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Opiliones, Laniatores) 2002. The Journal of Arachnology 30:435-451 THE HARVESTMAN FAMILY PHALANGODIDAE 4. A REVIEW OF THE GENUS BANKSULA (OPILIONES, LANIATORES) Darrell Ubick and Thomas S. Briggs: Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA. E-mail:[email protected] ABSTRACT. Two new species of Banksula are described. Banksula incredula enlarges the concept of the genus and is assigned to a new species group which is the likely sister group to the other Banksula. The species is unique in numerous morphological features, being the largest species in the genus, the first non-cavernicolous species, and occurring in the Coast Ranges of California, rather than in the Sierra Nevada foothills. The second new species, B. ttltankhamen, is a typical member of the californica group but with more pronounced troglomorphy. Clinal variation is documented for B. grahami Briggs 1974, with B. elliotti Briggs & Ubick 1981, now placed as its junior synonym, representing the northern and most troglomorphic populations. Keywords: Banksula, Phalangodidae, Opiliones, cavernicoles, California The phalangodid genus Banksula Roewer atively simple glans consisting of a stylus and 1949 was established for Scotolenwn califor­ a pair of parastylar lobes (Figs. 18-22). nica Banks 1900, a troglobitic species from Alabaster Cave in the Sierra Nevada, Califor­ METHODS nia. Eight additional species were subsequent­ Specimen preparation and observation fol­ ly added by Briggs (1974) and Briggs & lows the format described in earlier papers Ubick (1981). Although some of these have (Briggs & Ubick 1981; Ubick & Briggs large apparently functional eyes, all are re­ 1989). All measurements not labeled other­ stlicted to the caves of the central Sierra Ne­ wise are in millimeters. 'Fig.' and 'Figs.' refer vada foothills. Given the narrow distribution to this paper, ' fig' and ' figs' to previously and habitat preference for the genus, it came published works. as quite a surprise to discover an epigean spe­ All specimens are deposited at the Califor­ cies, and especially one from the Coast Ran­ nia Academy of Sciences, San Francisco, Cal­ ges of California. This species, which we are ifornia, apart from the type series of Banksula describing here as Banksula incredula new californica (Banks), which is at the Museum species, also extends the generic limits on of Comparative Zoology, Cambridge, Massa­ morphological grounds, being the largest spe­ chusetts. cies in the genus and having a leg hypertrophy The following abbreviations are utilized: which, in some individuals at least,exceeds AS = apical spine of VPP; AT = apical tooth that of most troglobitic Banksula.· Despite of ovipositor; EMH = eyemound height; these and other unique features, which are dis­ EML = eyemound length; GO = genital oper­ cussed below, the species is dearly a member . culum; GOW = genital operculuni width; of the genus Banksula gi ven that its ·palpal PSL = parastylar lobes; SL = scute length; femur bears a dorsal row of setose tubercles SW = scute width; TEL = total body length; (Figs. 2, 3). Although similar tubercles occur TC = tarsal count; VP = ventral plate; VPP in some species of Sitalcina (Figs. 4, 5), these = ventral plate prong; VPW = VPP width. do not bear setae artdthus are not homologous TAXONOMY to those in Banksula.Also,and even more fundamentally, the penis of B. incredula is Genus Banksula Roewer typical of Banksula, having a bifurcate ventral Banksula Roewer 1949: 33; Briggs 1974: I; Briggs plate with ventrally situated prongs and a rel- & Ubick 1981: 315. 435 436 THE JOURNAL OF ARACHNOLOGY UBICK & BRlGGS-REVIEW OF BANKSULA 437 1 Figure l.- Banksula incredula, lateral view (drawing by Jenny Speckels). Figure 9.-Map of central California showing distribution and. area cladogram of Banksula. Sym­ Type species.-Scotolemon californica The incredula species group bols: diamond ~ B. incredula; dots = B. califomica Banks 1900. group (1, B. galilei; 2, B. cali/omica; 3, B. grubbsi; Diagnosis.-Species of Banksula are Diagnosis.-The single species represent­ ing this group differs from other Banksula in 4, B. rudolphi; 5, B. tutankhamen; 6, B. martillo­ unique among the Nearctic Phalangodidae in rum; 7, B. tuolumne); oblong region =: B. melones several characters: it is large, has a higher tar­ having a row of setiferous dorsal tubercles on group (8 , B. melones; 9, B. grahami). sal count, occupies an epigean habitat, has Figures 2-5.-Phalangodid palpi. 2, Banksula in­ the paJpal femur (Figs 2, 3) and a penis with credula, lateral view of male left palp; 3, same, a bifurcate ventral plate with ventrally posi­ large eyes and a high eyemound, and has ad­ close-up showing dorsal femoral tubercle; 4, Sital­ culum width, 0.32. Leg IT length, 10.12. Leg tioned prongs (Figs. 18, 19). ditional palpal megaspines, with 4 ectobasal cina chalona Briggs, lateral view of male left pal pal II1scute length, 4.64. Tarsal count, 5-8 and Description.-Color of body yellowish to and 2 mesoapical on the femur As regards the femur; '. 5, same; clos.e-up showing dorsal tubercle. 9-5-6. Proximal megaspine of palpal femur orange; appendages white to yellowish white. genitalia, males have a ventral plate with large Scale bar: 2 = 610 j.1m; 3 = 38 j.1m; 4 = 150 j.1m; with mesal tubercle at base; lacking ectal tu­ Body length 1.5- 3.0 mm. Body covered with apical spines and long setae and females an 5 = 30 j.1m. bercle. tubercles. Scute with eye mound low and ovipositor with smooth cuticle and a pair of Penis (Figs. 18-22): VPP subparallel, sep­ rounded to high and conical; cornea and retina apical teeth. with 4 ventrobasal and 2 mesodistal; patella arated by width of one prong; with large present or absent. Anterior margin of scute Distribution.-Known only from a single with 1 ectal and 2 mesal; tibia with 3 ectal slightly curved AS; with 10 long setae in api­ with 0 to a few pairs of small anterior tuber­ locality in San Mateo County, California. and 3 mesal; tarsus with 2 ectal and 2 mesal. cal half and 12 shOlt setae basally on each cles; ozopores lateral. Pal pal megaspines: fe­ Banksula incredula new species Palpal femur with 5-7 setiferous tubercles prong; long setae clearly longer than to twice mur with one to two mesodistal and three to Figs. 1-3, 6, 9-22 dorsally. the length of AS; short setae about half the four ectobasal; patella with two mesal and one Male (holotype): Total body length, 2.59. length of AS. Glans with apical stylar region ectal; tibia with three mesal and three ectal; Type.-Male holotype from within a sand­ Scute length, 2.18; width, 2.01. Eyemound consisting of triangular pointed stylus and a tarsus with two mesal and two ectal. Palp with stone talus on San Bruno Mountain, San Ma­ length, 0.49; width, 0.64; height, 0.49. Oper- pair of ribbon-like apically rounded PSL. dorsal row of setiferous tubercles on femur teo County, California (10 May 1991; D and smaller tubercles dorsally on other seg­ Ubick) deposited in CAS. ments, laterally on femur, and between me­ Etymology.-The species name refers to gaspines; basal megaspine of femur with me­ the unexpected morphology, habitat, and dis­ sal and ectal tubercles in most species, mesal tribution of the species, and is considered tubercle absent in most females. Tarsal count feminine in gender 4-6-5-6, in most species, and 4 or 5-7 to 9- Diagnosis.-Same as for species group. 5-5 or 6 in B. incredula. Ovipositor surface Description.-Total body length, 2.56- wrinkled; smooth or covered with micro­ 3.00. Scute length, 1.79-2.18. Leg II length, spines; apical teeth present or absent; with 6 8.81-10.51. Leg II/Scute length, 4.64-5.64. pairs of apical and 1 pair of subapical setae; Tarsal count, 4 to 5-7 to 9-5-5 to 6. (N = 8) setae slightly curved, apically pointed. Penis Color orange, appendages pale orange, tarsi with folding glans; stylus shOlt, narrow or white. Body unifOlmly coarsely tuberculate. wide, with lateral carinae present or absent, Scute margin without apparent anterior tuber­ surrounded by 1 pair of parastylar lobes; ven­ cles. Eyemound a prominent cone, as high as tral plate bifurcate, with 1 pair of apical long. Cornea and retina large and conspicu­ spines, 7-12 pairs of long setae and up to 12 ous. Genital operculum relatively small: Figures 6-8.-Venters of female Banksula representing the three species groups. 6, B. in~redula; 7, B. pairs of short setae. GOW/SW = 0.2. Palpal megaspines: femur galilei (califomica group); 8, B. melones. Scale bar: 6 = 750 11m; 7 = 430 j.1m; 8 = 600 j.1m. 438 THE JOURNAL OF ARACHNOLOGY UBICK & BRIGGS-REVIEW OF BANKSULA 439 Figures 18-20.-Banksula incredula, penis. 18, dorsal view; 19, ventral view; 20, dorsal view of left ventral plate prong showing apical spine (AS). Scale bar: 18, 19 = 150 J.Lm; 20 = 30 J.Lm. Figures 1O-13.- Banksula incredula. 10, anteri­ Figures 14-17.- Banksula incredula, ovipositor. or half of f~male scute showing eyemound; 11, fe­ 14, lateral view; 15, close-up showing apical tooth male left palpus showing two mesoapical mega­ (AT); 16, apical view; 17, close-up oflateral surface Ubick); 1 2(18 Jan. 1992, T. Briggs, W. spines. They differ from the melones group in spines (1 , 2); 12, female left palpus showing four showing absence of misrospines. Scale bar: 14 = Rauscher, D . Ubick); 1 0", 1 2 (26 Jan. 1992, having a smaller GO, witb a GO/SW ratio of ectobasal megaspines (I, 2, 3, 4); 13 , male left pal­ 200 J.Lm; 15 = 60 J.Lm; 16 = 150 J.Lm; 17 = 20 J.Lm.
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