DOI: 10.18195/issn.0312-3162.24(4).2008.387-394 f\ccord.' ottflC WcstCrIl AlIstrllliall !V1l1SClIlll 24: 387-394 (2008) The first troglomorphic of the family Olpiidae (Pseudoscorpiones), with remarks on the composition of the family

Mark S. Harvey1,2 and Mei Chen Leng' [Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia. Email: [email protected]. 'School of Biology, University of Western Australia, Crawley, Western Australia 6009, Australia.

Abstract -A new genus and species of Olpiidae, Lillllllcolpilllll lillllllCi, is de- scribed from a pisolite mesa in the Pilbara region of Western Australia. The sole specimen exhibits troglomorphic facies with highly reduced eyes, pallid colouration and slightly attenuated appendages, differing from all other cave- dwelling olpiids which lack troglomorphic facies. The affinities of the new genus are difficult to determine, but it is most similar to a group of genera characterised by the position of trichobothrium ish: f'.SClIdollOrJlS, AlIS!rollOrJlS, XCllolpilllll, ElIryolpilllll and possibly Hc!crofpilllll. The family Olpiidae is di- vided into two subfamilies, Olpiinae and IIesperolpiinae.

INTRODUCTION (1986), who based his system on the observations of the family Olpiidae are of Heurtault (1980a, 1980b, 1982) and Heurtault virtually cosmopolitan in distribution, usually and Rebiere (1983). occurring in xeric environments under rocks, The specimen was examined with an Olympus in leaf litter or beneath the bark of trees. Cave- BH-2 compound microscope and illustrated with dwelling olpiids are, however, quite rare, with the aid of a drawing tube. Measurements were only five species recorded from subterranean taken at the highest possible magnification using habitats: Ca/oclzeirlls tenerifae Mahnert 2002 an ocular graticule. The specimen was examined by from the Canary Islands (Mahnert 2(02), and preparing a temporary slide mount by immersing l Progaryplls gracilis Mahnert 2001, P. /iliac Mahnert the specimen in 20 ;;, lactic acid at room temperature 2001, P. nigrimanlls Mahnert 2001, and P. sdifer for several days, and mounting it on a microscope Mahnert 2001 from Brazil (Mahnert 2(01). All of slide with 10 or 12 mm coverslips supported by these species have large eyes and seem to exhibit small sections of 0.25 mm or 0.50 mm diameter no troglomorphic morphological modifications. nylon fishing line. After study the specimen Among specimens collected from subterranean were returned to 75% ethanol with the dissected habitats within mesas in the Pilbara region portions placed in 12 x 3 mm glass genitalia of Western Australia was a single male of an microvials (BioQuip Products, Inc.). Images of the unusual olpiid. The specimen has reduced eyes, whole animal were taken using a Micropublisher is slightly pale in colouration, and has slightly 5.0 digital camera mounted on a Leica MZ16 attenuate appendages. To further document the microscope. subterranean pseudoscorpion fauna of Western Australia we present a description of this species Family Olpiidae Banks 1895 based upon a single male specimen. We also Subfamily Olpiinae Banks 1895 propose a division of the Olpiidae into two subfamilies, Olpiinae and Hesperolpiinae, based Remarks primarily upon the length of the venom ducts. For most of the past 80 years, the Olpiidae were The specimen examined in the present study defined by a series of characters states outlined by is lodged in the Western Australian Museum, Chamberlin (1930, 1931) and Beier (1932). Although Perth (WAM). Terminology and mensuration numerous new genera have been described since mostly follows Chamberlin (1931), with the the 1930's, the family definition has barely altered. exception of the nomenclature of the chelicera The Olpiidae were recently restricted by Judson (Judson 2(07), pedipalps, legs and with some (1992, 1(93) by the removal of the garypinines to m inor mod ifications to the term inology of the a separate family, Carypinidae, and the removal trichobothria (Harvey 1992). In particular, it of the hesperolpiines as a subfamily of C;,1l'ypidae. should be noted that the terminology for the The status of the garypinines as a family was trichobothria used by Harvey (1992) differs confirmed by Judson (200S), who referred to his slightly from that used by other workers. The unpublished thesis (Judson 1(92) in defining the setal notation of metatarsus IV follows Muchmore group. With the exclusion of the garypinines from 388 M.S. Harvey, M.C. Leng the family Olpiidae, and the transfer of a further that the type species of Xenolpium, X. pacificum five genera (Neominniza Beier 1930, Oreolpium (With 1907) from New Zealand, possessed leg Benedict and Malcolm 1978, Protogarypinus Beier morphology characteristic of the Olpiini requiring 1954, TeratolpiuJ1l Beier 1959 and Thaumatolpium the automatic synonymy of the two tribes. In Beier 1931) from the Olpiidae to the Garypinidae the meantime, Hoff (1964) included a third tribe by Harvey and Stahlavsky (unpublished data), the - Hesperolpiini Hoff 1964 - within the Olpiinae family now contains 33 genera (Table 1). based upon the long venom ducts within the Hoff (1945) divided the Olpiinae into the tribes chelal fingers, and the patella of leg I shorter than Olpiini Banks 1895 and Xenolpiini Hoff 1945 the femur, with a freely mobile joint. Hoff (1964) based upon the morphology of the first pair included Hesperolpium and Aphelolpium within of legs in which the femur are longer than the the Hesperolpiini, which was augmented by the patella in Olpiini, with a freely moveable joint, inclusion of Planctolpium Hoff 1964 by Muchmore but the two segments are subequal in Xenolpiini, (1979), previously described as a member of the thus allowing only restricted movement. Both Garypidae (Hoff 1964). Long venom ducts are tribes were rediagnosed by Hoff (1964), but were also found in a variety of other olpiids: Apolpium subsequently used only intermittently by later Chamberlin 1930 (Chamberlin 1931; Tooren researchers. They were abandoned by Heurtault 2002; Harvey, personal observation), Calocheirus (1980b), who presented new data on the type Chamberlin 1930 (Mahnert 1986, 2002), Cardiolpium species of several Old World genera, including Mahnert 1986 (Mahnert 1986), Ectactolpium Beier the type genera of the two tribes, Olpium L. 1947 (Harvey, personal observation), Nanolpium Koch 1873 and Xenolpium Chamberlin 1930. In Beier 1947 (Harvey, personal observation), particular, Heurtault (1979, 1980a, 1980b) found Progarypus Beier 1931 (Mahnert 2001; Harvey,

Table 1 Genera assigned to the Olpiidae.

Family Olpiidae Banks 1895 Subfamily Olpiinae Banks 1895 AntiLlolpium Muchmore 1991 Austrohoms Beier 1966 Banksolpium Muchmore 1986 Beierolpium Heurtault 1977 Calocheiridius Beier and Turk 1952 Euryolpium Redikorzev 1938 Halominniza Mahnert 1975 Heterolpium Sivaraman 1980 Hoffhorus Heurtault 1977 Homs Chamberlin 1930 Indolpium Hoff 1945 Leptolpium Tooren 2002 Linnaeolpiurn gen. novo Minniza Simon 1881 Neopachyolpium Hoff 1945 Nipponogarypus Morikawa 1955 Novohorus Hoff 1945 Olpiolum Beier 1931 Olpium L. Koch 1873 Pachyolpium Beier 1931 Parolpium Beier 1931 Pseudohorus Beier 1946 Xenolpium Chamberlin 1930

Subfamily Hesperolpiinae Hoff 1964 Aphelolpium Hoff 1964 Apolpium Chamberlin 1930 Calocheirus Chamberlin 1930 Cardiolpium Mahnert 1986 Ectactolpium Beier 1947 Hesperolpium Chamberlin 1930 Nanolpium Beier 1947 Planctolpium Hoff 1964 Progarypus Beier 1931 Stenolpiodes Beier 1959 Figure 1 sp. nov., holotype male Stenolpiun1 Beier 1955 Linnaeolpium linnaei (WAM T82354). Firsttroglomorphic Olpiidae 389 personal observation), Stcllolpiodcs Beier 1959 venom ducts characteristic of the rlesperolpiinae U~larvey, personal observation) and Stcllolpilllll are also found in Ceogarypidae, Carypidae, Beier 1955 (Mahnert 1984; Harvey, personal Larcidae and some Carypinidae (Allzlilyolpilll/1 observation). All of these genera, along with and Ncoal/1blyolpillllz). The lack of any features ApiTclolpilllll, f-icspcrolpilllll and Pla/lctolpilllll, arc providing firm evidence for the monophyly here regarded as hesperolpiines, based upon the of the Olpiidae suggests that the Olpiinae and criteria explicitly outlined by Hoff (1964). Hesperolpiinae may not be sister taxa, concordant Although we concur with Heurtault (1979, with the analyses of Murienne cl al. (2008), and 1980a, 1980b) that a satisfactory division between that other character systems need to be explored the Olpiini and Xenolpiini is unobtainable, we to establish their affinities and status within the agree with Hoff (1964) that the hesperolpiines Carypoidea. appear to represent a distinct diagnosable entity, which is here recognised as a subfamily of the Genus LilllwcolpiulIl gen. novo Olpiidae. The proposal by Judson (1992, 1993) that the ~Iesperolpiinae represents a member of Type species the Garypidae remains untested (but see below). Li/lllllColpilll11 lillllaci sp. novo The Hesperolpiinae contains 11 genera, whereas the relimited Olpiinae contains 22 genera (Table Diagnosis 1). Nearly all Olpiinae possess short venom ducts Lillllllcolpilll11 differ from all other olpiid genera where they generally do not reach ct in the fixed by the following combination of characters: two chelal finger or t in the moveable chelal finger. blades in the cheliceral rallum (Figure 8), reduced The most obvious exception is Mi/llliza barkizalllac eyes (Figure 2), the lack of enlarged tactile setae Mahnert 1991 from Saudi Arabia, in which the on the dorsal surface of the pedipalpal femur venom ducts are quite elongate (Mahnert 1991). (Figure 3), and the position of trichobothrium it The distinctiveness of the hesperolpiines is which is situated on the externo-dorsal face of the further emphasized by the results of Murienne fixed chelal finger (Figure 3). Lillllllcolpilllll can be ct al. (2008) who, using molecular sequence data explicitly distinguished from other olpiid genera from two nuclear ribosomal genes and one currently known to occur in Australia as follows: mitochondrial protein-encoding gene, found from Beierolpilll11 by the position of trichobothrium that the three hesperolpiines used in the study st, which is distal to sb in Lil1lllleolpilll11 and dorsal (Apolopill!1l parVlIl11 Hoff 1945, Na/lolpilll11 sp. and to sb in BeierolpizllIz (e.g. Heurtault 1982; Harvey Progaryplls sp.), formed a clade with the olpiine 1988); and from ElIrtlOlpilll11, Olpizull and XCllolpilll11 PacllllOlpilll11 sp., whereas the remaining olpiines by the position of trichobothrium cst which is (Bcicrolpilll11 bOnlcl11isszai (Beier 1966), Calocizciridills clearly basal to it in ElIryolpilll11, Olpilll11 and tcrl11itopizilllS Beier 1964, ElIryolpilll11 sp. and Xellolpilll11, but is on approximately the same level XCllolpilll11 sp.) formed a monophyletic clade that as it in Lillllaeolpilllll. There also appears to be was sister to Garypidae (Allagaryplls izClltwolci unnamed representatives of Illdolpilll11 in Australia Muchmore 1982 and Slf/lspizyro/llls apil11cllls (SLihlavsky et al. 2006; Harvey, unpublished data), Harvey 1987). This phylogeny was found with the which can be distinguished by the same criterion combined analysis of (a) the 18S rRNA, 28S rRNA and COl data, and (b) the 18S rRNA and 28S rRNA as ElIryolpilll11, Olpilll11 and Xellolpilll11. data. The COl analysis found Carypidae grouping Description with Pllcizyolpilll11 sp. and Apolpilll11 parvlIlIl, with NllllOlpilllll sp. and Progaryplls sp. grouping Male with the remaining olpiines. The placement of Pacizyolpilllll sp. with the hesperolpiines argue Cizclicera: with 5 setae on hand, all setae against the proposed division of the Olpiidae acuminate; movable finger with 1 subdistal seta; into two subfamilies, but presumably further subterminal tooth of movable finger not bifurcate molecular markers and additional taxa may help and not enlarged; rallum of 2 or 3 blades, anterior to resolve this conundrum. blade smooth except for 2 basal serrations on posterior margin; lamina exterior present, quite Further research into the affinities of both broad. olpiid subfamilies is necessary. The olpiines and hesperolpiines share a number of morphological Pedipalp: femur apparently without tactile setae. features with other garypoid families: the Fixed chelal finger with 8 trichobothria, movable short venom ducts found in olpiines are also chelal finger with 4 trichobothria; esb and isb found in Menthidae and some Carypinidae situated basally in straight row; cb and esb closely (all genera except AlIlblyolpilllll Simon 1898 spaced, less than 1 areolar diameter apart, csb and Ncoalllblyolpilll/1 Hoff 1956), and the long closer to isb than to est situated mediallv on external face of fixed finger, situated midway 390 M.S. Harvey, M.C. Leng

between et and esb; et situated subdistally; ist Chelicera: with 5 setae on hand (Figure 6), all setae situated on approximately same level of isb; ib acuminate; movable finger with 1 subdistal seta; situated dorsal to eb; st situated closer to chelal subterminal tooth of movable finger not bifurcate finger margin than b, sb and t. Venom apparatus and not enlarged; with 2 Iyrifissures on dorsal face present in both chelal fingers, venom ducts very and 1 Iyrifissure on ventral face; galea with bifurcate short, terminating in nodus ramosus almost tip and 1 sub-medial ramus; rallum composed of immediately. 2 blades, distal blade with 2 basal serrations on CephalotllOrax: carapace sub-rectangular; with posterior margin, basal blade smooth (Figure 8); 1 pair of eye-spots with very flat lenses situated serrula exterior with 16 blades; lamina exterior near anterior margin of carapace. present, moderately broad. Abdomen: pleural membrane longitudinally Pedipalp: trochanter, femur and patella striate. Tergites and sternites without any trace completely smooth, chela with several large of suture line; glandular setae absent. Spiracular granulations on mesal surface at base of fingers; helix present. setae very long and acicular; trochanter elongate, without tubercles; trochanter 2.73, femur 3.47, Genitalia: dorsal anterior glands absent; patella 2.35, chela (with pedicel) 3.60, chela ejaculatory canal atrium large; with 1 pair of (without pedicel) 3.50, hand 1.75 times longer internal glandular setae; median genital sac than broad, movable finger 1.06 times longer ovoid. than hand. Femur apparently without long tactile Legs: junction between femora and patellae I and 11 setae. Patella with three lyrifissures situated broad and apparently mobile; femur I approximately dorsally near pedicel. Fixed chelal finger with same length as patella I; tibiae III and IV without 8 trichobothria, movable chelal finger with 4 tactile seta; metatarsi III and IV with long tactile trichobothria (Figure 4): eb and esb situated seta situated very close to basal edge; setal basally; esb closer to cb than to isb; est slightly formula of metatarsus IV T-1-1-2-1 or T-1-1-3-1 or basal to it; trichobothria ib situated slightly basally T-0-1-1-3 or T-2-2-3-2; arolium much longer than to cb; ist slightly distal to isb; et situated near distal claws, not divided. end of finger; ca. 6 microsetae (chemosensory setae) present on fixed finger distal to et; sb Etymology situated closer to b than to st; t situated mid-way This genus is named for Carolus Linnaeus (1707- between st and tip of movable finger; microsetae 1778), founder of the modern system of taxonomy (chemosensory setae) not present on movable during the 250th anniversary of the publication finger; small "sensory spot" situated slightly of the 10th edition of Systema Naturae (Linnaeus distal to sb, consisting of single elliptical opening. 1758), and combined with the generic name Venom apparatus present in both chelal fingers, Olpium. It is neuter in gender. venom ducts very short, terminating in nodus ramosus almost immediately. Chelal teeth obtuse; Linnaeolpium linnaei sp. novo fixed finger with 25 teeth; movable finger with 23 teeth; accessory teeth absent. Figures 1-12 Cephalothorax: carapace (Figure 2) 1.38 times longer than broad; sub-rectangular; with 1 pair of eye- Material examined spots with flat lenses situated near anterior margin of carapace, posterior pair missing; with 16 setae, Holotype arranged 4: 4: 2: 4: 2; without furrows; with 5 pairs Australia: Western Australia: 0, Mesa K of lyrifissures. Manducatory process with 1 long (sample K0502 T4-2), near Pannawonica, 21°42'33"5, distal, 1 long sub-distal and 2 small internal setae; 116°16'16"E, 12 January 2007, D. Kamien (WAM remainder of maxilla with 6 setae. Chaetotaxy of T82354). coxae I-IV: 3: 5: 4: 5. Abdomen: pleural membrane longitudinally striate. Diagnosis Tergites and sternites without medial suture. Tergal Linnaeolpium linnaei is distinguished from all chaetotaxy: 2: 2: 2: 3: 4: 3: 4: 4: 4: TlT2TlT: 1T2Tl: 2; other Australasian olpiids by the reduced eyes. uniseriate; all setae acicular. SternaI chaetotaxy: 6: (0) 2 [1+1] (0): (0) 4 (0): 4: 4: 4: 4: 4: TlT2TlT: 1TlTlTl: Description 2; setae uniseriate and acuminate; glandular setae absent; anus not surrounded by sternite Xl. Adult male Genitalia: ejaculatory canal atrium large and Body moderately flattened. Colour with rounded; dorsal anterior glands absent; median sclerotized portions generally very pale, pedipalps genital sac ovoid, undivided (Figure 12). and anterior portion of carapace slightly darker. Legs: junction between femora and patellae I and First troglomorphic Olpiidae 391

11 broad and apparently sub-mobile; femur I about with 4 pairs of ventral setae; subterminal tarsal setae same length as patella I; fen1ur + patella of leg IV arcuate and acute; arolium much longer than claws, 3.56 times longer than broad; femora I and 11 with not divided (Figures 9-11). 2 perpendicular lyri fissures situated sub-distally; Dil11ellsiolls (111111): Male holotype: Body length tibiae III and IV with 2 moderately long tactile 1.32. Pedipalps: trochanter 0.192/0Jl70, femur setae, onc situated proximally, the other situated 0.288/0.083, patella 0.250/0.106, chela (with pedicel) sub-medially (Figure 10); metatarsi III and IV with 0.461/0.128, chela (without pedicel) 0.448, hand long subbasal tactile seta (Figure 10); metatarsus IV length 0.224, movable finger length 0.237. Chelicera with setal formula of T-1-1-2-1 (i.e. dorsal face with 0.115/(l064, movable finger length 0.090. Carapace 1 tactile seta and 2 regular setae; ventral face with 0.176/0.128; eye diameter 0.003. Leg I: femur not 2 paired setae; lateral face with 1 seta); tarsus IV

2

5

Figures 2-8 Lil1l1ilColpilll11 fil1l1ilCl sp. nov., holotype male (WAM T82354): 2, carapace, dorsal aspect; 3, right pedipalp, dorsal aspect; 4, left chela, lateral aspect; 5, left chelal fingers, detail; 6, right chelicera, dorsal aspect; 7, tip of movable cheliceraI finger, dorsal aspect; 8, right rallum, lateral aspect. Scale lines 0.1 mm (Figures 2-5), 0.5 mm (Figure 6),0.25 (Figure 8). M.S. Harvey, M.C. Leng measurable, patella 0.180/0.076, tibia 0.255/0.077, numerous Australian olpiines from a wide variety metatarsus 0.102/0.061, tarsus 0.128/0.051. Leg of habitats, we have not seen any specimens IV: femur + patella 0.249/0.070, tibia 0.173/0.047, apart from the holotype of L. linnaei that bears metatarsus 0.058/0.040, tarsus 0.090/0.031. reduced eyes. Other troglobitic pseudoscorpions have been recently recorded from the pisolitic Remarks mesas situated near Pannawonica, including Limzaeolpillm limzaei is currently known from Tyrannochthonills basme Edward and Harvey 2008 only a single location in north-western Western and Lagynochthonills aserna Edward and Harvey Australia where it was taken from a litter trap 2008 (Chthoniidae) (Edward and Harvey 2008), within a pisolite mesa, and as such can be readily Indohya sp. (Hyidae) (Harvey and Volschenk 2007), characterised as a short-range endemic species as and Ideoblothrlls pisolitus Harvey and Edward defined by Harvey (2002). The specimen displays 2007, 1. limzaei Harvey and Leng 2008, and I. sp. some troglomorphic features such as reduced eyes Mesa A () (Harvey and Edward 2007a; and pallid colouration. Despite having examined Harvey and Leng 2008). All of these species

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9 10

11

12

Figures 9-12 Lirmaeolpium linnaei sp. nov., holotype male (WAM T82354): 9, left leg I, patella-tarsus, lateral aspect; 10, left leg IV, lateral aspect; 11, left leg IV, metatarsus and tarsus. Scale lines = 0.1 mm (Figure 10),0.5 mm (Figures 9, 11), 0.05 (Figure 12). First troglol1lorphic Olpiidae 393 exhibit troglomorphic morphological features, as on Ideo{Jfollil'll.' (Pseudoscorpiones: Syarinidae) from do other troglobitic recorded from the subtl'l'ranean environments in Australia. I

Mahnert, V. (1986). Une nouvelle espece du genre Murienne, J., Harvey, M.S. and Giribet, G. (2008). Tyrannochthonius Chamb. des lies Canaries, avec First molecular phylogeny of the major clades remarques sur les genres Apolpiolum Beier et Calocheirus of Pseudoscorpiones (Arthropoda: Chelicerata). Chamberlin (Arachnida, Pseudoscorpiones). Mhnoires Molecular Phylogenetics and Evolution 49: 170-184. de la SocietC Royale Entomologique de Belgique 33: 143- Stahlavsky, E, Kral, J., Harvey, MS. and Haddad, CR. 153. (2006). A karyotype study on the pseudoscorpion Mahnert, V. (1991). Pseudoscorpions (Arachnida) from the families Geogarypidae, Garypinidae and Olpiidae Arabian Peninsula. Fauna ofSaudi Arabia 12: 171-199. (Arachnida: Pseudoscorpiones). European Journal of Mahnert, V. (2001). Cave-dwelling pseudoscorpions Entomology 103: 277-289. (Arachnida, Pseudoscorpiones) from Brazil. Revue Tooren, D. van den (2002). Pseudoscorpions Suisse de Zoologie 108: 95-148. (Pseudoscorpiones: Olpiidae) of the genus Apolpium Mahnert, V. (2002). Two new species of pseudoscorpions from Venezuela, and the genera Pachyolpium, (Arachnida, Pseudoscorpiones) from caves on Tenerife Leptolpium gen. novo and Serianus from Cura<;ao, Aruba and La Palma (Canary Islands, Spain), with some and Bonaire. Zoologische Mededelingen 76: 141-192. new records from the Canary Islands and the Azores (Portugal). Revue Suisse de Zoologie 109: 777-784. Manuscript received 26 May 2008; accepted 3 October 2008. Muchmore, W.B. (1979). Pseudoscorpions from Florida and the Caribbean area. 7. Floridian diplosphyronids. Florida Entomologist 62: 193-213. Muchmore, W.B. (1986). Redefinition of the genus Olpiolum and description of a new genus Banksolpium (Pseudoscorpionida, Olpiidae). Journal of Araclmology 14: 83-92.