223. P. yucatecana (sp. 34; holotype; Yucatan: Chichen Itza). 224. P. P. tillandsiae Carolina: sandaracina (sp. 35; Holotype; Campeche: Francisco Escarcega). 225. (sp. 36; South Cooper). 352 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Figures 236-241 . Scanning electron micrographs of female epigyna of Pelegrina species. View is mostly ventral, slightly oblique lateral. Scale bars 0.1 mm. 236. P. galathea (sp. 1; Michigan: Washtenaw Co.). 237. P. proxima (sp. 2; Cuba: Havana). 238. P. proterva (sp. 5; Iowa: Hancock Co.). 239. P. peckhamorum (sp. 6; Arkansas: Washington Co.). 240. P. pervaga (sp. 10; Texas: Erath Co.). 241. P. flavipedes (sp. 12; Alberta: Edmonton). Pelecrina Jumping Spiders • Maddison 353

Figures 242-247. Scanning electron micrographs of female epigyna of Pelegrina species. View is mostly ventral, slightly oblique lateral. Scale bars 0.1 mm. 242. P. flaviceps(sp. 13; Maine: Sagadahoc Co.). 243. P. exlgua(sp. 14; Virginia; Shenandoah Co.). 244. P. montana (sp. 15; Colorado; Boulder Co.). 245. P. insignis (sp. 16; New Hampshire; Cheshire Co.). 246. P. clemata (sp. 18; Colorado; Gunnison Co.). 247. P. aeneola (sp. 19; Oregon; Lane Co.). 354 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 '

Pelegrina Jumping Spiders • Maddison 355

Figures 254-257. Scanning electron micrographs of female epigyna of Pelegrina and Metaphidippus mannii group species. View is mostly ventral, slightly oblique lateral. Scale bars 0.1 mm. 254. P. tillandsiae {sp. 36; South Carolina: Cooper). 255. P. bunltes (sp. 37; Arizona: Mount Hopkins). 256. Metaphidippus mannii (sp. 40; Oregon: Lane Co.). 257. Metaphidippus chera (sp. 43; Arizona: Pima Co.).

Figures 248-253. Scanning electron micrographs of female epigyna of Pelegrina species. View is mostly ventral, slightly oblique lateral. Scale bars 0.1 mm. 248. P. balla (sp. 20; California: Plumas Co.). 249. P. furcata(iorm mimus; sp. 22; Arizona: Yavapai P. P. Co.). 250. P. furcata (sp. 22; Arizona: Yavapai Co.). 251. arizonensis (sp. 28; Minnesota: Anoka Co.). 252. verecunda (sp. 30; Arizona: Yavapai Co.). 253. P. variegata (sp. 33; Oaxaca: El Tule). 356 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

280. edrilana

Figures 258-263. Pelegrina galathea (sp. 1). 258. 6 face (Texas: San Patricio Co.). 259. Tip of embolus (Florida; Sebastian).

260. Bruce 261 , 262. dorsal ventral 9 Palp (Ontario: Co.). Epigynum, (261 ) and (262) (Arkansas: Washington Co.). 263. abdomen (Pennsylvania, Erie Co.).

264-269. Figures Pelegrina proxima (sp. 2). 264.

Figures 270-275. Pelegrina dithalea (sp. 3; Arizona: Santa Cruz Co., Sycamore Canyon). 270. 6 face (holotype). 271. Tip of embolus. 272. Palp. 273, 274. Epigynum, dorsal (273) and ventral (274). 275. 9 abdomen.

Figures 276-281. Pelegrina edrilana (sp. 4). 276-278. Holotype. 276. S face. 277. Tip of embolus. 278. Palp. 279. Epigynum, dorsal nr. El (Oaxaca: Tule). 280. Epigynum, ventral (Distrito Federal: San Jeronimo; see also Fig. 4). 281. 9 abdomen (Distrito Federal: San Jeronimo),

Scale bars. 0.1 mm, except for 6 face and i abdomen 0.5 mm and tip of embolus 0.01 mm. Pelegrina Jumping Spiders • Maddison 357

Figures 282-287. Pelegrina proterva (sp. 5). 282. 6 face (Massachusetts: Dukes Co.). 283. Tip of embolus (Massachusetts: Middlesex Co.). 284. Palp (Ontario: near Barrie). 285. Epigynum, dorsal (Ontario: Muskoka District). 286. Epigynum, ventral (Iowa: Hancock Co.). 287. 5 abdomen (Michigan: Mackinac Co.).

Figures 288-293. Pelegrina peckhamorum (sp. 6; Arkansas, except 289 and 293 Massachusetts). 288. 6 face. 289. Tip of embolus. 290. Palp. 291. Epigynum, dorsal. 292. Epigynum, ventral. 293. 2 abdomen.

Figures 294-298. Pelegrina neoleonis (sp. 7). 294. c5 face (Chipinque Mesa). 295. Palp (Cerro PotosO- 296, 297. Epigynum, dorsal (296) and ventral (297) (Cerro Potosi). 298. 9 abdomen (Oaxaca).

Figures 299-303. Pelegrina tristis (sp. 8). 299. i face (holotype). 300. Palp (holotype). 301, 302. Epigynum, dorsal (301) and ventral (302) (paratype; arrow/ shows pale surface that descends deeply). 303. 2 abdomen (Arizona: Madera Canyon).

Scale bars. 0.1 mm, except for 6 face and 2 abdomen 0.5 mm and tip of embolus 0.01 mm. 358 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

flavipedes

Figures 304-308. Pelegrina sabinema (sp. 9). 304. 6 face (New Mexico: Santa Fe Co.). 305. Palp (Arizona; Coconino Co.). 306, 307. Epigynum, dorsal (306) and ventral (307) (New Mexico: Santa Fe Co.). 308. 9 abdomen (New Mexico: nr. Edgewood).

Figures 309-31 3. Pelegrina pervaga (sp. 1 0). 309. 6 face (Texas: Kerr Co.). 31 0. Palp (Texas: Kerr Co.). 311,312. Epigynum, dorsal (311) and ventral (312) (Texas: Val Verde Co.). 313. 9 abdomen (Texas: Val Verde Co.).

Figures 314-318. Pelegrina kastoni(sp. 11). 314. 3 face (Arizona: Sycamore Canyon). 315. Palp (Arizona: Cfiiricahua Mtns.). 316, 317. Epigynum, dorsal (316) and ventral (317) (Arizona: nr. Cienega Lake). 318. 9 abdomen (Arizona: Madera Canyon).

Figures 319-323. Pelegrina flavipedes (sp. 12). 319. <5 face (Alberta: Cypress Hills). 320. Palp (Ontario: Muskoka District). 321, 322. Epigynum, dorsal (321) and ventral (322) (Ontario: Bruce Co.). 323. 9 abdomen (Michigan: Crawford Co.). See also Figures 338 and 339.

Scale bars. 0.1 mm, except for 6 face and i abdomen 0.5 mm. Pelegrina Jumping Spiders • Maddison 359

338

Figures 324-328. Pelegrina flaviceps (sp. 13; Ontario: Kingston, except 324 New Hampshire: Surry). 324. 6 face. 325. Palp. 326. Epigynum, dorsal. 327. Epigynum, ventral. 328. 2 abdomen. See also Figures 340 and 341.

Figures 329-335. Pelegrina exigua (sp. 14). 329-333. Dull form (Virginia: Shenandoah Co., except 333 Kentucky: Rowan Co.). 329. $ face. 330. Palp. 331. Epigynum, dorsal. 332. Epigynum, ventral. 333. 5 abdomen. 334, 335. Striped form. 334. 5 face (Maryland: Montgomery Co.). 400. s abdomen (Missouri: Jefferson City). See also Figures 336, 337, and 342.

Figures 336-342. Spermathecal ducts of cleared epigyna, dorsal view, of flavipedes group species. 336, 337, 342. P. exigua (336, North Carolina: Durham Co.; 337, Kentucky: Rowan Co.; 342, holotype. New York: Ithaca). 338, 339. P. flavipedes {336. Ontario: Sudbury District; 339, Manitoba: Neepawa). 340, 341. P. flawceps 340, Maine: Sagadahoc Co.; 341, New Hampshire: Strafford Co.

Scale bars. 0.1 mm, except for 6 face and 9 abdomen 0.5 mm. 360 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

Figures 343-347. Pelegrina montana (sp. 15). 343. 6 face (Vermont: Chittendon Co.). 344. Palp (Alberta: Waterton Lake). 345, 346. Epigynum, dorsal (345) and ventral (346) (Northwest Territories: Sawmill Bay). 347. 9 abdomen (British Columbia: Pink Mtn.).

Figures 348-353. Pelegrina Insignis (sp. 1 6). 348. 6 face (New Brunswick: nr. Chipman). 349. Embolus (Michigan: Midland Co.). 350. Palp (Ontario: Barrie). 351, 352. Epigynum, dorsal (351) and vental (352) (Barrie). 353. 2 abdomen (Minnesota: Olmsted Co.).

Figures 354-358. Pelegrina chaimona (sp. 17). 354. s face (holotype). 355. Palp (holotype). 356, 357. Epigynum, dorsal (356) and ventral (357) (Anzona: Cochise Co.). 358. 9 abdomen (Arizona: Cochise Co.).

Figures 359-364. Pelegrina clemata(sp. 1 8; Colorado: Saguache Co., except 359 Colorado: Gunnison Co. and 360 Washington: Yakima Co.). 359. 6 face. 360. Embolus. 361. Palp. 362. Epigynum, dorsal. 363. Epigynum, ventral. 364. 9 abdomen.

Scale bars. 0.1 mm, except for s face and 9 abdomen 0.5 mm. Pelegrina Jumping Spiders • Maddison 361

Figures 365-377. Pelegrina aeneola (sp. 19). 365. <5 face (California: Ventura Co.). 366-372. Palpi (366, British Columbia: Fountain Valley; 367, California: Ventura co.; 368, 369, Oregon: Lake Co.; 370, 371 Idaho: Franklin Co.; 372 Wyoming: Sheridan Co.). 373, 374. Epigynum dorsal (373) and ventral (374) (British Columbia: Fountain Valley). 375, 376. Epigynum, dorsal (375) and ventral (376) (South Dakota: Custer Co.). 377. 5 abdomen (South Dakota: Custer Co.).

Figures 378-382. Pelegrina balia (sp. 20; California: Santa Barbara Co., holotype and paratype). 378. s face (arrow shows distinctive flange on fang). 379. Palp. 380. Epigynum, dorsal. 381 . Epigynum, ventral. 382. 9 abdomen.

Figures 383-387. Pelegrina chalceola (sp. 21). 383. <5 face (holotype). 384. Palp (holotype). 385, 386. Epigynum, dorsal (385) and ventral (386) (Arizona: Chiricahua Mtns.). 387. $ abdomen (Arizona: Chiricahua Mtns.).

Scale bars. 0.1 mm, excpet for 6 face and 9 abdomen 0.5 mm. 362 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4

405 bicuspidata

Figures 388-402. Pelegrina furcata (sp. 22). 388. <5 face (Oaxaca: 31 km N of Guelatao de Jaurez). 389. Tibial apophysis nr. de El Arizona: Santa (Puebia: Xicotepec Juarez). 390-394. Palpi (390, Colorado: Paso Co.; 391 , Arizona: Yavapai Co.; 392, Cruz Co.; 393, Guatemala; labeled "Type"; 394, Oaxaca: 31 km N of Guelatao de Juarez). 395, 396 (Arizona: Yavapai Co.). 395. Epigynum, ventral. 396. 9 abdomen. 397-399 (Arizona: Santa Cruz Co.). 397. Epigynum, dorsal. 398. Epigynum, ventral. 399. 9 abdomen. 400-402 (Oaxaca: 31 km N of Guelatao de Juarez). 400. Epigynum, dorsal. 401. Epigynum, ventral. 402. 2 abdomen.

Figures 403, 404. Pelegrina volcana (sp. 23; Panama: El Volcan). 403. 6 face. 404. Palp.

Figures 405, 406. Pelegrina bicuspidata (sp. 24). 405. s face (Chiapas: nr. Arriaga). 406. Right palp, image photographically reversed (holotype).

Figures 407-409. Pelegrina ochracea (sp. 25). 407, 408. Epigynum, ventral view after clearing (407) and before clearing (408) (holotype). 409. 9 abdomen (Chiapas: San Cristobal).

Scale bars. 0.1 mm, except for 5 face and 9 abdomen 0.5 mm. Pelegrina Jumping Spideks • Maddison 363

430 helenae

Figures 410-414. Pelegrina morelos (sp. 26; holotype and paratype). 410. 6 face. 411. Palp. 412. Epigynum, dorsal. 413. Epigynum, ventral. 414. 9 abdomen.

Figures 415-419. Pelegrina huachuca (sp. 27). 415. $ face (holotype). 416. Palp (holotype). 417, 418. Epigynum, dorsal (417) and ventral (418) (Arizona: Madera Canyon). 419. 9 abdomen (Arizona: Santa Catalina Mtns.).

Figures 420-425. Pelegrina arizonensis (sp. 28; Minnesota: Anoka Co.). 420. S face. 421. Tibial apophysis. 422. Palp. 423. Epigynum, dorsal. 424. Epigynum, ventral. 425. 9 abdomen.

Figures 426-431. Pelegrina helenae (sp. 29). 426. $ face (Washington: Franklin Co.). 427-430 (Wyoming: Bighorn Co.). 427. Tibial apophysis. 428. Palp. 429. Epigynum, dorsal. 430. Epigynum, ventral. 431. 9 abdomen (Nevada: Washoe Co.).

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Figures 432-436. Pelegrina verecunda (sp. 30; Arizona: Yavapai Co., except 436 Arizona: Cochise Co.). 432. s face. 433. Palp. 434. Epigynum, dorsal. 435. Epigynum, ventral. 436. ? abdomen.

Figures 437-441. Pelegrina clavator{sp. 31; Nuevo Leon: Chlpinque Mesa). 437. s face. 438. Palp. 439. Epigynum, dorsal.

440. Epigynum, ventral. 441 . 9 abdomen.

Figures 442-446. Pelegrina pallidata (sp. 32). 442. $ face (Nicaragua: Matagalpa). 443. Palp (Nicaragua: Matagalpa). 444, 445. Epigynum, dorsal (444) and ventral (445) (Guatemala: Chichicatenango). 446. 9 abdomen (Chiapas: near San Cristobal).

Figures 447-451 . Pelegrina variegata (sp. 33; Oaxaca: nr. El Tule). 447. 6 face. 448. Palp. 449. Epigynum, dorsal. 450. Epigynum,

ventral. 451 . 9 abdomen.

Scale bars. 0.1 mm, except for s face and 9 abdomen 0.5 mm. Pelegrina Jumping Spiders • Maddison 365

455. dorsal Figures 452^56. Pelegrina yucatecana (sp. 34). 452. 6 face (holotype). 453. Palp (holotype). 454, Epigynum, (454) and ventral (455) (Yucatan: nr. Xocenpich). 456. 5 abdomen (Yucatan: or. Chichen Itza).

458. 459^61 Grutas de Figures 457-463. Pelegrina sandaracina (sp. 35). 457. 6 face (holotype). Palp (holotype). (Yucatan: 462. dorsal. Loltun). 459. Epigynum, dorsal. 460. Epigynum, ventral. 461. 5 abdomen. 462^63 (Chiapas: Arriaga). Epigynum, 463. Epigynum, ventral.

Mexico. 464. dorsal. 465. ventral. 466. 5 Figures 464-471. Pelegrina species. 464-466. From Neriaco, Epigynum, Epigynum, abdomen. 467, 468. From Jalisco and Guerrero. 467. Epigynum, ventral (Guerrero: 11 mi W of Chllpancingo). 468. ? abdomen El dorsal. 470. ventral. (Jalisco: 3 mi S of Mazamitia). 469-471. From Durango (10 mi E of Salto). 469. Epigynum, Epigynum, 471. 9 abdomen.

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Nagaina incunda

Figures 472-477. Pelegrina tlllandsiae (sp. 36), 472. <5 face (North Carolina: Polluckville). 473. Embolus (Florida: Lake Placid). 474. Palp (Louisiana: Baton Rouge). 475, 476. Epigynum, dorsal (475) and ventral (476) (Louisiana: Baton Rouge). 477. 9 abdomen (Florida: Lake Placid).

Figures 478-482. Pelegrina bunites (sp. 37; Oaxaca: 50 km NW of Oaxaca, except 478 Arizona: Santa Cruz Co.). 478. 3 face. 479. Palp. 480. Epigynum, dorsal. 481. Epigynum, ventral. 482. 9 abdomen.

Figures 483-487. Pelegrina orestes (sp. 38; Arizona: Madera Canyon). 483. <5 face. 484. Palp. 485. Epigynum, dorsal. 486. Epigynum, ventral. 487. 9 abdomen.

Figures 488-492. Nagaina incunda (sp. 39). 488. 6 face (Chiapas: 76 km S of Palenque). 489. Palp (Quintana Roo: Kohunlich ruins). 490, 491. Epigynum, dorsal (490) and ventral (491) (Quintana Roo: Kohunlich ruins). 492. 9 abdomen (Chiapas: 105 km SE of Palenque).

Scale bars. 0.1 mm, except for 6 face and 9 abdomen 0.5 mm. Pelegrina Jumping Spiders • Maddison 367

Figures 493-502. Metaphidippus mannii {sp. 40). 493-497. Form vers/co/or (California: Mendocino Co., except 495 and 496 Humboldt Co.). 493. s face (arrow shows bulge on chelicerae typical of mann/V group). 494. Palp. 495. Epigynum, dorsal. 496. Epigynum, ventral. 497. 9 abdomen. 498-502. Form mannii (Arizona: Sycamore Canyon, except 498 holotype from Arizona). 498, 499. Palpi. 500. Epigynum, dorsal. 501. Epigynum, ventral. 502. 9 abdomen.

41 California: 1 mi S of San from Figures 503-508. Metaphidippus diplacis (sp. ; Baja Quintin, except 504 holotype San Diego, California). 503. 3 face. 504. Embolus. 505. Palp. 506. Epigynum, dorsal. 507. Epigynum, ventral. 508. 9 abdomen.

Figures 509-513. Metaphidippus tricolor {sp. 42; California: Marin Co., except 509 and 513 Monterey Co.). 509. 6 face. 510. Palp. 511. Epigynum, dorsal. 512. Epigynum, ventral. 513. 9 abdomen.

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Figures 514-528. Metaphidippus chera (sp. 43). 514. 5 face (New Mexico: Dona Ana Co.). 515. Tibial apophysis (New Mexico: Doiia Ana Co.). 516, 517. Palpi (Nevada: Churchill Co.; arrow in 516 shows bulge at base of tibial apophysis, typical of mannii group). 518-523. Emboli of <5^ from one locality showing variation in shape of embolus and embolic base (Texas: Wichita Co.). 524, 525. Epigynum, dorsal (524) and ventral (525) (Arizona: Pima Co.). 526. Epigynum, ventral (California: Riverside Co.). 527, 528. V abdomen (Arizona: Pima Co.).

Figures 529-533. Metaphidippus carmenensis (sp. 44; Californa: Riverside Co., except 533, Baja California: San Felipe). 529. 6 face. 530. Palp. 531. Epigynum, dorsal. 532. Epigynum, ventral. 533. 9 abdomen.

Figures 534-538. Metaphidippus emmiltus (sp. 45; holotype and paratype). 534. s face. 535. Palp. 536. Epigynum, dorsal. 537. Epigynum, ventral. 538. 9 abdomen.

Scale bars. 0.1 mm, except for $ face and 9 abdomen 0.5 mm.