GENERIC PLACEMENT OF THE EMPIRE CAVE PSEUDOSCORPION, MICROCREAGRIS IMPERIALIS (NEOBISIIDAE), A POTENTIALLY ENDANGERED ARACHNI D

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1995. The Journal of Arachnology 23 :171 17 6 GENERIC PLACEMENT OF THE EMPIRE CAVE PSEUDOSCORPION, MICROCREAGRIS IMPERIALIS (NEOBISIIDAE), A POTENTIALLY ENDANGERED ARACHNI D William B. Muchmore : Department of Biology, University of Rochester, Rochester, New York 14627 USA James C. Cokendolpher : Adjunct Professor, Department of Biology, Midwester n State University, Wichita Falls, Texas 76308 US A ABSTRACT. Types, topotypes, and some other material of the pseudoscorpion Microcreagris imperialis Muchmore have been studied, and the species is transferred to the genus Fissilicreagris Curcic. Supplemental descriptio n and illustrations are presented, including the first information about females. This species is known only from three caves in Cave Gulch, Santa Cruz County, California. The cave habitat for F. imperialis is threatened by vandalism, development, and closure; and the U. S. Fish and Wildlife Service has proposed this pseudoscorpio n as a candidate for listing as an endangered or threatened species. More than 1500 species and subspecies of animals in the United States are proposed by th e U. S. Fish and Wildlife Service for listing as endangered or threatened (Drewry 1994). In most instances, these taxa have been submitted for consideration in the absence of any validation o f taxonomic status. One of the candidate species of pseudoscorpions being reviewed for possibl e addition to the List of Endangered and Threatened Wildlife under the Endangered Species Act of 1973, as emended, is Microcreagris imperialis Muchmore, from Empire Cave, Santa Cru z County, California. As Mahnert (1979) and Curcic (1983) hav e demonstrated, the genus Microcreagris Balzan is restricted to two species in China and Afghanistan, and the numerous American species which had been placed in that genus are improperly assigned. Curcic (1978 1989) created quite a few new genera and placed many of the America n species in them; he was, however, unable to make definite generic assignments for some species, including M. imperialis (1984:165). The purpose of this paper is to clarify the taxonomic position of M. imperialis so that thi s potentially endangered species can be identified properly in the literature. Furthermore, we wish to provide sufficient identification characteristics so that field biologists might more easily recognize this species in its native habitat. METHOD S Specimens have been borrowed from th e American Museum of Natural History, Ne w York, New York (AMNH); California Academy of Sciences, San Francisco, California (CAS); D. Ubick personal collection, San Francisco, California (CDU) ; and the Florida State Collection of Arthropods, Gainesville, Florida (FSCA). Unless otherwise stated, the specimens have been dissected, cleared, and mounted in Canad a balsam on microscope slides. 171 Fissilicreagris imperialis (Muchmore), new combinatio n Empire Cave Pseudoscorpion Figs. 1 9 Microcreagris imperialis Muchmore 1969 :13-15, 21, fig. 10; Arnett 1984:21666 ; Briggs & Ubick 1988 :44; Drewry 1989:566 ; Coddington, Larcher & Coken - dolpher 1990 :11 ; Drewry 1991 :58833 ; Harvey 1991 : 342 ; Drewry 1994 :59025. Microcreagris' imperialis : Curcic 1984 :164, 165, figs. 20, 41. Pseudoscorpion : Briggs 1990:180. Type locality. Empire Cave, in Cave Gulch, one mile NW of Santa Cruz, Santa Cruz County, California. Material examined. Holotype male and one paratype male from Empire Cave, 26 Augus t

172 THE JOURNAL OF ARACHNOLOGY 1 1--\I T-T- 3 4 Figures 1-4.-Fissilicreagris imperialis (Muchmore), male holotype. 1, Tip of movable finger of chelicera, with galea; 2, Cheliceral flagellum; 3, Central part of sternite 3; 4, Sternites 5-9, showing chaetotaxy (setae omitted). 1963, R. E. Graham, mounted on slides (AMNH); one paratype male, same data (FSCA); one topotype female from Empire Cave, September 1972, R. Lem, mounted on slide (CAS); one topotype male from Empire Cave, 8 July 1989, D. Ubick, et al., in alcohol (CDU); two topotype males fro m Empire Cave, 8 September 1991, D. Ubick and S. Fend, mounted on slides (CAS); one female from Dolloff Cave, across Cave Gulch from Em - pire Cave, 22 April 1979, D. C. Rudolph et al., mounted on slide (CAS); three females from IXL Cave, in Cave Gulch one-half mile S of Dollof Cave, 21 April 1979, D. C. Rudolph et al., tw o mounted, one in alcohol (CAS). Supplementary description. The topotypes and specimens from Dolloff and IXL caves are generally similar to the holotype and paratypes. Dimensions, proportions, and chaetotaxies o f body and appendages vary slightly from the values given in the original description, but all appear conspecific with the types. A few feature s of the additional specimens are, however, worth mentioning. Apex of palpal coxa (manducatory process ) bears three setae in all specimens, as in types. Cheliceral galea short and twice bifid (Fig. 1); galeae of types also like this, not just "with four or five terminal spinules" as characterized i n original description (Muchmore 1969:15). Cheliceral flagellum composed of 7-8 serrate setae (Fig. 2). Sternites 6, 7, and 8 with two setae on face near middle (discal setae) as in types (one topotype with two setae on face of sternite 5) ; sternites 9 and 10 with two corresponding seta e slightly anterior to the marginal row (Fig. 4). Trichobothria on palpal chela of holotype as shown in Fig. 5 (and Curcio 1984 : fig. 41); there is a little variation in position of trichobothria o n fixed fingers of other specimens (Fig. 6). Genital opercula (sternites 2 and 3) of male about as illustrated for holotype (Curcio 1984: fig. 20); sternite 3 of holotype with 22 setae scattere d broadly (paratypes and topotypes with 18-22 se - tae); sternite 3 of holotype with five small seta e near middle and 11 larger setae along posterio r margin (paratypes and topotypes with 5-9 smal setae near middle and 12-13 along margin); anterior margin of sternite 3 slightly concave at middle (Fig. 3), but not as distinctly indented as in Fissilicreagris chamberlini (Beier) (see Curci o 1984 : fig. 3). Genital opercula of female as show n in Fig. 8, similar to those of F. chamberlini (see Curcio 1984: fig. 4); sternite 2 with 8-11 small setae in two groups, on either side of midline; sternite 3 with 11-14 setae along posterior margin. In both sexes, on sternites 3 and 4, there ar e 4-7 small setae on each spiracular plate. Internal] genitalia of holotype male shown in Fig. 7; gen.. erally similar to those of Saetigerocreagris phyl lisae (Chamberlin) (see Chamberlin 1962 : fig. 12) Tartarocreagris texana (Muchmore) (see Much more 1992 : fig. 4), and Fissilicreagris macilent,, (Simon) (see Muchmore 1994 : fig. 4); the dors z sacs are thin-walled and not as clearly separat

MUCHMORE & COKENDOLPHER EMPIRE CAVE PSEUDOSCORPION 17 3 Figures 5 8. Fissilicreagris imperialis (Muchmore). 5, Left palpal chela of holotype (male), lateral view, showing positions of trichobothria (darkened areoles are underneath) ; 6, Right palpal chela of paratype (male), lateral view ; 7, Internal genitalia of holotype (male), ventral view (dorsal genital sacs stippled) ; 8, Central part s of sternites 2 and 3 of female (IXL Cave). as in the three species mentioned ; lateral sac s long and narrow. Internal genitalia of female no t distinguished. Measurements (mm). Male: Figures give n first for holotype, followed in parentheses b y ranges for the two paratypes and two topotypes. Body L 3.50 (2.50 3.45). Carapace L 0.935 (0.89 0.96). Chelicera L 0.52 (0.495 0.56). Palp: trochanter 0.58 (0.56-0.63)/0.215 (0.205 0.23); femur 1.12 (1.08 1.19)/0.23 (0.215 0.24); patell a 1.07 (1.00 1.11)/0.29 (0.28 0.305); chela (without pedicel) 1.73 (1.69 1.86)/0.435 (0.43 0.47) ; hand (without pedicel) 0.73 (0.70 0.815)/0.38 5 (0.38 0.43); pedicel L 0.16 (0.16 0.18); movable finger L 1.05 (1.03 1.15). Leg IV: femur + patella 0.89 (0.81 0.92)/0.215 (0.205 0.23); tibia 0.835 (0.79 0.89)/0.115 (0.11 0.125); basitarsus 0.29 (0.295 0.31)/0.095 (0.08 0.095); telotarsus 0.43 (0.40 0.445)/0.07 (0.07 0.075). Female : Figures given first for topotype, followed in parentheses by ranges for three speci - 4 Figure 9. Fissilicreagris imperialis (Muchmore), female (IXL Cave). Dorsal view (setae are transparen t and can only been seen with a microscope, not visibl e in field examinations).

174 THE JOURNAL OF ARACHNOLOG Y mens from Dolloff and IXL caves. Body L 3.3 3 (3.52 3.62). Carapace L 0.89 (1.00 1.05). Chelicera L 0.525 (0.53 0.585). Palp : trochanter 0.5 5 (0.59 0.63)/0.215 (0.22 0.26); femur 1.03 (1.09 1.21)/0.215 (0.24 0.26); patella 0.95 (1.03 1.14)/ 0.265 (0.29 0.325); chela (without pedicel) 1.6 7 (1.76 1.91)/0.43 (0.48 0.55); hand (without pedicel) 0.70 (0.76 0.87)/0.385 (0.43 0.50) ; pedicel L 0.15 (0.17 0.20); movable finger L 0.9 6 (1.04 1.16). Leg IV: femur + patella 0.79 (0.87 0.96)/0.18 (0.21 0.235) ; tibia 0.74 (0.85 0.925)/ 0.11 (0.12 0.125); basitarsus 0.27 (0.30 0.32)/ 0.08 (0.09 0.105) ; telotarsus 0.39 (0.415 0.445)/ 0.075 (0.08). Remarks. Attempting to place M. imperialis in a genus, Curcio (1984:165) stated "In most of its diagnostic characters (shape of flagellum, presence of anterior discal setae on abdomina l sternites, chaetotaxy of manducatory process, an d trichobothriotaxy), it is closest to the genus Australinocreagris Curcio 1984]." This is generally correct, but it is also true that in these sam e characters M. imperialis is very similar to representatives of the genus Fissilicreagris, which are found in the same general area of California (see Curcio 1984 :154 156 ; Muchmore 1994 :63 64). In addition, the internal genitalia of mal e M. imperialis are more like those of F. macilenta (see Muchmore 1994) and F. chamberlini than those ofaustralinocreagris grahami (Muchmore) (unpubl. obs.); in particular, the dorsal genital sac of the latter species is entire and round i n outline, while that of the first two species and M. imperialis is bilobed or divided into two separat e round sacs. Microcreagris imperialis is also sim - ilar to the two species of the genus Saetigerocreagris Curcio in respect to the male genitalia, but it differs from them in proportions of bod y and appendages and in the chaetotaxies of various parts. Its relation to the genus Tartarocreagris, which also has similar male genitalia, but is presently known only from Texas, is uncertain. It is concluded that Microcreagris imperialis Muchmore is most similar to Fissilicreagris macilenta and F. chamberlini and should be considered congeneric with them. Its major difference from them is its lack of eyes, a conditio n which is presumably an adaptation to life in caves. Field recognition. Although preserved ma - terial is required for positive identification, persons conducting a census of cave faunas or othe r work associated with protection of this specie s can be fairly certain that they have Fissilicreagris imperialis if: (1) it is in a cave in Cave Gulch, and (2) it appears like Fig. 9, eyeless, and about 3.0 3.5 mm in length. The only other pseudo - scorpion known from Cave Gulch is an undescribed, blind species of Neochthonius Chamberlin (Chthoniidae) in Empire Cave (unpubl. obs.); this is easily distinguished from F. imperialis by its much smaller size (only one-third a s long as the latter). There are other species undoubtedly present at surface locations, but thes e should have eyes. DISCUSSIO N Fissilicreagris imperialis is known only from Empire, Dolloff, and IXL caves in Cave Gulch, Santa Cruz County, California. It may occur in one or more of the other caves in Cave Gulch, but it is certainly restricted to this small, isolated karst area. The three caves are all within one - half mile of each other. In addition to the new records listed under specimens examined, it i s important to note that D. Ubick (pers. comm. 1995) observed but did not collect more than si x specimens in Empire Cave on 3 July 1993. Microcreagris imperialis was first listed by th e U.S. Fish and Wildlife Service as a candidate for review as an endangered or threatened specie s over a decade ago (Arnett 1984). It is still onl y a candidate species and therefore it receives n o substantive or procedural protection under the Endangered Species Act. Drewry (1989, 1991, 1994) continued to list this species, without a change in its status of review. The history of Empire Cave is a tragic one. It has been known and vandalized for over 120 years (Halliday 1962). According to Graham (1967), during August 1962 the entrance to the cave was capped by a cement barrier throug h which a small portal (about one meter square ) allowed access to the cave. This change in the entrance greatly decreased the available light in the entrance and presumably restricted air flo w and increased humidity. By August 1963, a dramatic shift was noted in the distribution of cav e arthropods and gastropods (Graham 1967, 1968a). Presumably the restricted entrance also altered the energy input into the cave. Graham (1968a) also noted that in 1966 the cave wa s blackened and filled with a strong odor, possibl y gasoline. He further stated that this is the most heavily vandalized cave in the state. Despite repeated attempts to seal the cave, it has been dug open in each case (Graham 1968a). Adamson (1982) referred to the caves in Cav e Gulch as small, badly vandalized, and often lit-

MUCHMORE & COKENDOLPHER EMPIRE CAVE PSEUDOSCORPION 17 5 tered and trashed caves. She went on to report on a clean-up trip to the caves (including Empir e and Dolloff caves) during March of 1982. At that time Empire Cave was "really filled with tras h including papers, wood, cigarette butts, orange peels, etc. and most dangerous, the shards of man y a beer bottle. This poor cave is reputedly used for parties etc. by UCSC students." As in so many efforts to clean caves, apparently no attentio n was given to the fauna and how this "trash" ma y be affecting them. While some of the collection s of this pseudoscorpion have been taken from th e undersides of rocks (D. Ubick pers. comm. 1995), most specimens thus far reported from Empir e Cave have been taken on wood in the cave. How many pseudoscorpions or their prey items were accidentally removed with the wood during the 1982 clean-up? It is hoped that future efforts will be better guided. Briggs & Ubick (1988) stated that Cave Gulch on the Gray Whale Ranch and nearby Empire Cave were in danger. At that time there were plans to log the area. Those authors felt that thi s activity could collapse caves and disturb root systems on which primary consumers feed, alter drainage, and block entrances. If the areas were logged it could also lead to further development. Briggs (1990) continued to state that the cav e habitat for this species was threatened by development and closure. A Timber Harvest Plan t o log the Gray Whale Ranch and an adjacent area on Empire Grade was approved in late 1994 by the California Department of Forestry (Anonymous 1995). According to the plan, harvesting can start at anytime. The owner of the ranch has stated that the entire ranch will be logged withi n four years (Anonymous 1995). Little data on the biology of pseudoscorpion s in Cave Gulch caves are available. The topotype female reported herein from Empire Cave was taken under wood at 20 C on 25 September 1972. The type series was collected "about middle of twilight zone. Each captured on the floo r either on side or bottom of wood, or in one case traveling over dripstone on floor. Temp. 53.5 F, air saturated, floor very damp." (Muchmore 1969). There are a few other observations avail - able on the habitat, but nothing specific to th e pseudoscorpion. These caves are in limeston e (Graham 1966). The Cave Gulch is subject t o intermittent flooding. During heavy flooding, al - most all of the 75 m of Empire Cave fills wit h water (Halliday 1962). Dolloff Cave's entrance is almost at the level of an intermittent side stream and must be re-excavated after every major floo d (Halliday 1962). The lower portion of Empire Cave has clay floors (Graham 1967). Graham (1967, 1968a, 1968b) provided maps to Empire Cave and recorded the conditions : 29 January 1960 (6.9 7.3 C, 94 99% R. H.), 7 August 196 2 (9.0 9.3 and 12 C, 100% R. H.), 26 August 196 3 (9.3 10.9 and 12.8 C, 96 100% R. H.). In th e same publications, he also recorded the Dollof Cave conditions as : 28 August 1963 (8.6 10. 2 and 12.6 C, 90% R. H.), 16 October 1966 (1 6 C, 85% R. H.). ACKNOWLEDGMENT S We are indebted to Darrell Ubick, Californi a Academy of Sciences, San Francisco, California ; Norman I. Platnick, American Museum of Natural History, New York, New York; and G. B. Edwards, Florida State Collection of Arthropods, Gainesville, Florida for the loan of material. We sincerely thank Norman V. Homer (Midwestern State University, Wichita Falls, Texas) and Darrell Ubick (California Academy of Sciences, San Francisco) for reviewing the manuscript. LITERATURE CITED Adamson, M. 1982. Santa Cruz clean-up. Devil' s Advocate, 15 :34 35. Anonymous. 1995. Gray Whale Ranch in holding pattern ; the bad news. Updates from Save the Gray Whale Parklands, Sierra Club, 7 :1. Arnett, G. R. 1984. Endangered and threatened wild - life and plants; review of invertebrate wildlife fo r listing as endangered or threatened species. Dept. Interior Fish & Wildlife Serv. 50 CFR, part 17. Fed. Reg., 49 :21664 21675. Briggs, T. S. 1990. Biology of northern California. Protecting California cave biology. P. 180 In NSS [Nat. Speleo. Soc.] 1990 Convention Guidebook (V. Johnson, ed.), Yreka, California. Briggs, T. S. & D. Ubick. 1988. Cavernicoles from Cave Gulch, Santa Cruz County. California Caver, 38 :43 44. Chamberlin, J. C. 1962. New and little-known false scorpions, principally from caves, belonging to the families Chthoniidae and Neobisiidae (Arachnida, Chelonethida). Bull. American Mus. Nat. Hist., 123 : 299-352. Coddington, J. A., S. F. Larcher, & J. C. Cokendolpher. 1990. The systematic status of Arachnida, exclusive of Acari, in North America north of Mexico. Pp. 5 20 In Systematics of the North American insects and arachnids: Status and needs. (M. Kosztarab & C. W. Schaefer, eds.), Virginia Agricul. Exp. Stat. Info. Ser. 90-1, Virginia Polytech. Inst. & State Univ., Blacksburg, Virginia.

176 THE JOURNAL OF ARACHNOLOG Y Curcic, B. P. M. 1978. Tuberocreagris, a new genu s of pseudoscorpions from the United States (Arach - nida, Pseudoscorpiones, Neobisiidae). Fragm. Balcanica, 10 :111-121. Curcic, B. P. M. 1983. A revision of some Asian species ofmicrocreagris Balzan, 1892 (Neobisiidae, Pseudoscorpiones). Bull. British Arachnol. Soc., 6:23-36. Curcic, B. P. M. 1984. A revision of some North American species of Microcreagris Balzan, 189 2 (Arachnida: Pseudoscorpiones: Neobisiidae). Bull. British Arachnol. Soc., 6:149-166. Curcic, B. P. M. 1989. Further revision of some North American false scorpions originally assigned to Microcreagris Balzan (Pseudoscorpiones, Neobisiidae). J. Arachnol., 17:351-362. Drewry, G. 1989. Endangered and threatened wildlife and plants ; animal notice of review. Dept. Interior Fish & Wildlife Serv. 50 CFR, part 17. Fed. Reg., 54:554-579. Drewry, G. 1991. Endangered and threatened wildlife and plants; animal candidate review for listing as endangered or threatened species. Dept. Interior Fish & Wildlife Serv. 50 CFR, part 17. Fed. Reg., 56 : 58804-58836. Drewry, G. 1994. Endangered and threatened wildlife and plants; animal candidate review for listing a s endangered or threatened species. Dept. Interior Fish & Wildlife Serv. 50 CFR, part 17. Fed. Reg., 59: 58982-59028. Graham, R. E. 1966. Observations on the roosting habits of the big-eared bat, Plecotus townsendii, in California limestone caves. Cave Notes, 8 :17-22. Graham, R. E. 1967. The subterranean niche of Pseu - dometa biologica (Arachnida ; Araneidae) in the Sant a Cruz caves, California, with comments on ecologica l equivalence in the cave environment. Caves Karst, 9:17-22. Graham, R. E. 1968a. Spatial biometrics of subterranean demes of Triphosa haesitata (Lepidoptera: Geometridae). Caves Karst, 10:21-28. Graham, R. E. 1968b. The twilight moth, Triphosa haesitata, (Lepidoptera: Geometridae) from Cali - fornia and Nevada caves. Caves Karst, 10:41-48. Halliday, W. R. 1962. Caves of California. Privately printed, Seattle, Washington. 194 pp. Harvey, M. S. 1991. Catalogue of the Pseudoscorpionida. Manchester Univ. Press, Manchester, En - gland, 726 pp. Mahnert, V. 1979. The identity of Microcreagris gigas Balzan (Pseudoscorpiones, Neobisiidae). Bull. British Arachnol. Soc., 4:339-341. Muchmore, W. B. 1969. New species and records o f cavernicolous pseudoscorpions of the genus Microcreagris (Arachnida, Chelonethida, Neobisiidae, Ideobisiinae). American Mus. Novit., 2392:1-21. Muchmore, W. B. 1992. Cavernicolous pseudoscorpions from Texas and New Mexico (Arachnida : Pseudoscorpionida). Texas Mem. Mus., Speleol. Monogr., 3:127-153. Muchmore, W. B. 1994. On four species of pseudoscorpions from California described by E. Simon in 1878 (Pseudoscorpionida: Neobisiidae, Chernetidae, Cheliferidae). J. Arachnol., 22:60-69. Manuscript received 15 July 1995, revised 12 Augus t 1995.