"The Journal of ARACHNOLOGY OFFICIAL ORGAN OF THE AMERICAN ARACHNOLOGICAL SOCIETY VOLUME 34 2006 NUMBER 1 THE JOURNAL OF ARACHNOLOGY EDITOR-IN-CHIEF: Daniel J. Mott, Texas A&M International University MANAGING EDITOR: Paula Cushing, Denver Museum of Nature & Science SUBJECT EDITORS: Ecology — Soren Toft, University of Aarhus; Systematics — Mark Harvey, Western Australian Museum; Behavior — Gail Stratton, University of Mississippi; Morphology and Physiology — Jeffrey Shultz, University of Maryland EDITORIAL BOARD: Alan Cady, Miami University (Ohio); James Carrel, University of Missouri; Jonathan Coddington, Smithsonian Institution; William Eberhard, Univer- sidad de Costa Rica; Rosemary Gillespie, University of California, Berkeley; Charles Griswold, California Academy of Sciences; Marshal Hedin, San Diego State University; Herbert Levi, Harvard University; Brent Opell, Virginia Polytechnic Institute & State University; Norman Platnick, American Museum of Natural History; Ann Rypstra, Mi- ami University (Ohio); Paul Selden, University of Manchester (UK.); Matthias Schaefer, Universitset Goettingen (Germany); William Shear, Hampden- Sydney College; Petra Si- erwald. Field Museum; I-Min Tso, Tunghai University (Taiwan). The Journal of Arachnology (ISSN 0161-8202), a publication devoted to the study of Arachnida, is published three times each year by The American Arachnological Society. Memberships (yearly): Membership is open to all those interested in Arachnida. Sub- scriptions to The Journal of Arachnology mid American Arachnology (the newsletter), and annual meeting notices, are included with membership in the Society. Regular, $40; Stu- dents, $25; Institutional, $125 . Inquiries should be directed to the Membership Secretary (see below). Back Issues: Patricia Miller, PO. Box 5354, Northwest Mississippi Commu- nity College, Senatobia, Mississippi 38668 USA. Telephone: (601) 562-3382. Undelivered Issues: Allen Press, Inc., 1041 New Hampshire Street, PO. Box 368, Lawrence, Kansas 66044 USA. THE AMERICAN ARACHNOLOGICAL SOCIETY PRESIDENT: Elizabeth Jakob (2006-2008), Department of Psychology, University of Massachusetts, Amherst, MA 01003 USA. PRESIDENT-ELECT: Paula Cushing (2006-2008), Denver Museum of Nature & Sci- ence, Denver, CO 80205 USA. MEMBERSHIP SECRETARY: Jeffrey W. Shultz (appointed). Department of Entomology, University of Maryland, College Park, MD 20742 USA. TREASURER: Karen Cangialosi, Department of Biology, Keene State College, Keene, NH 03435-2001 USA. SECRETARY: Alan Cady, Dept, of Zoology, Miami University, Middletown, Ohio 45042 USA. ARCHIVIST: Lenny Vincent, Fullerton College, Fullerton, California 92634 USA. DIRECTORS: Gary Miller (2005-2007), Deborah Smith (2004-2006), Christopher Buddie (2005-2007). PAST DIRECTOR AND PARLIAMENTARIAN: H. Don Cameron (appointed), Ann Arbor, Michigan 48105 USA. HONORARY MEMBERS: C.D. Dondale, H.W. Levi, A.R Millidge. Cover photo: Spiderlings emerging from egg sac guarded by female southern crevice spider, Kukulcania hibernalis. Photo by Jim Carrel. Publication date: 23 August 2006 ©This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 2006. The Journal of Arachnology 34:1-36 THE WOLF SPIDERS OF ARTESIAN SPRINGS IN ARID SOUTH AUSTRALIA, WITH A REVALIDATION OF TETRALYCOSA (ARANEAE, LYCOSIDAE) Volker W. Framenau: Department of Terrestrial Invertebrates, Western Australian Museum, Perth, Western Australia 6000, Australia. E-mail: volker.framenau@museum.wa.gov.au Travis B. Gotch and Andrew D. Austin: Centre for Evolutionary Biology & Biodiversity, School of Earth & Environmental Science, The University of Adelaide, South Australia 5005, Australia ABSTRACT. Artesian springs, commonly referred to as mound springs, are isolated unique threatened wetlands in arid central Australia that harbor a large number of endemic and relict species. Wolf spiders (Lycosidae) are the dominant invertebrate predators in mound springs and are the most abundant spider family present. Nine species are common, five of which are known to occur in other Australian wetland habitats, such as river floodplains and lakeshores: Artoria howquaensis Framenau 2002, Hogna crispipes (L. Koch 1877) new combination (= Trochosa pulveresparsa (L. Koch 1877) new synonymy; = Geoly- cosa tongatabuensis (Strand 1911) new synonymy; = Tarentula tanna Strand 1913 new synonymy; = Lycosa waitei Rainbow 1917 new synonymy; = Lycosa strenua Rainbow 1920 new synonymy; = Lycosa rainbowi (Roewer 1951) new synonymy), Venatrix arenaris (Hogg 1905), V. fontis Framenau & Vink 2001, and V. goyderi (Hickman 1944). Four species commonly found in mound springs are described as new: Artoria victoriensis new species, Hogna diyari new species, H. kuyani new species, and Tetralycosa arabanae new species. Venatrix fontis and T. arabanae are mainly found at mound springs and have only rarely been recorded from other wetland habitats. Tetralycosa Roewer 1960 is revalidated with Lycosa. meracula Simon 1909 as type species. The genus is defined by its unique male pedipalp morphology with a deeply divided tegulum that carries a mesally directed spur on its retrolateral section opposing the hook- shaped median apophysis. Three Australian species are transferred to Tetralycosa: T. alteripa (McKay 1976) new combination, T. eyrei (Hickman 1944) new combination and T. oraria (L. Koch 1876) new combination {— Trochosa candicans (L. Koch 1877) new synonymy; = Lycosa meracula Simon 1909 new synonymy). Hogna pexa (Hickman 1944) new combination, an Australian wolf spider closely related to Hogna kuyani new species, is transferred from Pardosa. Keywords: Artoria, Venatrix, Hogna, systematics, new species, mound springs Central Australia is one of the driest places on earth. In the northern regions of South Australia the mean annual rainfall is between 100-150 mm and has an annual evaporation rate in excess of 3600 mm (Kotwicki 1987). The largest single source of water in this re- gion is located below the surface in an enor- mous aquifer known as the Great Artesian Ba- sin. This basin is a single continuous aquifer spanning 1.76 million km^ across the states of Queensland, New South Wales, South Austra- lia and the Northern Territory (Habermehl 1980, 1982; Harris 1992). The water from this basin discharges naturally from artesian springs (referred to as mound springs in South Australia) and artificially from free flowing bores (known locally as bore drains) (Fig. 1). These springs and bores form habitats that are analogous to islands in an otherwise desert en- vironment for species that are dependent on permanent water for their survival (Harris 1981). Artesian springs in this region form at frac- tures and fault lines along the margin of the basin creating wetlands of varying sizes. The typical artesian spring in South Australia is a low mound with water flowing from the top and forming a wetland around the base (Fig. 2). The mound is formed as water with high mineral and bicarbonate content precipitates minerals on the surface that over time create a raised area. Additionally, vegetation around 2 THE JOURNAL OF ARACHNOLOGY Figure I . — Geographical locations of the South Australian artesian springs and bore drains. (A) shows the distribution of artesian spring complexes (•, capital letters) and flowing bore brains ( 0 , lower case letters) included in this study; (B) is an expanded view of the spring complexes and bore drains around Lake Eyre South. Artesian spring complexes: A = Dalhousie, B = Neales River, C = Lake Cadibarra- wirracanna, D = Francis Swamp, E = Billa Kalina, F = Margaret River, G = Lake Eyre South, H = FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 3 the discharge region traps and collects wind- blown sand (Habermehl 1982). The flow rate from these artesian springs ranges from aL most insignificant for small soaks to greater than 50 million litres per day such as at DaL housie Springs (Sibenaler 1996). The wetland area is dependent on water flow and is cate- gorized into two areas. The vent is the area where water issues from the ground and can vary in form from an active spring with a pool of open water to a damp soak, and there can be one to a number of vents associated with a particular mound. The tail is the part of the spring that results from the outflow of water away from the vent. It can be a channel or a uniform flow radiating out from the vent, and can range in area from less than 1 m^ to great- er than 70 ha and over 16 km in length (Si- benaler 1996). These areas support substantial water dependent vegetation that is usually dominated by only one or two species (Symon 1985; Fatchen & Fatchen 1993). Artesian springs in South Australia host nu- merous endemic species of high conservation status due to their very restricted distribution and potential threats to the integrity of their fragile habitats. They include endemic gastro- pods, crustaceans and fish (Ferguson 1985; Ponder 1985; Boyd 1990; Kinhill Engineers 1997). Artesian springs are threatened by a number of human impacts, most importantly excessive water consumption by cattle, min- ing companies and gas abstraction operations. This may result in a localized reduction of wa- ter pressure in the Great Artesian Basin fol- lowed by reduced flows and, in rare instances, spring extinction (Kinhill Engineers 1997). Recent studies of artesian springs and bore drains in South Australia have shown that wolf spiders (Lycosidae) are the most abun- dant predatory group. They include a number of undescribed taxa and are associated with vegetated areas of Cyperus laevigatus, Phrag- mites australis and Typha domingensis both at the vent and on the tail (Lamb 1998; Gotch 2000). Other spider families have been re- ported from artesian springs and bore drains in lower numbers, including Hahniidae, Pi- sauridae, Linyphiidae, Clubionidae, Saltici- dae, Zodariidae, Oxyopidae, Gnaphosidae, Desidae, Corinnidae, Araneidae, Tetragnathi- dae and Prodidomidae (Lamb 1998; Gotch 2000; D. Niejalke & D. Hirst, pers. comm.). Here we provide a complete taxonomic treatment of wolf spiders of artesian springs in South Australia to facilitate their identifi- cation as part of on-going research to develop procedures for environmental monitoring of these unique habitats. METHODS Typical artesian spring lycosids as defined for this study are species which are faculta- tively dependent on the occurrence of open spring or bore water and will only be found in the confined space where it is available. These do not include the mostly burrowing species of the arid environment surrounding the springs, which on rare occasions can be found at the springs (for example Lycosa woonda McKay 1979; VWF, TBG pers. obs.). This study is mainly based on material col- lected during three studies on the arthropod communities of South Australian artesian springs (Lamb 1998; Gotch 2000) lodged at the South Australian Museum. In addition, the collections of all other major museums in Australia were examined thoroughly for con- specifics of the artesian spring species as part of an ongoing revision of the Lycosidae of Australia. Descriptions are based on specimens pre- served in 70% EtOH. Internal female genitalia were prepared for examination by submersion in 10% KOH overnight at room temperature. For clarity, the illustrations of epigyna and male pedipalps omit the setae. The morpho- logical nomenclature follows Framenau & Vink (2001) and Framenau (2002). All type material was examined unless otherwise stat- ed. All measurements are in millimeters (mm). Abbreviations. — Eyes: anterior (AE), an- terior median (AME), anterior lateral (ALE), posterior (PE), posterior median (PME), pos- terior lateral (PLE). Measurements (adult spi- ders, if not otherwise stated): total length Hermit Hill, I = DavenportAVangianna. Bore drains: a = Hamilton, b = Welcome, c = Elizabeth, d = Coward, e = Charles Angus, f = Morris Creek, g = Crows Nest, h = Coranna, i = Muloorina, j = Lake Letty no. 3, k = Clayton. 4 THE JOURNAL OF ARACHNOLOGY Figure 2. — McLachlan Springs, a typical artesian spring in South Australia showing the vent and tail microhabitats. (TL), carapace length (CL) and width (CW), abdomen length (AL) and width (AW). Aus- tralian States and Territories: Australian Cap- ital Territory (ACT), New South Wales (NSW), Northern Territory (NT), Queensland (Qld), South Australia (SA), Tasmania (Tas), Victoria (Vic), Western Australia (WA). Collections: AM = Australian Museum, Sydney; ANIC = Australian National Insect Collection, Canberra; BMNH = Natural His- tory Museum, London; CVIC = Central Vic- torian Insect Collection, LaTrobe University, Bendigo; MNHP = Museum National d’Histoire Naturelle, Paris; MV = Museum Victoria, Melbourne; QM = Queensland Mu- seum, Brisbane; QVMAG = Queen Victoria Museum and Art Gallery, Launceston; SAM = South Australian Museum, Adelaide; SMF = Senckenberg Museum, Frankfurt; TMAG = Tasmanian Museum and Art Gallery, Hobart; WAM = Western Australian Museum, Perth; ZMB = Museum ftir Naturkunde, Zentralin- stitut der Humboldt-Universitat, Berlin; ZMH = Zoologisches Institut und Zoologisches Museum, Universitat Hamburg. KEY TO LYCOSIDAE OF SOUTH AUSTRALIAN ARTESIAN SPRINGS 1. Male pedipalp with basoembolic apophysis that reaches around the base of the median apophysis; median apophysis with a long narrow base, originating apically at tegulum (Figs. 59, 65, 67); epigynum a simple posterior atrium that is sclerotized laterally (Fig. 61), or with a posterior sclerotized rim that reaches anteriorly into a white, oval center (Fig. 68) Genus Artoria Thorell 1 877 .................................................. 8 Male pedipalp without basoembolic apophysis, median apophysis originating laterally at tegulum (Figs. 4, 11, 15, 20, 35, 42), or a basally directed broad hook (Fig. 49); epigynum inverted T-shaped (Figs. 8, 13, 25, 27-31, 38, 45), or a triangular atrium (Figs. 17, 54, 55) 2. Tegulum of male pedipalp with deep and wide longitudinal division in retrolateral half, median apophysis forms a basally directed hook opposing a distinct tip on the retrolateral part of the tegulum (Fig. 49); female epigynum forms a triangular atrium, hoods clearly FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 5 eyes (PLE) posterior lateral Posterior m^ian eyes (PME) anterior lateral eyes (ALE) anterior median eyes (AME) cymbium copulatory spermatheca posterior lateral part M ' Figures 3-9. — Venatrix arenaris (Hogg 1905): Male from Horse Springs, SA (WAM T47290): 3. hab- itus; 4. left pedipalp, ventral; 5. left pedipalp, retrolateral; 6. eye arrangement; 7. apical part of bulb. Female from Fred Springs, SA (WAM T47292): 8. ventral view of epigynum; 9. dorsal view of epigynum. Scale bar: (3) 2.07 mm, (4, 5) 0.58 mm, (6) 0.98 mm, (7) 0.36 mm, (8, 9) 0.46 mm. separated (Figs. 54, 55); carapace and abdomen light yellowish-brown, carapace with in- distinct dark radial pattern, abdomen with indistinct white patches (Fig. 48); small spiders; TL 4.8-11.5 mm. Main distribution at artesian springs, occasionally near salt lakes (only recorded from SA) ................................ Tetralycosa arabanae new species Tegulum not divided, median apophysis directed retrolaterally, much broader at the base than tip and with a ventrally directed process (e. g. Figs. 4, 11, 15, 20, 35, 42); epigynum inverted T-shaped (e. g. Figs. 8, 13, 25, 27-31, 38, 45), or a triangular atrium with distinct anterior hoods separated from atrium (Fig. 17); carapace brown with light median band or uniformly dark grey to black; small to medium-sized spiders; TL 5.5-20.0 Subfamily Ly- cosinae 3 3. Tip of male cymbium with large, claw-like setae (Figs. 4-5, 11, 15), outer edge of fangs in males with tubercle (Fig. 16); posterior lateral edges of epigynum bulging anteriorly (Figs. 8, 13), or epigynum a triangular atrium with distinct anterior hoods separated from atrium (Fig. 17) Genus Venatrix Roewer 1960 4 Tip of male cymbium without claw-like setae, but with a variable number of macrosetae (Figs. 20, 21, 35, 36, 42, 43); posterior lateral edges of epigynum not bulging anteriorly, i.e. posterior lateral parts thickest at their base near the median septum (Figs. 25, 27-31, 38, 45) Genus Hogna Simon 1885 6 4. Carapace brown with a wide median band that constricts anteriorly of fovea and forms a star-like pattern around the fovea (Fig. 3); terminal apophysis of the male pedipalp forms a large roof over the tip of the embolus (Figs. 4, 7); bulging posterior lateral ends of epigynum whitish, median septum of equal width along its whole length (Fig. 8). TL 8.0-15.0. Aus- 6 THE JOURNAL OF ARACHNOLOGY tralia-wide on sand and small gravel near rivers, ponds and springs (Fig. 10) ........... Venatrix arenaris (Hogg 1905) Carapace brown to dark brown with a narrow light brown median band ............... 5 5. Terminal apophysis of male pedipalp sickle-shaped (Figs. 11, 12); epigynum inverted T- shaped, the median septum widening anteriorly (Figs. 13); TL 8.0-17.0 mm. Mainly at artesian springs, rarely found near rivers (NSW, SA, Vic) (Fig. 14) ............................................ Venatrix fontis Framenau & Vink 2001 Terminal apophysis of male pedipalp forms a roof over the tip of the embolus (Figs. 15); female epigynum a triangular atrium (Fig. 17); TL 5.0—11.0 mm. Open, vegetated areas near water, Australia-wide (Fig. 18), also in New Zealand and New Caledonia ................................................. Venatrix goyderi (Hickman 1944) 6. Carapace dark reddish-brown, appears dark grey to black due to a dense cover of silver- grey setae (in particular in fresh material); no light median band; abdomen dark grey with indistinct light and dark patches (Fig. 41); pedipalp Figs. 42-44; epigynum Figs. 45, 46; TL 8.5-20.0 mm. Near water (SA, Qld, NSW, WA) (Fig. 47) . . . Hogna kuyani new species Carapace brown with a distinct light median band 7 7. Light median band on carapace wide, covering approx, one third of carapace width (Fig. 33); venter yellow with two black spots behind epigastric furrow and a variable number of black spots laterally (Fig. 34); pedipalp Figs. 35-37; epigynum Figs. 38, 39; TL 9.5-18.0 mm. Near water (SA, Qld, NSW, Vic) (Fig. 40) ............... Hogna diyari new species Light median band on carapace narrow, covering less than a quarter of carapace width (Fig. 19), submarginal band with three dark blotches (sometimes not very distinct); venter uni- formly yellow-brown; pedipalp Figs. 20, 21, 24; epigynum Figs. 25-31; TL 7.0-20.0 mm. Open areas near water on sand or grass, inland and coastal (Australia-wide, including off- shore islands and reefs (Fig. 32); also in New Zealand and Pacific islands) ......... Hogna crispipes (L. Koch 1877) 8. Median apophysis of male pedipalp with triangular apical process and a broad, ventrally bent tip (Figs. 59); pedipalp patella and tibia bright yellow; pedipalp tibia and basal half of cymbium with dense cover of white setae (very conspicuous in unpreserved specimens); epigynum forms an indistinct, lightly sclerotized posterior atrium (Fig. 61); carapace black with light marginal bands due to a dense cover of white setae; TL 3. 5-6.0 mm. Open, but shaded areas near water, mound springs and lowland river floodplains (SA, Vic) (Fig. 62) Artoria howquaensis Framenau 2002 Median apophysis of male pedipalp in ventral view shaped like an upside-down sock (Fig. 65), pedipalp patella light brown, cymbium without white setae; epigynum forms an oval atrium with a sclerotized posterior rim that reaches into the center of the atrium (Fig. 68); carapace brown with light median and submarginal bands and dark radial pattern (Fig. 63); femora of all legs with dark annulations (particularly distinct on ventral side of leg III and IV); TL 3. 5-8. 5 mm. Rare at artesian Springs, but very common in open, moderately moist cultural landscapes and suburban areas (NSW, SA, Tas, Vic) (Fig. 70) .................................................. Artoria victoriensis new species TAXONOMY Subfamily Lycosinae Simon 1898 Remarks, — The Lycosinae appear to be well-defined since Dondale (1986) established synapomorphic characters for the male pedi- palp (p. 331): “median apophysis transverse, with ventrally directed spur” and “median apophysis with sinuous channel on dorsal sur- face”. However, there are difficulties in estab- lishing monophyletic taxa below the subfam- ily level. Molecular analysis suggests that Dondale’s (1986) "Trochosa-' and Hycosa- groups’ within the Lycosinae, based on the shape of the terminal apophysis, are paraphy- letic (Vink et al. 2002). Alternatively, Zyuzin (1993) distinguished his tribes Trochosini and Lycosini based on the shape of the median apophyis (Tegular apophysis, TA’ sensu Zyu- zin 1993) and the female epigynum. Genus Venatrix Roewer 1960 Venatrix Roewer 1960: 745 (name first listed as a nomen nudum in Roewer 1955: 307). FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 7 Figure 10. — Records of Venatrix arenaris (Hogg 1905) in Australia. Remarks* — Venatrix was established by Roewer (1960) and recently revised to include 22 Australian species, of which one, V. goy- deri, is also found in New Zealand and New Caledonia (Frameeau & Vink 2001; Vink 2002; CJ. Vink, pers. comm.). Males of Ven- atrix have a tubercle on the outer edge of the fangs (Fig. 16) and large, claw-like setae at the tip of the cymbium (Figs. 4, 5, 11, 15). Three species of Venatrix are present at arte- sian springs and bore drains of South Austra- lia, V. arenaris, V. fontis and V. goyderi. Full taxonomic bibliographies for these species can be found in Framenau & Vink (2001), but up- dated distribution maps are provided here. Venatrix arenaris Hogg 1905 Figs. 3-10 Lycosa arenaris Hogg 1905: 586-588, fig. 88; Mc- Kay, 1974: 1-6, figs. la~m. Lycosa celaenica Rainbow 1917: 488-489, plate 32, figs. 10, 11. Venatrix arenaris (Hogg 1905): Framenau and Vink 2001: 960-962, figs. 40a-f, 41. Diagnosis. — Venatrix arenaris is a medi- um-sized spider (TL 8.0-15.0). Its mottled, indistinct coloration varies from very dark to light beige (Fig. 3) and blends very easily with its preferred sandy habitat. Most speci- mens, except very dark spiders, have a light narrow band on the anterior half of the ster- num. Males are distinguished by their broad terminal apophysis, which bends ventrally forming a roof over the tip of the embolus (Figs. 4, 5, 7). The female epigynum forms an inverted ‘T’, with a narrow median septum. The lateral tips of the posterior transverse part bulge anteriorly (Fig. 8). Distribution and habitat preferences. — Venatrix arenaris is found Australia-wide (Fig. 10). It is present in most artesian springs and bore drains in South Australia, and is the dominant species in the south-eastern springs from the Blanche Cup in the west to Mulligan Springs in the east (Table 1). Within springs K arenaris typically resides next to the edges of open wet spaces and small open water pools. It is rarely active during the day and usually conceals itself under C. laevigatas. At night this species can be observed foraging on the surface of still water. Remarks. — Recent preliminary allozyme studies indicate that V. arenaris populations from the South Australia and those inhabiting lowland floodplains of the Great Dividing Range in south-eastern Australia (illustrated in Framenau & Vink 2001) possibly represent two different species (Gotch 2003; M. Adams pers. comm.). Thus, the status and distribution of V. arenaris as presently defined on mor- phological grounds needs to be reassessed in conjunction with more detailed allozyme stud- ies. Representative male and female speci- mens from South Australia are illustrated here (Figs. 3-9). Venatrix fontis Framenau & Vink 2001 Figs. 11-14 Venatrix fontis Framenau & Vink 2001: 959-960, figs. 38-f, 39. Diagnosis. — This is a medium- sized wolf spider (TL 8.0-17.0 mm). The carapace varies from nearly black to a light olive-grey and a narrow, yellow median band is always pre- sent. The abdomen is dark grey and has a lan- ceolate yellow heart mark in its anterior half. The body coloration resembles V. goyderi, however, V. fontis is generally larger. In con- trast to V. arenaris and V. goyderi, the ter- minal apophysis of the male pedipalp of V. fontis is sickle-shaped (Figs. 11, 12). The fe- male epigynum is inverted T-shaped, but in contrast to V. arenaris, its median guide wid- ens anteriorly (Figs. 13). Distribution and habitat preferences. — Venatrix fontis appears to have its main dis- tribution at the South Australian artesian springs; however, single specimens have been found in Victoria and New South Wales (Fig. 14). It is the dominant species in the western and northern springs, from Coward Springs in the south to the Mt. Dutton spring complex in the north (Table 1). Venatrix fontis is a noc- 8 THE JOURNAL OF ARACHNOLOGY embolus embolus median / septum posterior lateral part Figures 11-13. — Venatrix fontis Framenau & Vink 2001: Male from Freeling Springs, SA (SAM NN9908): 11. left pedipalp, ventral; 12. apical part of bulb. Females from Freeling Springs, SA (SAM NN9910): 13. ventral view of epigynum. Scale bar: (11) 0.59 mm, (12) 0.29 mm, (13) 0.57 mm. turnal species that is associated with less densely vegetated springs, especially those with gravel or travertine substrates. During the day large adult V. fontis can be found under sheets of travertine and rocks while juveniles shelter in clumps of C. laevigatus at the spring margins, Venatrix goyderi (Hickman 1944) Figs. 15-18 Lycosa goyderi Hickman 1944: 33-34: plate 2, fig. 20. Lycosa howensis McKay 1979b: 237-238, figs. la-e. Figure 14. — Records of Venatrix fontis Framenau & Vink 2001 in Australia. Venatrix goyderi (Hickman 1944): Framenau & Vink, 2001: 963-965, figs. 44a-e, 45. Diagnosis. — -This is the smallest (TL 5.0- 11.0 mm) of the three Venatrix species found regularly at South Australian artesian springs and bore drains. This species is brown to dark brown. The carapace has a narrow, light me- dian band. The abdomen bears a light median heart mark and pairs of light brown patches. The terminal apophysis of the male pedipalp forms a roof-like structure over the embolus (Fig. 15). The female of V. goyderi is the only member of the subfamily Lycosinae at the ar- tesian springs that does not have an inverted T-shaped epigynum (Fig. 17). Distribution and habitat preferences. — Venatrix goyderi has been found in all states of mainland Australia as well as Lord Howe Island (Fig. 18), the North Island of New Zea- land (Framenau & Vink 2001) and recently in New Caledonia (C.J. Vink pers. comm.). In the arid zone of South Australia it is found associated with wetlands across the north-east of the state, particularly the Coopers Creek and Diamantina River systems, and with ar- tificial wetlands such as bore drains where it is the dominant wolf spider. This species is FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 9 Figures 15-17. — Venatrix goyderi (Hickman 1944): Male from Howqua River, Vic (AM KS58209): 15. left pedipalp, ventral; 16. fang with tubercle. Female from Howqua River, Vic (AM KS58206): 17. ventral view of epigynum. Scale bar: (15) 0.43 mm, (16) 0.41 mm, (17) 0.31 mm. also found in large numbers at springs that have been exposed to significant disturbance from over grazing, dredging or from severe floods such as at Buttercup Springs (Table 1). Remarks. — The holotype female of V. goy- deri had been reported lost (McKay 1985; Framenau & Vink 2001), however, it was re- cently discovered at the Australian Museum in Sydney (AM KS49705, VWF, examined) confirming the identity of this species. Genus Hogna Simon 1885 Remarks. — Hogna was first listed by Si- mon (1885), and subsequently (Simon 1898: 347) defined mainly based on somatic char- acters, in particular the arrangement of the eyes and the correlation of the length of leg segments of the fourth leg. The type species is H. radiata (Latreille 1817), a common spe- cies in the Mediterranean region, that is found across Central Asia and Central Africa (Plat- nick 2004). Currently, Hogna includes more than 200 species (Plateick 2005), however, the genus is in need of revision (Dondale & Redner 1990). Here we place three lycosids from South Aus- tralian artesian springs in this genus due to the similarity of their male and female genitalia with those of H. radiata as illustrated by Fuhn & Niculescu-Burlacu (1971) and Miller (1971). One of the artesian spring species, H. crispipes, is transferred from Lycosa Latreille 1804, the two other species are new to sci- Figure 18. — Records of Venatrix goyderi Hick- man in 1944 Australia. ence, H. diyari new species and H. kuyani new species. The examination of type material of Aus- tralian wolf spiders to establish the identity of the artesian spring species revealed that the genitalic morphology of Pardosa pexa Hick- man 1944 (holotype male, AM KS17123, from 'Burts Waterhole’ (SA), examined by VWF) is very similar to the species here placed in Hogna. Consequently, we transfer Pardosa pexa to Hogna: Hogna pexa (Hick- man 1944) new combination. This new ge- neric placement also reflects the true subfam- ily status of this species, as it clearly belongs to the Lycosinae and not Pardosinae {sensu Dondale 1986). Hogna crispipes (L. Koch 1877) new combination Figs. 19-32 Lycosa crispipes L. Koch 1877: 923-925, plate 79, figs. 8, 8a, plate 80, figs. 1, la; Rainbow 1911: 266; McKay 1979a: 253, figs. 4e-f, m; McKay 1985: 76; Platnick 1989: 370. not Lycosa crispipes L. Koch 1877 sensu McKay 1979a 252-255, figs. 4a-d, g-1 (misidentification, not L. crispipes but two undescribed species). Lycosa pulvere-sparsa L. Koch 1877: 941—942, plate 79, fig. 2; Rainbow 1911: 272. NEW SYN- ONYMY. Tarentula tongatabuensis Strand 1911: 207; Strand 1915: 258, plate 14, fig. 21, plate 19, fig. 99. NEW SYNONYMY Tarentula tanna Strand 1913: 121-122; Strand 1915: 260, plate 19, fig. 96a-b; Ledoux & Halle 1995: 7. NEW SYNONYMY. Lycosa waitei Rainbow 1917: 487-788, plate 32, figs. 7-9; Roewer 1955: 272; McKay 1973: 380; Bonnet 1957: 2669; McKay 1985: 84. NEW SYNONYMY 10 THE JOURNAL OF ARACHNOLOGY Table 1 . — Distribution and relative abundance of lycosid species at a selection of artesian springs and bore drains in South Australia ( + + + dominant species, ++ subordinate species, + rare species); see Fig. 1 for geographical location for each site. Species Artoria Artoria Tetraly- Venatrix how- victor- Hogna Hogna Hogna cosa Venatrix Venatrix goyderi quaensis iensis crispipes diyari kuyani arabanae arenaris fontis (Hick- Framen- new (L. Koch new new new (Hogg Fr. & V. man Sample Locations au 2002 species 1877) species species species 1905) 2001 1944) Artesian Springs Dalhousie (A) Dalhousie Kingfisher + + + + + Neales River (B) Freeling + + + + + Hawker + + + + Outside + + + + + Lake Cadibarrawirracanna (C) Lake Cadi + + Francis Swamp (D) Big Depot Francis + + + + + Swamp + + + + + + + + + Billa Kalina (E) Billa Kalina + + + + + Margaret River (F) Blanche Cup + + + + + + + + Bubbler + + + + + + + Buttercup + + + + + + + Coward + + + + Elizabeth + + + + + + + + Horse + + + + T + + + + + Jersey + + + + + + + + + + + Kewson Hill + + Little Bubbler + + + + Lake Eyre South (G) Fred + + + + + + + + + + + + Gosse + + + + + + + + + + + + McLachlan + -f + + + + Smith + + + Hermit Hill (H) Bopeechee + + + + + Dead Boy + + + + + Hermit Hill + + + + + + + + + T Old Finniss + + + + + + + + + Old Woman + + + + + Sulphuric + + + + + + West Finniss + + + + + + + Wangianna/Davenport (I) Davenport + + T Welcome + + + FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 1 Table 1. — Continued. Species Artoria Artoria Tetraly- Venatrix how- victor- Hogna Hogna Hogna cosa Venatrix Venatrix goyderi quaensis iensis crispipes diyari kuyani arabanae arenaris fontis (Hick- Framen- new (L. Koch new new new (Hogg Fr. & V man Sample Locations au 2002 species 1877) species species species 1905) 2001 1944) Bore Drains Hamilton (a) + + + Welcome (b) Elizabeth (c) + + + + + + + Coward (d) Charles + + + + + + Angus (e) Morris Creek + + + + + + + + (f) Crows Nest + + + + + + + + + + + + + + + (g) + + Coranna (h) Muloorina (i) Lake Letty + + + + + #3 (j) + + Clayton (k) + + A + + + Lycosa (?) immansueta Simon 1909: Rainbow 1915: 787 (misidentification). Lycosa strenua Rainbow 1920: 260-261, plate 30, figs. 92-93 (preoccupied by Lycosa strenua Nic- olet 1849 and Lycosa strenua Thorell 1872). NEW SYNONYMY. Lycosa tanna (Strand 1913): Berland 1938: 182- 183, figs. 147-149; Bonnet 1957: 2666. Tarentula rainbowi Roewer 1951: 442 (replacement name for Lycosa strenua Rainbow 1920). NEW SYNONYMY Hygrolycosa crispipes (L. Koch 1877): Roewer 1955: 261; Rack 1961: 37; McKay 1973: 380. Lycosa rainbowi (Roewer 1951): Roewer 1955: 272; McKay 1985: 82. Scaptocosa tongatabuensis (Strand 1911): Roewer 1955: 291. Varacosa pulveresparsa (L. Koch 1877): Roewer 1955: 305; Rack 1961: 38; McKay 1973: 381. Varacosa tanna (Strand 1913): Roewer 1955: 305; Chrysanthus 1967: 424, figs. 73, 78-79. Lycosa tongatabuensis (Strand 1911): Bonnet 1957: 2667. Lycosa pulveresparsa L. Koch 1877: McKay, 1985: 82. '‘‘Lycosa'' tongatabuensis (Strand 1911): Ledoux & Halle 1995: 7, figs. 5a-c. Geolycosa tongatabuensis (Strand 1911): Platnick 1998: 554; Vink 2002: 36-37, figs. 31, 38, 65, 92. Types examined. — Lectotype (designated here) of Lycosa crispipes, 1 female, Queens- land, Bowen, 20°00'S, 148°14'E, BMNH, 1919.9.18.222. Paralectotype of Lycosa cris- pipes, 1 female, Queensland, Bowen, 20°00'S, 148°14'E, Museum Godeffroy 14572, Rack (1961)-catalogue 450 (ZMH). Syntype of Lycosa pulveresparsa, 1 fe- male, Rockhampton 23°22'S, 150°30'E, Mu- seum Godeffroy 14554, Rack (1961)-cata- logue 476 (ZMH). The whereabouts of a second syntype of Lycosa pulvere-sparsa from ‘Bradley's Collection’ listed by L. Koch (1877) is unknown to VWE Lectotype (designated by Vink 2002) of Tarentula tongatabuensis, 1 female, Tonga, Tongatupu Nuku’alofa, 21°07'S, 175°12'W, 4.vi.l909, E Wolf, 1909 (SMF 2199). Para- lectotype of Tarentula tongatabuensis, 1 ju- venile, same data as lectotype (SMF). Holotype of Tarentula tanna, 1 female, Va- nuatu, Tanna, 19°30'S, 169°20'E, 23.V.1909, E. Wolf (SMF 2167). Holotype of Lycosa waitei Rainbow 1917, 1 female. South Australia, Coopers Creek, ca. 28°23'S 137°4UE, September/October 1916, South Australian Museum Expedition to the Interior (SAM NN380). Holotype of Lycosa strenua Rainbow 1920, 12 THE JOURNAL OF ARACHNOLOGY Figures 19-31. — Hogna crispipes (L. Koch 1877): Male from Cullyamurra Waterhole, SA (SAM NN 13955): 19. habitus; 20. left pedipalp, ventral; 21. left pedipalp, retrolateral; 22. eye arrangement; 23. lateral view of carapace; 24. apical part of bulb. Female: 25. ventral view of epigynum (WAM T47310, from Fred Springs, SA); 26. dorsal view of epigynum (WAM T47310); 27. ventral view of epigynum (lectotype from Bowen, Qld; BMNH 1919.9.18.222); 28. ventral view of epigynum (paralectotype from Bowen, Qld; ZMH 450); 29. ventral view of epigynum (lectotype of Tarentula tongatabuensis Strand FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 13 1 female, New South Wales, Norfolk Island, 29°02^S, 167°57'E, A.M. Lea, December 1915-January 1916 (SAM NN277). All types examined. Other material examined. — AUSTRA- LIA: South Australia: 1 S , Bakers Creek, N of Wilpena (QM S2I408); 1 6, 1 $, 1 juv. Charles Angus Bore (SAM NN 13945-6); 22 d, 8 $, 1 $ with eggsac, 39 juv., Coongie Lake (SAM NN 13979-82, NN 13983-4006, NN14108-10); 16,19, Coongie Lake, L77 km W (SAM NN13977, NN20979); 1 $, Coongie Lake, 50 m SSW (SAM NN13976); 1 $, Coongie Lake, 700m E (SAM NN13978); 1 $, Coongie Lake, 7 km SE (SAM NN13975); 6 c3, 8 juv., Coranna Bore (WAM T47304, T47308-9); 1 $ with 116 spiderlings. Coward Springs (SAM NN13948); 3 ?, 1 juv., Culburra (QM S61110); 15 d, 5 9, 1 juv. (SAM NN13955- 74); 1 d, Dickinna Hill, 9.5 km SW (SAM NN13954); 1 9, Elizabeth Springs (Nth B) (SAM NN13938); 5 9, Einnis Springs (SAM NNl 3 139-40); 6 d, 2 9, Fred Springs (SAM NN13941-2, WAM T47302-3, T47305, T47307, T47310); 1 d, Freeling Spring (SAM NN13947); 1 d, Gosse East Spring (SAM NNl 3937); 1 d, 1 juv., Greenfields Wetlands, Dry Creek, Salisbury (SAM NN14010); 1 9, Johnsons Dam, Granite Downs Station (SAM NNl 3953); 1 d, 1 juv., Karroongooloo Sta- tion, via Adelaide (MV K8147); 1 d, Lake Hope channel, 3.9 km S Lake Appadare (SAM NN13949); 1 d. Lake S Siccus River (Koonamoore Station?) (SAM NN14007); 1 d, Maslins Beach (SAM NNl 40 12); 1 d, 2 9, Morris Creek Bore (SAM NNl 3943-4; WAM T47306); 1 d, Mt Fairview, Paney Sta- tion (SAM NN14008); 1 9, Scott Creek Weir (AM KS32122); 1 9, Todmordon, 90 miles W Oodnadatta, Capt. SA White Expedition, published in Rainbow (1915) as Lycosa (?) immansueta (SAM NN411); 1 d. Twin Hill (SAM NNl 3951); 1 9, Windsor Gardens, Adelaide (SAM NN14011). New South Wales: 2 9, no location (NSW?), W.J. Rainbow man- uscript no. 115 (AM KS84107); 1 9, no lo- cation (NSW?), W.J. Rainbow manuscript no. 78 (AM KS84109); 1 9, Armidale (AM KS84106); 15 d, 17 9, Bowra Station, 350 m past entrance, N of Carinda (AM KS76337-40, KS76743); 1 9, Broken Hill (SAM NN14123); 1 d, Clarence River, Cop- manhurst (SAM NN14013); 1 9, Coolaba Ramsey Park (AM KS42374); 1 d, Coota- mundra (AM KS84103); 1 d, 1 9, Darling River, 1.5 km South of Trilby Station (AM KS76557, KS76562); 3 d, 1 9 , Gwydir High- way, 300 m N of Minnamurra Station turnoff (AM KS76554, KS76561, KS76564); 2 d, 1 9, Hunter Valley AM KS7322); 1 9, Lord Howe Island (AM KS68547); 3 d, 1 9, Merri Merri Creek, 2.5 km North of Quambone (AM KS76553, KS76559-60); 14 d, 3 9, Mullin- gar Station, Lower Murray-Darling region (AM KS67036-7); 1 d, 1 9, 5 juv., Narrabri (AM KS84102); 3 d, 3 9, Narran Lakes Re- serve access track, 6.5 km from Narran Lakes Road (AM KS76550-2, KS76563); 3 9,19 with eggsac, Norfolk Island (AM KS43951- 2, KS43954, KS68883); 1 9, Norfolk Island, Burnt Pine (AM KS49891); 2 9, Norfolk Is- land, Captain Cook Memorial (ANIC); 1 9, Norfolk Island, Duncombe Bay (AM KS49895); 2 9, Norfolk Island, Mill Road (AM KS43953); 1 9, Nyngan-Canonba Road, 2.9 km South of Fairview Station junction (AM KS76555); 1 d, Road to Wanaaring, 12.7 km W of Mitchell Hwy junction (AM KS76284); 1 d, Spring Hill Station, Lower Murray-Darling region (AM KS66736); 1 d, Sturt National Park, 19.2 km S of Fort Gray Homestead on Cameron Corner Rd (AM KS84105); 1 d, Warren-Quambone Road, 0.7 km N of turnoff to Wyndabyne Station (AM KS76556); Northern Territory: 1 9, Cox Riv- er (SAM NN13129); 1 9, Curtin Springs (ANIC); 1 9, 2 juv., Tobermory Station, No. 8 Dam (QM S61119). Queensland: 1 9, Ap- pel Channel, Mornington Island (SAM NN14015); 1 d, 2 juv., Barrow Creek (QM S21407); 6 9, Birdsville (QM W7186); 1 d, 1 9, Birdsville, near town (QM W6117); 2 d, 1 9 with eggsac, 2 9,1 juv., Bowen (QM <— from Tonga; SMF 2199); 30. ventral view of epigynum (holotype of Lycosa watei Rainbow from Coopers Creek, SA; SAM NN380); 31. ventral view of epigynum (holotype of Lycosa strenua Rainbow from Norfolk Island, NSW; SAM NN277). Scale bar: (19) 2.13 mm, (20-21) 0.48 mm, (22) 1.37 mm, (23) 2.55 mm, (24) 0.14 mm, (25-31) 0.65 mm. 14 THE JOURNAL OF ARACHNOLOGY S21412); 1 9, Bushy Island, Great Barrier Reef (QM S61116); 1 9, Cape Tribulation (QM S61108); 1 9, Claudie River mouth (QM S61131); 1 d, 2 9, Cluny Station Bil- labong (QM S61066); 1 9, Coopers Creek, between Cluny Station and Monkira (QM S61104); 3 (J, 4 9, 7 juv., Curtis Island, S end of township (QM S61096S61 103); 1 d, Eulo, ‘Cookara’ (QM S61098); 1 9 with eggsac, 2 9, Eurithethera Soak, Toomba Range (QM W7185); 2 9,1 juv., Farmer Is- land, Great Barrier Reef (QM S61128); 1 9, 1 Juv., Frederick Reef, North Reef Cay, Coral Sea (ANIC); 26 d, 7 9, 2 juv., Gatton, Queensland Agricultural College (QM S61069~71, S61074-9, S61081-5, S61087- 90, S6112--4); 1 d. Grey Range, central tank, ‘Orient’ (QM S61099); 2 9, Halfway Islet, Great Barrier Reef (QM S61126); 1 d, 3 9, 1 juv., Hannah Point, North Molle Island (QM S61100); 6 d, 1 9, Jondaryan, 20 km S (QM W7189); 3 d, 5 9, 2 juv., Jumbo Bore, ‘Nor- ley’, Thargomindah (QM S61101); 1 d. Lake Broadwater (QM S61095); 1 d, Lake Broad- water, near cottage (QM S61080); 1 9, Lake Hutter, N of Aramac (QM S61113); 1 d, 1 9, Lake Nuga Nuga (QM S61068); 1 d, Long- reach (SAM NN14014); 1 9, Lucinda (QM S21414); 19,1 juv., Lydeman Island, Great Barrier Reef (QM S61129); 1 9, MacArthur Cay, Great Barrier Reef (QM S61121); 2 9, Magra Islet, Great Barrier Reef (QM S61 127); 1 9 with eggsac, 1 juv., Maydelaine Island (ANIC); 28 d, 31 9, 4 juv., Muncoonie Lakes (QM W6413-6, W7187); 1 d, Mundingburra (AM KS86384); 1 d, Murrumba Downs (QM S61093); 1 9, Pelican Island (QM S61125); 1 d, 1 9, 2 9 with eggsac, 3 juv., Raine Is- land (WAM T55434; QM S61073, S61145- 7); 1 9,1 juv., Saunders Islet, Great Barrier Reef (QM S61118); 1 9, Sherrard Island, Great Barrier Reef (QM S61 120); 1 9 , Stainer Islet (QM S61107); 1 9, Thargomindah (QM W7188); 1 9, 1 juv,, Thursday Island, Nth side (QM S 17225); 1 9, Tingalpa (QM S26112); 2 9, Townsville, common wetlands (QM S61094); 1 9, Townsville, Community Environmental Park (QM S61114); 1 9, 1 9 with eggsac, 1 9 with spiderlings, Townsville, near Fishermans Wharf (QM S61106, S61 133); 1 d. Turtle Islet, Lihou Reef, Coral Sea (ANIC); 1 9, Vanrook Station, Gilbert River Crossing West side (AM KS44298); 1 9, Wenlock River (QM S21409). Victoria: 1 d, Avon River near Valencia Creek (WAM T47111); 2 d, 1 9, Barmah Forest (WAM T47112-3); 3 d, 2 juv., Booths Rd, 0.2 km S Murray Valley Hwy (MV K8771); 4 9, Mur- ray Valley Hwy, 0.3 km NNW Walshs Bridge (MV K8691); 1 9, Murray Valley Hwy, Deep Ck Crossing (MV K8774); 1 9, Redcliffs (MV K8258). Western Australia: 1 d, 1 9, Amelia Heights (WAM 69I2Q12, 71/939); 1 9 with spiderlings, Argyle Downs Homestead, edge of Behn River (QM W5058); 2 9, Ash- more Reef, East Islet (AM KS68684); 1 d, 3 9, Attadale (WAM 71/900, 71/985-6, 71/ 1448); 1 9, Avon River, Northam (WAM 71/ 987); 1 d, Baskerville (WAM T47248); 6 d, 3 9, Beacon, ca. 15 km S, Askew Road (WAM T47147); 1 9 with spiderlings, Behn River, Argyle Downs, Ord River area (QM W5060); 1 d, Broome (WAM T47240); 2 9 with eggsac, Cannington (WAM 71/774, 71/ 838); 1 9, Carmel (AM KS86382); 1 d, Chillmoney Road, North, SW Binnu (WAM T47132); 5 d, Chillmoney Road, SW Binnu (WAM T47227); 1 9, 1 juv., Christmas Is- land, 1.5 miles N of South Point (ANIC); 1 9, Christmas Island (QM S61132); 1 d, 3 9, 2 juv.. Cocos Keeling Island (QM S61134); 1 d, 6 9, Coolinup Nature Reserve (WAM T47130, T47141); 1 9, Cottesloe (WAM T53622); 4 d, Dumbleyung Lake North (WAM T47131, T47238); 1 9, Eneabba, AMC mine (WAM T553137); 1 9, Esperance (WAM 71/898); 1 9, Faure Island, Shark Bay (SAM NN14128); 12 d, 40 9, 5 juv., Goon- garrie (WAM T48123, T48166); 1 9, Grass Patch, E of, ‘Sieda’ (WAM T53580); 1 9, Gunyidi, ca. 12 km W (WAM T47142); 1 9, Gutha, 37 miles North (WAM T51547); 2 9, Home Island, Cocos-Keeling Islands (ANIC); 1 d, 1 9, Jarrahdale (WAM T55768-9); 1 9, Kirwan (WAM 71/984); 4 d. Lake Bryde East Nature Reserve, Lake Bryde Rd (WAM T47146); 2 d, Lake Bryde West Nature Re- serve, Lake Bryde Rd (WAM T47139); 1 9, 12 juv.. Lake Cronin (WAM T48124); 1 9, Lake Gruszka (WAM T51548); 1 d. Lake Gulson, 65 km SE of Hyden (WAM T51474); 1 d. Lake Mollerin (WAM T47222); 6 d, 2 9, 1 juv., Lake Ninan Shire reserve (WAM T47143); 12 d, 2 9, 3 juv.. Little Sandy De- sert, 23.1 km ESE of Burranbar Pool (WAM T53420-2); 7 d, 24 9,8 juv.. Little Sandy Desert, 23.3 km ESE of Burranbar Pool (WAM T53417-9); 19,2 juv., Lorna Glen FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 15 Station (WAM T55132); 1 c?, Maitland River (WAM 71/1517); 1 Marangaroo (AM KS86383); 1 $, Mellish Reef (WAM T51413); 1 d, 1 9, Morawa-Perenjori Road (WAM T47144); 1 6, Mortlock Creek, Won= gan Hills (WAM 99/1103); 1 $ with spider- lings, Murchison River (QM S61111); 1 9, Myaree (WAM T53532); 1 d, Nedlands (WAM T53461); 1 d, Noranda (WAM T53535); 1 d, Nugadong West Rd, SW Wub- in (WAM T47136); 1 d, Nullagine (WAM T55307); 1 d, 1 9, Nullewa Lake (WAM T47140); 1 d, Oakajee Nature Reserve (WAM T47221); 1 9, Parry Creek Billabong (WAM T53691); 2 d, 2 9, 3 juv., R.G.C. Mine, 10 km S of Eneabba (WAM T51397- 9); 5 d, 5 9,3 9 with eggsac, Rossmoyne (WAM 71/561-2, 71/740, 71/835-7, 71/867- 70, 71/940, 71/1446, T48122); 1 9, Separa- tion Well (WAM T53510); 1 9, South Lake, near Perth (WAM T53508); 1 d, 2 juv., The Loop, Murchison River (WAM T53662); 1 9, 1 juv., Thirsty Point Waterhole, 1.5 miles E (WAM 71/1447); 1 9, Toolibin Lake (WAM T47133); 1 d, 3 9, Walkaway Nature Reserve (WAM T47135, T47235); 1 9, Wanneroo Lake (WAM 69/2071); 1 9 with eggsac, War- burton Ranges (WAM T53812); 1 d, 1 juv., Warr Well (WAM T51555); 1 9, Weelhamby Lake (WAM T47274); 1 9, Wittenoom Rd near Dempster Rd junction, E Gibson (WAM T47148); 1 9, Yannarie River at North West Coastal Hwy (WAM T53493). NEW CALE- DONIA: 1 9 , no exact location (SAM NN13935). VANUATU: 1 d, Espiritu Santo, Malac Village (SAM NN13936); 2 9, 2 juv., Malekula (AM KS84104). SOLOMON IS- LANDS: 1 9, Vanikoro, Santa Cruz Group (AM KS84108). Diagnosis. — The male and female genitalia of H. crispipes are very similar to those of H. diyari and H. kuyani. However, all three spe- cies can be easily distinguished by their color pattern (Figs. 19, 33, 41). Whereas the median band in H. crispipes is narrower than one- fourth the carapace width, it is one-third the width of the carapace in H. diyari and absent in H, kuyani. Description. — Male: Carapace (Figs. 19, 23): Dorsal line straight in lateral view; dark brown, with light brown narrow median band; distinct light brown submarginal bands with three dark blotches; carapace covered with brown setae in dark areas and white setae in light brown parts and eye region; few black bristles in anterior half of median band; black bristles in head region between PE and pos- terior of PLE; 1 long bristle between AME. Sternum: Yellow-brown; covered with white setae, denser and longer towards margins; few brown bristles. Labium: Brown; front end truncate and white. Chelicerae: Light brown; covered with white setae and few brown bris- tles in basal half; three retromarginal teeth, with the median slightly larger; three promar- ginal teeth, with the median largest. Pedipalp (Figs. 20, 21, 24): Cymbium elongated, tip with 2-6 macrosetae; terminal apophysis sick- le-shaped (Figs. 20, 24). Abdomen: Irregular dark grey; irregular yellow-brown median band; brown lanceolate heart mark with indis- tinct darker edges, that continues into a tri- angular, dark grey pattern in posterior half; covered with white setae and additional brown setae in darker area; venter uniformly yellow- brown and covered with white setae; spinner- ets yellow-brown. Legs: Leg formula IV > I > II > III; all femora, patellae and tibiae brown, dorsally with indistinct grey annula- tions; metatarsi dark brown, metatarsus I with long dense hair-like setae; scopulous setae on all tarsi; spination of leg I (based on SAM NN 13955): Femur: 6 dorsal, 2 apicoprolateral; patella: 1 prolateral, 1 retrolateral; tibia: 3 ventral pairs, 2 prolateral, 2 retrolateral, 1 dor- sal; metatarsus: 2 ventral pairs, 1 apico ventral, 2 prolateral, 2 retrolateral, 1 apicoprolateral, 1 apicoretrolateral. Female: Carapace: As male, submarginal blotches less distinct as the submarginal band is darker. Sternum: coloration light brown, covered with brown bristles of increasing length and density towards margins. Labium: Dark brown, front end truncate and white. Chelicerae: Dark brown, setae and bristles as male; three retromarginal teeth with the me- dian largest, three promarginal teeth, with the median largest. Epigynum (Figs. 25-31): Ven- tral view: inverted T-shaped (Figs. 25, 27-31); dorsal view: round spermathecae, copulatory ducts short and twisted (Fig. 26). Abdomen: As male, pattern less distinct in particular in posterior half; venter yellowish-grey covered with brown setae; spinnerets as male. Legs: Leg formula and coloration as male; spination of leg I (based on SAM NN13970): Femur: 6 dorsal, 2 apicoprolateral; tibia: 3 ventral pairs, 1 (small) prolateral; metatarsus: 2 ventral 16 THE JOURNAL OF ARACHNOLOGY pairs, 1 apicoventral, 1 apicoprolateral, 1 ap- icoretrolateral. Measurements: Male SAM NN 13955 (fe- male SAM NN13970): TL 10.1 (12.6), CL 5.1 (6.0), CW 4.1 (4.5). Eyes: AME 0.23 (0.26), ALE 0.17 (0.17), PME 0.46 (0.43), PLE 0.37 (0.34). Row of eyes: AE 0.92 (1.12), PME 1.09 (1.26), PLE 1.37 (1.60). Sternum (length/ width) 2.4/1.95 (2.55/1.95). Labium (length/ width) 0.63/0.57 (0.80/0.83). AL 5.25 (6.60), AW 2.55 (3.75). Legs: Lengths of segments (femur + patella/tibia + metatarsus + tarsus = total length): Pedipalp 1.64 + 1.95 + — + 1.5 - 5.1, I 5.50 + 5.70 + 4.05 + 2.25 - 16.80, II 4.35 + 5.25 + 3.90 + 2.4 = 15.90, III 3.75 + 4.50 + 4.05 + 2.40 - 14.70, IV 5.10 + 6.15 + 6.00 + 3.00 = 20.25 (Pedipalp 1.95 + 2.25 -f — + 1.50 = 5.70, I 4.05 + 5.25 + 3.00 + 2.10 - 14.40, II 3.90 + 4.95 4 3.00 + 1.95 = 13.80, III 3.60 + 4.20 + 3.15 + 1.95 - 12.90, IV 4.95 + 6.45 + 5.40 -f 2.55 = 19.35). Variation: Males (females) (range, mean ± s.d.): TL 6.0-17.1, 9.5 ± 1.7; n = 55; CL 3.5- 8.7, 4.9 ± 0.8; n = 56; CW 2.6-6.3, 3.7 ± 0.6; n = 56 (TL 8.0-21.0, 13.6 ± 2.7, n - 86; CL 4.1-10.4, 6.0 ± 1.1, n = 86; CW 2.9- 7.4, 4.4 ± 0.9; n = 86). The size variation within H. crispipes is considerable and populations from offshore is- lands and reefs appear to be on average larger than the mainland specimens, a pattern also observed in vertebrates (e.g. Lomolino 1985, Boback 2003). The three dark blotches on the lateral margins of the carapace may not be as distinct as in the specimen illustrated (Fig. 19), and may be absent in some cases. Distribution and habitat preferences. — Hogna crispipes is found on mainland Aus- tralia and offshore islands and reefs in the East and West of Australia (Fig. 32), as well as in New Zealand (Vink 2002) and on several Pa- cific islands (e.g. Tonga, New Caledonia, Va- nuatu, and the Solomon Islands). While un- common in artesian springs, this species is widely distributed across all of the major spring groups within South Australia (Table 1). It is usually found on the edges of the springs and in the ephemeral wet zone that exists beyond the permanent vegetated wet- land. Remarks. — The original description of H. crispipes was based upon male and female syntypes from Bowen and Rockhampton de- Figure 32. — Records of Hogna crispipes (L. Koch 1877) in Australia. posited at the Museum Godeffroy (L. Koch 1877); the precise number of specimens was not stated. While two female syntypes lodged in the BMNH and ZMH were examined dur- ing this study, we could not find any male syntypes in either of these museums or in the ZMB, where the majority of the material from the Museum Godeffroy is currently lodged. These must be considered lost. In addition, L. Koch’s (1877) original description and illus- trations suggest that the male syntype(s) is (are) not conspecific with the females. An in- verted color pattern (light instead of dark me- dian abdominal heart mark) and the structure of the pedipalp suggest that the male specimen illustrated is Venatrix goyderi and not Hogna crispipes. To provide nomenclatural stability for the name H. crispipes, one of the female syntypes is here designated as the lectotype. Hogna crispipes was redescribed and illus- trated by McKay (1979a). A re-examination of the specimens included in McKay’s revi- sion revealed that the majority of the material is not conspecific with the type material of H. crispipes. Only two females from Behn River, Argyle Downs (WA) (QM W5058, W5060; McKay 1979a, fig. m) are conspecific with the female type material of H. crispipes. All other specimens described and illustrated belong to two undescribed Hogna. Although the genital morphology of both males and females of these undescribed species is very similar to H. crispipes, the carapace does not display the typical narrow median and blotched marginal bands, but is very similar to Venatrix arenaris (see Fig. 3). The type material of Lycosa crispipes, Ly~ cosa pulveresparsa L. Koch 1877, Tarentula tongatabuensis Strand 1911 (= senior syno- FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 17 sperma theca Figures 33-39. — Hogna diyari new species: Male: 33. habitus and 34. ventral view of abdomen (ho- lotype from Coongie Lake, SA; SAM NN14115); left pedipalp: 35. ventral and 36. retrolateral view, and 37. apical part of bulb (paratype from Coongie Lake, SA; SAM NN14116). Female (WAM T48035): 38. ventral and 39. dorsal view of epigynum. Scale bar: (33-34) 2.68 mm, (35-36) 0.73 mm, (37) 0.24 mm, (38-39) 0.70 mm. nym of Tarentula tanna Strand 1913 (Ledoux & Halle 1995)), Lycosa waitei Rainbow 1917 and Lycosa rainbowi Roewer 1951 (replace- ment name for Lycosa strenua Rainbow 1920), does not show any differences in so- matic and genitalic characters that warrant sta- tus as different species. Therefore, T. tonga- tabuensis, L. pulvere-sparsa, L. waitei and L. rainbowi are considered junior synonyms of H. crispipes. Recently, Vink (2002) placed H. crispipes (sub G. tongatabuensis) in the genus Geoly- cosa. However, this species does not conform very well to the generic description of Geo- lycosa (e.g. Dondale & Redner 1990). Vink (2002) argued that the genitalia of H. tonga- tabuensis conform more to Geolycosa than Hogna. However, comparison of the genitalic structure of H. crispipes with that of H. ra- diata as illustrated in Fuhn & Niculescu-Bur- lacu (1971) and Miller (1971) showed very good agreement. In addition, Geolycosa is characterized by a sloping dorsal line of the carapace, the absence of light median and sub- marginal bands on the carapace, and the ab- sence of macrosetae at the tip of the male cymbium (Dondale & Redner 1990). None of these characters fit H. crispipes, which has a horizontal dorsal carapace profile, distinct light median and submarginal bands on the carapace, and 2-6 macrosetae on the tip of the cymbium. Therefore, H. crispipes, as well as the closely related H. diyari and H. kuyani are placed in Hogna. Hogna diyari new species Figs. 32-40 Types examined. — Holotype male, Austra- lia, South Australia, Coongie Lake, 27°12'S, 140°10'E, 26-28 October 1995, on shoreline. 18 THE JOURNAL OF ARACHNOLOGY D. Hirst (SAM NN14115). Paratypes: 5 males, 2 females, data as holotype (SAM NN141 11-14, NN14116-8), Other material examined. — AUSTRA- LIA: South Australia: 1 6, Blanche Cup Mound Spring (SAM NN14083); 3 d, 1 $, 14 juv. Clayton Bore, 33 miles N of Marree (WAM 71/573-590); 1 d, Clifton Hills Sta- tion (SAM NN14103); 3 d, 1 juv., Coongie Lake (SAM NN14105-7); 1 d, Coongie, 6.2 km NW (SAM NN14104); 4 d, 6 $, 2 juv.. Coward Springs Railway Bore (SAM NN14085-94); 1 $, Francis Swamp Mound Spring tail (SAM NN14084); 3 d, 2 $, Fred Springs (SAM NN14080-1; WAM T48034- 6); 1 $, Jersey Spring (SAM NN14082); 1 d. Lake Cadibarrawirracanna (SAM NN14102); 2 $, 4 juv.. Lake Hart (SAM NN141 19-20); 3 d , 2 $ , 2 $ with eggsac. Lake Hope chan- nel, 3.9 km S Lake Appadare (SAM NN14095-101); 1 9, Seven Mile Creek, Clif- ton Hills (SAM NN141); 1 $, Stirtons Old Campsite, E edge of Cannuwalkaninna Dune (WAM 73/232). New South Wales: 1 d, 2 9, Broken Hill (SAM NN14079, NN14121~2); 2 d, 1 9, Kinchega National Park (AM KS69252, KS8410). Queensland: 1 d, Cluny Station Billabong (QM S61148); 1 d, Dyne- vor Lakes, E of Thargomindah (QM S61 1 15). Victoria: 1 9, labeled ‘Mcmillan Park, Sale. V’ [possibly Sale, East Gippsland] (MV K8156). Etymology. — The specific name is a noun in apposition honoring the Diyari people, an Aboriginal tribe representing the traditional custodians of parts of the land on which the South Australian artesian springs are found. Diagnosis. — In contrast to the other two darker colored Hogna species of artesian springs, H. crispipes and H. kuyani, the car- apace coloration of //. diyari is light brown with a wide yellow median band that con- stricts anteriorly of the fovea and narrows slightly posteriorly (Eig. 33). Most distin- guishable is a pair of small black spots behind the epigastric furrow and up to eight spots along the lateral border of the yellow venter (Fig. 34). Description. — Male: Carapace (Fig. 33): Brown, with wide yellow-brown median band that constricts anteriorly of fovea and narrows slightly in posterior half; irregular light mar- ginal bands; head region dark brown; carapace covered with white setae, particularly in head region; additional brown setae in dark areas; six brown bristles in median band anteriorly of fovea with the posterior ones in a pair; black bristles in head region between PME and PLE, between PME and below AE. Ster- num: Yellow-brown; densely covered with white setae, denser and longer towards mar- gins. Labium: Light brown, basal half darker; front end truncate and white. Chelicerae: Bas- al half light brown with a dense cover of white setae and fewer brown bristles, apical half dark brown with few brown bristles; three re- tromarginal teeth, with the basal largest; three promarginal teeth, with the median largest. Pedipalp (Pigs. 35-37): Cymbium elongated, tip with two macrosetae; median apophysis with ventral process; terminal apophysis sick- le-shaped, embolus long and slender (Figs. 35, 37). Abdomen: Irregular grey brown; yellow- brown median band; brown lanceolate heart mark with dark grey patchy borders in anterior half, continuing into a triangular, dark grey pattern in posterior half; covered with white setae and additional brown setae in darker ar- eas; few longer, brown bristles; venter yellow- brown with a pair of black spots behind epi- gastric furrow and irregular black spots laterally (Fig. 34); covered with white setae, black setae on black spots; spinnerets yellow- brown, with grey setae towards tips. Legs: Leg formula IV > I > II > III; tarsi and meta- tarsi dark brown, tibiae basally brown and api- cally dark brown, femora brown, femora III and IV with faint grey annulation dorsally; dense scopulous setae on all tarsi and meta- tarsi I and II; dense and hair-like setae dor- sally on tarsi and metatarsi I and IL spination of leg I (based on holotype SAM NN14115): Femur: 6 dorsal, 2 apicoprolateral; patella: 1 prolateral, 1 retrolateral; tibia: 3 ventral pairs, 2 prolateral, 2 retrolateral; metatarsus: 2 ven- tral pairs, 1 apicoventral, 2 prolateral, 2 retro- lateral, 1 apicoventral, 1 apicoretrolateral. Female: Carapace: As male. Sternum: col- oration as male, but fewer and shorter white setae. Labium, chelicerae and their dentition: as male. Epigynum (Eigs. 38, 39): Ventral view: inverted T-shaped (Fig. 38); dorsal view: ovoid spermathecae, copulatory ducts connected posteriorly (Fig. 39). Abdomen: As male, pattern less distinct in particular in pos- terior half; venter and spinnerets as male. Legs: Leg formula IV > I > II > III; color- ation as male; spination of leg I (based on FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 19 Figure 40. — Records of Hogna diyari new spe- cies in Australia. paratype SAM NN14118): Femur: 6 dorsal, 2 apicoprolateral; tibia: 3 ventral pairs, 2 pro- lateral, 2 retrolateral; metatarsus: 2 ventral pairs, 1 apicoventral, 1 apicoprolateral, 1 ap- icoretrolateral. Measurements: Male holotype SAM NN14115 (female paratype SAM NN14118): TL 13.5 (15.5), CL 6.6 (7.1), CW 5.1 (5.1). Eyes: AME 0.34 (0.29), ALE 0.26 (0.23), PME 0.71 (0.69), PLE 0.57 (0.60). Row of eyes: AE 1.34 (1.40), PME 1.66 (1.74), PLE 2.03 (2.4). Sternum (length/width) 2. 8/2.6 (3.0/2.6). Labium (length/width) 0.83/0.83 (1.06/1.06). AL 7.1 (8.7), AW 4.1 (5.6). Legs: Lengths of segments (femur + patella/tibia + metatarsus + tarsus — total length): Pedipalp 2.55 + 2.1 + — + 2.1 = 6.75, I 5.85 + 7.50 + 5.40 + 2.85 - 21.60, II 5.4 + 6.75 + 5.1 + 2.7 - 19.95, III 5.10 + 6.00 + 5.40 + 2.55 = 19.05, IV 6.30 + 7.65 + 7.50 + 3.15 - 24.60 (pedipalp 1.80 + 2.55 + — + 1.80 = 5.15, I 5.25 + 6.75 + 6.90 + 2.25 - 21.15, II 5.10 + 6.15 + 5.10 + 2.10 - 18.45, III 4.50 + 5.40 + 3.90 + 2.10 = 15.09, IV 5.85 + 7.35 + 6.15 + 2.85 - 22.20). Variation: Males (females) (range, mean ± s.d.): TL 9.8-15.1, 12.1 ± 1.4; n = 23; CL 4.1- 7.2, 6.0 ± 0.7; n = 25; CW 3.0-5.3, 4.3 ± 0.5; « - 25 (TL 12.7-17.6, 14.9 ± 1.4, « = 20; CL 5.9-8.1, 6.8 ± 0.7, n - 20; CW 4.2- 6.5, 5.0 ± 0.6; n = 20). Distribution and habitat preferences. — Most specimens of H. diyari have been found near water bodies in the dry interior of South Australia, Queensland and New South Wales. The single female from temperate Victoria may be erroneous, as the label is not entirely conclusive (Fig. 40). While uncommon in ar- tesian springs this species is widely distrib- uted across most of the major spring groups within South Australia (Table 1). It is rarely found around spring vents, preferring the ephemeral wet zone that exists beyond the permanent vegetated wetland. Hogna kuyani new species Figs. 41-47 Types examined. — Holotype male, Austra- lia, South Australia, Coongie Lake, 27°12'S, 140°10'E, 26.-28.X.1995, on shore, D. Hirst (SAM NN14044). Paratypes: 1 male, Coongie Lake, 27°12'S, 140°10'E, iii.1987, pitfall trap, J Reid/Coongie Lake Study (SAM NN 14029); 2 females, Coongie, 4.7 km SE, 27°12'10"S, 140°11'00"E, 14 March 1987, pitfall trap, J Reid/Coongie Lake Study HE (SAM NN14032-3). Other material examined. — AUSTRA- LIA: South Australia: 1 d, Appamurna Wa- terhole (SAM NN14027); 1 d, Clayton Bore, 33 miles N of Maree (WAM T51437); 1 d, Clifton Hill Outstation, 1.1 km E (SAM NN14069); 3 d, 1 ?, 1 juv., Clifton Hills Out- station, 4.8 km NE (SAM NN14062-5); 3 d, Clifton Hills Outstation, 5.4 km E (SAM NN 14066-8); 1 d, Clifton Hills Outstation, 8 km ENE (SAM NN14070); 3 d, Coongie, 1.77 km W (SAM NN14041-3); 1 9, Coon- gie, 11.4 km SE (SAM NN14034); 1 9, Coongie, 5.3 km SE (SAM NN14030); 1 d, Coongie, 7.89 km N (SAM NN 14037); 2 d, Coongie, 9.95 km NNE (SAM NN14035-6); 1 d, Dickinna Hill, 15.5 km SSE (SAM NN 14046); 1 d, Dudley Park Cemetery, Ade- laide (SAM NN14049); 1 9, Emu Bay, 4 km SE, Kangaroo Island (SAM NN14050); 1 d, Greenfields Wetlands, Dry Creeklisbury (SAM NN 14048); 1 9, Lake Palankarinna (SAM NN14026); 1 9, Moomba, 50 km N (SAM NN14025); 2 d, 2 juv., Morris Creek Bore (SAM NN 14021); 1 d, near Roxby Downs adj. Borefield Rd (SAM NN14022); 6 d, 1 9,1 juv.. New Altona Downs, 32.5 km SW (SAM NN14052-8); 1 d, New Altona Downs, 13 km SE (SAM NN14028); 3 d, 1 juv.. New Altona Downs, 36 km SW (SAM NN14059-61); 1 9, Whyalla (MV K8248). New South Wales: 1 d, Arcoola Creek Cross- ing on George Loop Road, Sturt National Park (AM KS84100); 2 d, 5 9, Broken Hill (SAM NN14072-8); 1 d, Connia Creek, 14.8 km S of Olive Downs Homestead, via Jump-Up Loop Road, Sturt NP (AM KS71564); 4 d. 20 THE JOURNAL OF ARACHNOLOGY Figures 41-46. — Hogna kuyani new species: Male: 41. habitus (holotype from Coongie Lake, SA; SAM NN 14044); left pedipalp: 42. ventral and 43. retrolateral view, and 44. apical part of bulb (paratype from 1.77 km W of Coongie, SA; SAM NN14041). Female from 4.8 km SE Coongie (SAM NN14031): 45. ventral and 46. dorsal view of epigynum. Scale bar: (41) 2.84 mm, (42, 43) 0.60 mm, (44) 0.32 mm, (45, 46) 0.43 mm. Mullingar Station, Lower Murray-Darling re- gion (AM KS67040, KS84099); 3 c?, 1 9, Sturt National Park (AM KS71040); 6 S, \ 9, 1 juv., Sturt National Park, 19.2 km S of Fort Grey Homestead, on Camerons Corner Road (AM KS51348); 1 d, Trilby, track to New Chum, 6.4 km from highway junction. Queensland: 1 d, Baryulahl gas well, 38 km S Ballera (SAM NN14360); 1 d, Cluny Sta- tion Billabong (QM S61 149); 1 d, Muncoonie Lakes (QM S61150). Western Australia: 3 d, Camel Lake Nature Reserve (WAM T47229); 2 d, Coolinup Nature Reserve (WAM T47232); 1 d, 2 9, Coyrecup Lake Nature Reserve (WAM T47145); 36 d, 7 9, Dum- bleyung Lake North (WAM T47226, T47237); 1 d. Grass Patch, E of, ‘Sieda‘, ‘10 bagger dam’ (WAM T53586); 1 d, 1 9, Gul- son Lake Nature Reserve (WAM T47218, T48084); 1 d, 3 9, Lake Bryde West Nature Reserve, Lake Bryde Rd (WAM T47138, T47225); 1 d. Lake Daringdella (SAM NN 14051); 1 9, Lake Moore (WAM T47217); 2 9, Midland (WAM 72/248); 2 d, 4 9,5 juv., Molpar (WAM 71/1449-54); 44 d, 14 9, 7 juv., Nugadong West Rd, SW Wubin (WAM T47137, T47228); 1 9, Palla- rup Nature Reserve, Lake Pallarup (WAM T47231); 1 d, 1 9, Reservoir Rd, W Kodinin (WAM T47233-4); 19,1 juv., R.G.C. Mine, 10 km S of Eneabba (WAM T51396); 13 d, 6 9, 34 juv., Taarblin Lake, 10 km SW of Toolibin Lake (WAM T51450); 21 d, 10 9, 22 juv., Taarblin Lake, south-west shore (WAM T48055-7, T48060); 1 d 1 9, Walk- away Nature Reserve (WAM T47134, T47236); 5 d, Wittenoom Hill Nature Re- serve, Wittenoom Rd (WAM T47230); 19,3 juv., Yuinmery (WAM T48125). Etymology. — The specific name is a noun in apposition honoring the Kuyani people, an Aboriginal tribe representing the traditional custodians of parts of the land on which the South Australian artesian springs are found. Diagnosis. — Hogna kuyani can be distin- guished from //. crispipes and H. diyari by its FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 21 uniform, dark reddish-brown carapace color- ation with a dense cover of silver-grey setae and the absence of median or submarginal bands. The median septum of the female epi- gynum is comparatively longer than that of the other two Hogna species (Fig. 45). The genitalic structure of H. kuyani is very similar to that of H. pexa Hickman 1944, which dif- fers in its considerably lighter body colora- tion, in particular of the abdomen. This does not seem to be an artifact of its preservation as Hickman’s (1944) original description con- firms a “yellow [abdomen] with a median lon- gitudinal brown patch in anterior half” in con- trast to the dark grey abdomen of H. kuyani. Description. — Male: Carapace (Fig. 41): Dark reddish-brown, with indistinct darker ra- dial pattern; head region black; carapace cov- ered with a thick layer of silver-grey setae (that rub off easily and may not be present in older specimens), brown bristles in head re- gion; one long bristle between AME, four long bristles below AE. Sternum: Light brown with irregular grey pigmentation; densely cov- ered with white setae, denser and longer to- wards margins. Labium: Dark brown, darkest in basal half; front end truncate and white. Chelicerae: Dark reddish-brown with a dense cover of white setae mainly in basal half; three retromarginal teeth, with the median largest; three promarginal teeth, with the median larg- est. Pedipalp (Figs. 42^44): Cymbium tip with ca. 6-8 macrosetae; median apophysis with ventral process that points basally; terminal apophysis sickle-shaped, embolus with a very thin tip (Figs. 42, 44). Abdomen: Dark grey- brown with an irregular pattern of dark and light spots; densely covered with silver-grey setae and fewer brown bristles in particular in anterior half; venter uniformly light brown and covered with white setae; spinnerets light brown. Legs: Leg formula IV > I > II > III; tarsi and metatarsi brown, tibiae and femora dark brown, femora with indistinct grey an- nulations; dense scopulous setae on all tarsi and metatarsi I and II; metatarsi I with long hair-like setae; femora, and less dense on tib- iae and patellae, dorsally with dense, silver- grey setae, spination of leg I (based on holo- type SAM NN 14044): Femur: 6 dorsal, 2 apicoprolateral; patella: 1 prolateral, 1 retro- lateral; tibia: 3 ventral pairs, 2 dorsal, 2 pro- lateral, 2 retrolateral; metatarsus: 2 ventral Figure 47. — Records of Hogna kuyani new spe- cies in Australia. pairs, 1 apicoventral, 2 prolateral, 2 retrola- teral, 1 apicoventral, 1 apicoretrolateral. Female: Carapace, sternum, labium, and chelicerae: Coloration, setae and cheliceral dentition as male (carapace medially slightly lighter). Epigynum (Figs. 45, 46): Ventral view: Inverted T-shaped with long median septum (Fig. 45); ventral view: circular sper- mathecae, copulatory ducts connected poste- riorly (Fig, 46). Abdomen: Coloration and se- tae as male. Legs: Leg formula IV > I > II > III; coloration as male; all tarsi and meta- tarsi I and II (only in apical half) with dense scopulae; spination of leg I (based on paratype SAM NN14032): Femur: 6 dorsal, 2 apico- prolateral; tibia: 3 ventral pairs, 1 (small) pro- lateral; metatarsus: 2 ventral pairs, 1 apicov- entral, 1 apicoprolateral, 1 apicoretrolateral, 1 (small) prolateral. Measurements: Male holotype SAM NN14044 (female paratype SAM NN14032): TL 11.3 (18.0), CL 6.2 (7.2), CW 4.7 (5.4). Eyes: AME 0.26 (0.29), ALE 0.17 (0.23), PME 0.63 (0.77), PLE 0.49 (0.60). Row of eyes: AE 1.17 (1.32), PME 1.63 (1.95), PLE 1.95 (2.35). Sternum (length/width) 2. 9/2.0 (3. 2/2. 7). Labium (length/width) 0.74/0.74 (0.97/0.97). AL 5.3 (9.8), AW 3.3 (6.8). Legs: Lengths of segments (femur + patella/tibia + metatarsus + tarsus = total length): Pedipalp 2.25 + 2.25 + — + 1.95 - 6.45, I 5.25 + 7.35 + 5.25 + 2.70 = 20.55, II 4.95 + 6.30 + 4.50 + 2.55 = 18.30, III 4.50 + 5.55 + 4.80 + 2.40 - 17.25, IV 5.85 + 7.35 T 7.50 + 2.85 = 23.55 (Pedipalp 2.85 + 2.85 + — + 1.80 = 7.50, I 5.10 + 7.20 + 3.90 + 1.95 - 18.15, II 5.10 + 6.30 + 4.05 + 1.95 = 17.40, III 4.50 + 5.55 + 4.35 + 1.95 - 16.35, IV 5.70 + 7.95 + 7.50 + 2.55 - 23.70). 22 THE JOURNAL OF ARACHNOLOGY Variation: Males (females) (range, mean ± s.d.): TL 8.5-22.3, 15.5 ± 3.3; n = 28; CL 5.1-10.5, 8.5 ± 1.6; n = 28; CW 3.6-8.3, 6.4 ± 1.3; n - 28 (TL 12.0-19.7, 16.2 ± 2.2, n = 14; CL 5.9-9.9, 7.8 ± 1.3, n = 14; CW 4.4-7.2, 5.8 ± 0.9; n = 14). As in H. crispipes, the size variation in H. kuyani is remarkable. For example, males range from 8.5-22.3 mm body length, mean- ing that the largest specimens are nearly three times the size of their smallest conspecifics. Distribution and habitat preferences. — This species is found widely across New South Wales, South Australia, Western Aus- tralia, and Queensland (Fig. 47). It is present in low numbers across a wide range of arte- sian springs in South Australia, but is most common in the springs around Hermit Hill (Table 1). Hogna kuyani can be found in un- saturated areas of wetland vegetation, where it makes shallow, wide burrows that are con- cealed by sheets of web covered in mud and litter. Unknown subfamily Genus Tetralycosa Roewer 1960 Tetralycosa Roewer 1960: 949 (name first listed as nomen nudum in Roewer 1955: 296). Type species. — Lycosa meracula Simon 1909, by monotypy (Roewer 1960). Diagnosis. — Males of Tetralycosa differ from all other lycosid genera by the combi- nation of the following characters: reduced palea with well developed, thin embolus; con- ductor forms a shaft for the resting embolus; terminal apophysis absent; tegulum deeply di- vided; medium apophysis originating apically on tegulum and hook-shaped, opposing an ap- icomedially directed pointy protrusion on the retrolateral section of the tegulum. Females: epigynum with a wide median septum, some- times partially hidden behind circular or oval sclerotized atrium. Generic description. — Small to large wolf spiders (TL 4.5-22.0 mm). Males smaller than females. Carapace longer than wide, dorsal profile more or less straight in lateral view in smaller species {T. arahanae, T. oraria) (Fig. 53), but with an elevated head region and downward slope towards posterior end in larg- er species {T. alteripa, T. eyrei). Carapace col- oration variable from a light yellowish brown {T. arahanae) to very dark brown {T. eyrei), without or with only an indistinct light median band. Abdomen coloration variable, generally with dark heart mark. AME larger than ALE, row of AE straight or slightly procurved (Figs. 52). Chelicerae generally with three promar- ginal and three retromarginal teeth, but 2-4 teeth on individual chelicerae possible on both margins. Leg formula IV > I > II > III or IV > II > I > III {T. alteripa, T. eyrei). Tegulum deeply divided longitudinally in retrolateral half. Median apophysis located apically at tegulum and forming a ventrally directed hook that opposes an apicomedially directed pointy protrusion on the retrolateral section of the tegulum. Median apophysis with a basal lobe. Palea reduced. Embolus originating prolaterally on and curving ven- trally around palea, long and slim. Ventrally directed lobe at the base of the embolus. Ter- minal apophysis well developed and forming a sclerotized shaft in which the embolus rests. Cymbium dorsally with dense, scopulous se- tae in apical half and without or only a few macrosetae on tip. Epigynum variable with a wide median septum sometimes only partially visible behind the sclerotized margins of the epigynum which only leave a round or oval atrium. Small round or oval spermathecae. Copulatory ducts short and twisted. Remarks. — The monotypic genus Tetraly- cosa, with the type species Lycosa meracula Simon 1909, was initially listed by Roewer (1955). Subsequently, Roewer (1960) provid- ed a diagnosis for this genus, characterized by the number of retromarginal cheliceral teeth and the arrangement of the eyes. McKay (1973) listed the species in Lycorma Simon 1885, following Guy (1966) who considered Tetralycosa a subgenus of Lycorma. Subse- quently, McKay (1979c) synonymized Tetra- lycosa with Lycosa. This decision was based on the examination of two juvenile syntypes of L. meracula. The syntype series, however, also contains a mature male and a recent in- vestigation of this specimen lodged at the MHNP revealed Lycosa meracula to be a ju- nior synonym of Trochosa oraria (L. Koch 1876). Due to the unique pedipalp morphol- ogy of T. oraria, Tetralycosa is here reinstated as the valid genus and monophyletic group of Australian wolf spiders, most of which appear to favor saline conditions near salt lakes, mound springs or sea shores. Roewer (1960) based the generic descrip- tion of Tetralycosa mainly on somatic char- FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 23 Figures 48—56. — Tetralycosa arabanae new species: Holotype male from Jersey Spring, SA (SAM NN13871): 48. habitus; 49. left pedipalp, ventral; 50. left pedipalp, retrolateral; 51. male, apical part of bulb (WAM T47295, from Morris Creek Bore, SA); 52. eye arrangement; 53. lateral profile of crapace. Females: ventral view of epigynum: 54. WAM T47299; 55. WAM T47297 (from Gosse Springs, SA); 56. dorsal view of epigynum (WAM T47296, from Morris Creek Bore, SA). Scale bar: (48) 1.42 mm, (49-50) 0.35 mm (51) 0.13 mm, (52) 1.09 mm, (53) 1.49 mm, (54-56) 0.40 m. acters, in particular the number of retromar- ginal teeth on the chelicerae and the arrangement of the eyes. Here, we revalidate this genus based on the unique morphology of the male pedipalp. The subfamilial placement of Tetralycosa is unclear. The genus appears to have some af- finities with the subfamily Pardosinae, as the conductor is “shaftlike, lying transversely along basal magin of palea” (Dondale 1986) and a “thick welLsclerotized basal part of pa- lea concealed by tegulum” (Zyuzin 1993). However, a preliminary molecular analysis of a dataset from the 12S rRNA gene subunit, that included T. oraria, was not conclusive in regard to the subfamilial placement of this ge- nus (Vink et al. 2002). Parsimony analysis re- sulted in the placement of T oraria, together with the presumably lycosine species Arctosa leopardus (Sundevall 1833), as a sister-group to traditional lycosine {Alopecosa, Lycosa, Trochosa, Varacosa, Venatrix) and pardosine (Pardosa) genera combined. In contrast, a strict consensus tree of six maximum likeli- hood trees places T. oraria basally, as sister- taxon to all other lycosid species (Vink et al. 2002). Tetralycosa also shows some affinity with Artoria. The laminar lobe at the base of the embolus in Tetralycosa (Figs. 51, 58) may be homologous to the basoembolic apophysis in Artoria. In addition, the shaft-like conduc- tor of Tetralycosa is situated similarly as the terminal apophysis (sensu Framenau 2002) in Artoria. Four species are here included in Tetraly- cosa: T. alteripa (McKay 1976) new combi- nation; T. arabanae new species; T. eyrei (Hickman 1944) new combination, and T. or- aria (L. Koch 1876) new combination. All four species appear to be halophilic species. 24 THE JOURNAL OF ARACHNOLOGY Figure 57. — Records of Tetralycosa arabanae new species in Australia. Two representatives, T. alteripa and T. eyrei, are exclusively found on the surface of salt lakes in the dry interior of Australia (McKay 1976; Hudson 1996, 1997; Hudson & Adams 1996) and T. oraria is typically found on the foreshore and in sand dunes of ocean beaches in southern Australia and Tasmania (McKay 1979c). A full revision of this genus will be the subject of a forthcoming publication. Tetralycosa arabanae new species Figs. 48-57 Types examined. — Holotype male, Austra- lia, South Australia: Jersey Spring, 29°20'S 136°45'E, 18.vii.l996, D. Niejalke (SAM NN 13871). Paratypes: 1 male, 2 juveniles, same location as holotype, 18.vii.l996, D. Niejalke (SAM NN13872); 7 males, 2 females 1 female with 41 spiderlings, same location, 12.xi.l997, K-J Lamb (SAM NN13887-96). Other material examined. — AUSTRA- LIA: South Australia: 1 6, Blanche Cup Mound Springs (SAM NN13884); 1 d, But- tercup Mound Spring (SAM NN13885); 1 $, Coongie Lake (SAM NN13869); 4 6, 1 9, Elizabeth Springs (North A) (SAM NN13878-82); 1 9, Elizabeth Springs (North B) (SAM NN13883); 19,1 juv., Elizabeth Springs Bore (SAM NN 13870); 2 S, 4 juv., Francis Swamp mound spring (SAM NN 13876-7); 1 9 with 67 spiderlings, Gosse East Spring (SAM NN 13886); 2 9, Gosse Springs (WAM T47297); 19,1 juv.. Hermit Hill Springs (SAM NN13873); 20 d, 10 9, 1 juv., Horse East Spring (SAM NN 13897- 916); 2 9, Horse Springs (WAM T47299); 1 9, Lake Frome (SAM NN 13867); 1 9, Lake Hart (SAM NN 13933); 1 9, 1 juv., Lake Hope Channel, 3.9 km S Lake Appadare (SAM NN13868); 2 d, 1 9, 1 juv., Mc- Lachlan Springs (WAM T47298); 1 d, 1 9, Morris Creek Bore (WAM T47295-6); 1 6, 4 juv.. Old Linnis Spring (SAM NN 13874); 1 d. Smith Spring (SAM NN13875, NN13929- 32); 5 6,2 9,5 juv., Tregolana Salt Lake (SAM NNl 3862-4). Etymology. — The specific name is a noun in apposition honoring the Arabana people, an Aboriginal tribe representing the traditional custodians of parts of the land where the South Australian artesian springs are found. Diagnosis. — Tetralycosa arabanae is very similar to T. oraria in particular in regard to male pedipalp morphology. However, the low- er tip of the conductor of the male pedipalp of r. arabanae has a triangular process point- ing apically (Fig. 51). This process is absent in T. oraria (Fig, 58). Female genitalia of both species are easily distinguished as the trian- gular epigynum of T. arabanae is approxi- mately as wide as long (Figs. 54, 55), whereas the ovoid epigynum of T. oraria is much wid- er than long (McKay, 1979c, figs, lb, d, f). Description. — Male: Carapace (Figs. 48, 53): Dorsal line straight in lateral view; light yellow-brown, sometimes with indistinct light brown radial pattern; eye field very dark brown; carapace covered with mainly white setae, few black setae posterior of fovea; few black bristles in eye field; four long brown bristles below AE, one long bristle between AML. Sternum: Yellow; white setae and fewer brown bristles both denser and longer towards margins. Labium: Brown; front end truncate and white. Chelicerae: Light brown; white se- tae basally and laterally, black setae apically near fangs; two retromarginal teeth of similar size; three promarginal teeth, with the median largest. Pedipalp (Figs. 49-51): Median apophysis a broad, ventrally directed hook and with basal lobe; embolus with a basal bulge and resting in a shaft formed by the conduc- tor; lower tip of conductor triangular (Fig. 51); cymbium dorsally with scopulous setae in apical half, and few apical macrosetae. Ab- domen: Olive-yellow; faint brownish heart mark in anterior half; three to four pairs of yellow spots, of which the anterior and pos- terior pair are largest; covered with white se- tae and few longer, brown bristles; venter yel- low; setae as dorsally, but brown bristles lighter and shorter; spinnerets yellow. Legs: Leg formula IV > I > II > III; light yellow- brown, with faint annulations centrally and in FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 25 apical half of femora, in basal half of patella, and basally and centrally on tibia; tarsus and metatarsus of legs I and II darker (brown); spination of leg I (based on holotype SAM NN 13871): Femur: 4 dorsal, 1 apicoprolateral, 1 apicoretrolateral; patella: 1 prolateral, 1 re- trolateral; tibia: 3 ventral pairs, 2 dorsal, 2 prolateral, 2 retrolateral; metatarsus: 3 ventral pairs, 3 prolateral, 3 retrolateral; 1 apicovee- traL Female: Carapace, sternum and labium: as male. Chelicerae: Dark brown, generally much darker than in males; setae and bristles as male; cheliceral dentition as male. Epigyn- um (Figs. 54-56): Ventral view: triangular with convex posterior margin (Figs. 54, 55); dorsal view: small oval spermathecae, copu- latory ducts directed anteromedially, small spermathecal organs (Fig. 56). Abdomen: As male. Legs: Leg formula IV > I > II > III; coloration as male, however tarsus and meta- tarsus of leg I and II not darker; spination of leg I (based on female SAM NN1389, reduced in comparison to males): Femur: 4 dorsal, 1 apicoprolateral; tibia: 1 apico ventral pair, 1 prolateral; metatarsus: 2 ventral pairs, 1 api- coventral. Measurements: male holotype SAM NN13871 (female SAM NN13894): TL 7.3 (7.6), CL 4.0 (3.7), CW 2.8 (2.7). Eyes: AME 0.2 (0.2), ALE 0.14 (0.16), PME 0.28 (0.32), RLE 0.26 (0.16). Row of eyes: AE 0.82 (0.88), PME 0.76 (0.80), PLE 1.10 (1.20). Sternum (length/width) 2. 0/1. 5 (1.6/1. 4). Labium (length/width) 0.51/0.52 (0.54/0.66). AL 3.1 (3.4), AW 2.3 (2.6). Legs: Lengths of seg- ments (femur + patella/tibia + metatarsus + tarsus = total length): Pedipalp 1.50 + 1.25 + -- + 1.20 = 3.95, I 3.05 + 3.90 + 2.90 + I. 55 = 11.40, II 3.0 + 3.65 + 3.0 + 1.5 = II. 15, III 2.85 + 3.25 + 3.05 + 1.4 - 10.55, IV 3.55 + 4.20 + 4.0 + 1.8 = 13.55 (PedL palp 1.25 + 1.25 + — + 1.0 = 3.5, I 2.35 + 3.15 + 1.95 + 1.1 = 8.65, II 2.30 + 2.90 + 1.9 + 1.05 = 8.15, III 2.25 + 2.25 + 2.05 + 1.05 - 7.9, IV 2.9 + 3.4 + 3.1 + 1.30 - 10.7). Variation: males (females) (range, mean ± s.d.): TL 4.8-7.7, 6.3 ± 0.8; n = 23; CL 2.5- 4.2, 3.3 ± 0.5; n - 24; CW 2.0-3.2, 2.5 ± 0.3; « - 24 (TL 6.6-11.1, 7.9 ± 1.2, n = 17; CL 3. 1-5.8, 3.8 ± 0.6, n = 19; CW 2.2-3.6, 2.7 ± 0.4; n - 19). Distribution and habitat preferences. — Figures 58. — Tetralycosa oraria (L. Koch 1876): Male from Australind, WA (WAM 71/360), apical part of bulb. Scale bar: 0.21 mm. This species is restricted to arid South Aus- tralia (Fig. 57). It is found in the southern and eastern springs from Jersey Springs in the west to Mulligan Springs in the east (Table 1). Tetralycosa arabanae is largely restricted to the lower parts of the spring tail and the ephemeral wet regions beyond the permanent vegetated wetland. It has also been found near semi-permanent saline waterholes near Hermit Hill Springs. NON-ARTESIAN SPRING LYCOSIDAE OF THE GENUS TETRALYCOSA The following species are not part of the artesian spring fauna of South Australia, but are transferred to the Tetralycosa as they show the unique pedipalp morphology characteristic for this genus. Tetralycosa alteripa (McKay 1976) new combination Lycosa alteripa McKay 1976: 418-420, figs. 2, 2a- e; Brignoli 1983: 450; McKay 1985: 74. Remarks. — -Tetralycosa alteripa shows the typical pedipalp and epigynum structure of Tetralycosa (McKay 1976, 418-420, figs, 2, 2a-e; holotype male (WAM 70/41) and para- type males and females (WAM 70/42-46, 74/ 501) examined by VWF) and is therefore transferred from the northern hemisphere ge- nus Lycosa to Tetralycosa. This species is typ- ically found on the surface of salt lakes in South Australia and Western Australia (Mc- Kay 1976; Hudson 1997). An allozyme study suggests the existence of an undescribed, cryptic sister-species of T alteripa in Western Australia (Hudson & Adams 1996). 26 THE JOURNAL OF ARACHNOLOGY Tetralycosa eyrei (Hickman 1944) new combination Pardosa eyrei Hickman 1944: 24,25, plate 1, figs. 11-13; Roewer 1955: 185; McKay 1973: 378. Lycosa eyrei (Hickman 1944): McKay 1985: 76; Platnick 1989: 370. Remarks* — The pedipalp and epigynum structure of T. eyri is similar to that of T. al- teripa (Hickman 1944: 24, 25, plate 1, figs. 11-13; holotype male (AM KS5738) and con- specific males and females (SAM NN13809- 15, NN17384-5; MV K8126, K 8183, ex- amined by VWF). As in T. alteripa, this species is typically found on the surface of salt lakes in South Australia and Victoria (Hudson 1996; Hudson & Adams 1996), al- though, allozyme data suggest the co-occur- ance of two cryptic species within T. eyrei (Hudson & Adams 1996). Tetralycosa eyrei has a sympatric distribution with the salt-lake dwelling scorpion Australobuthus xerolim- niorum Locket 1990 (Hudson 1997). Tetralycosa oraria (L. Koch 1876) new combination Fig. 58 Lycosa oraria L. Koch 1876: 883-886, plate 76, figs. 2, 2a, 3, 3a; Simon 1909: 188; Rainbow 1911: 270; Bonnet 1957: 2656. Lycosa candicans L. Koch 1877: 888-890, plate 76, figs. 5, 5a, 6, 6a, b; Rainbow 1911: 266; Hickman 1950: 5; Bonnet 1957: 2637. NEW SYNONY- MY. Lycosa sibyllina Simon 1909: 188, 189, fig. 7; Rainbow 1911: 272; Bonnet 1957: 2664; McKay 1973: 379; Moritz 1992: 325. Synonymized by McKay 1979c: 279. Lycosa meracula Simon 1909: 190, 191; Rainbow 1911: 270; McKay 1985: 80; Platnick 1989: 372; Moritz 1992: 320. NEW SYNONYMY not Lycosa meracula Simon, sensu McKay 1979c: 264, figs. 9a-k (misidentification; not L. mera- cula but an undescribed species). Crocodilosa oraria (L. Koch 1877): Roewer 1955; 238. Tetralycosa meracula (Simon 1909): Roewer 1955: 296; Roewer 1960: 949; Rack 1961: 38. Hogna sibyllina (Simon 1909): Roewer 1955: 253. Trochosula candicans (L. Koch 1877): Roewer 1955: 304. Trochosomma oraria (L. Koch 1877); Roewer 1960: 847; Roewer 1961: 14. Ocyale oraria (L. Koch 1877); McKay 1973: 380. Lycorma meracula (Simon 1877): McKay 1973: 380. Trochosa candicans (L. Koch 1877): McKay 1973: 381; McKay 1979c: 293-294, fig. 4e; McKay 1985: 85; Platnick 1989: 390. Trochosa oraria (L. Koch 1877): McKay 1979c: 279-282, figs, la-h; McKay 1985: 86; Platnick 1989: 391; Platnick 1993: 510. Remarks.' — -The male pedipalp of T. oraria is very similar to that of T. arabanae. It can mainly be distinguished by the lower tip of the conductor, which has a triangular protru- sion in T. arabanae (Figs. 51), but not so in T. oraria (Figs. 58). The wide, oval median septum of the epigynum of T, oraria (McKay 1979c: 279-282, figs, lb, d, f) conforms to the general pattern of Tetralycosa. Simon (1909) described Lycosa meracula based on one male and some immature spiders from (p. 191) “Stat. 5, Denham, ad litus in detritus; Stat. 65, Albany” collected during the ‘Hamburger Siidwest-Australische Expe- dition 1905' (Michaelsen & Hartmeyer 1907; Simon 1909). Three immatures from Denham are deposited in Hamburg (ZMH, Rack (1961)-catalogue 466), Berlin (ZMB 11085) and Perth (WAM 11/4303) (VWF, all exam- ined). Therefore, the adult male lodged in Par- is (MHNP 24964, labeled "'Lycosa meracula E.S., Austr. occid. (Michaelsen)”, VWF, ex- amined) without accurate locality data must be regarded as the syntype from Albany. This adult male is coespecific with T. oraria L. Koch 1876, as there is no difference in so- matic and genitalic characters between these species. Consequently, L. meracula, the type species of Tetralycosa, is considered a junior synonym of T. oraria. This also agrees with the type localities of both species, as T. oraria was described from King George Sound, the harbor bay of Albany (L. Koch 1876). Not being aware of the existence of the ma- ture male syntype at the MHNP, McKay (1979a) redescribed T meracula based on adult material collected near the type locality, Denham, of the immature specimens which are lodged in Berlin and Perth. However, the species he illustrated is not coespecific with the adult male syntype of T. meracula from Albany, and therefore his treatment of this species must be regarded as erroneous. He also stated that (p. 267) “The record of this species from Albany ] is erroneous, as the southern limit of this species appears to be just north of the Murchison River. [. . . ] Sta- tion 65 ‘Albany’ refers to station 65 Denham FRAMENAU ET AL.-— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 27 ] and not station 165 Albany,” Although Simon (1909) ¥/as incorrect citing ‘‘Stat. 5” for Denham (actually Stat, 65; Stat, 5 refers to a marine surface collection near Denham) and “Stat. 65” for Albany (actually Stat. 165), McKay’s (1979a) redesignation of the type lo- cality for the male syetype from Albany to Denham appears to be incorrect. Simon’s (1909) “Stat. 65, Albany” is most likely a transcription error of “Stat. 165, Albany”. It is likely that the immature syntypes of L. meracula are not, as McKay (1979a) suspect- ed, coespecific with the adult male from Al- bany, as r, oraria has not been reported from as far north as Denham. However, the adult male remains the name-bearing specimen as it was described earlier (Simon 1909, pp. 190, 191) than the juveniles (p. 191) (recommen- dation 69A.10, ICZN 1999). More important- ly, only the male syetype of L. meracula al- lows for an accurate identification of this species. The original illustrations of T. candicans (L. Koch 1877) with the hook-shaped mediae apophysis of the male pedipalp and the oval epigynum strongly suggest a synonymy with T. oraria. McKay (1979c) also stressed the similarity of T. oraria and T. candicans but did not synonymize both species, as he was not able to investigate more than one female specimen (listed as T. candicans in Hickman (1950)) of this presumably eastern Australian species. Tetralycosa oraria was then only known from Western Australia. A comparison of type material of both species is not possible as no syntypes of L. candicans could be lo- cated in the Naturhistorisches Museum, Vi- enna (J. Gruber, personal communication) or ‘Bradleys Collection’ (whereabouts of this collection unknown) where, according to L. Koch (1877), they should be housed. Our re- cent investigations uncovered a large number of recently collected T. oraria in the AM, MV, SAM, TMAG, and QVMAG from eastern Australian states including Tasmania which leave no doubt that T. candicans and T. oraria are conspecific. Therefore, T. candicans is considered a junior synonym of T. oraria. Tetralycosa oraria is mainly found on beaches and sand dunes along the southern coast of mainland Australia (Vic, SA, NSW, WA) and Tasmania. Figures 59--61. — Artoria howquaensis Framenau 2002: Male from Howqua River, Vic (MV K7467): 59. left pedipalp, ventral; 60. eye arrangement. Fe- male from from Howqua River, Vic (MV K7468): 61. dorsal view of epigynum Scale bar: (59) 0.30 mm, (60) 0.82 mm, (61) 0.30 mm. Genus Artoria Thorell 1877 Artoria Thorell 1877: 531. Artoriella Roewer 1960: 563 (name listed as nomen nudurn in Roewer 1955: 233). Trabaeola Roewer 1960: 582. Remarks* — The genus Artoria Thorell was established with the description of the male of A. parvula Thorell 1877 from Sulawesi. Fra- menau (2002) reviewed the genus including the description of seven new species from floodplain habitats in Victoria. An alpine Ar- toria was recently recorded from Mt. Kosci- usko (NSW) (Framenau 2004). The genus ap- pears to be widespread in southeast Asia and the Australasian region with probably more than 50 undescribed species in Australia alone (Framenau 2002). Vink (2002) recently re- corded three new species from New Zealand. The palpal morphology of Artoria is unique within the Lycosidae, and a preliminary mo- lecular analysis suggests that this genus forms a monophyletic clade with Anoteropsis Koch, 1877 and Notocosa Vink 2002 (Vink et al. 2002). Artoria does not fit any of the current subfamilies defined by Dondale (1986), Al- derweireldt and Jocque (1993) or Zyuzin (1985, 1993). 28 THE JOURNAL OF ARACHNOLOGY Figure 62. — Records of Artoria howquaensis Framenaii 2002 in Australia. Artoria howquaensis Framenau 2002 Figs. 59-61 Artoria howquaensis Framenau 2002; 217, 218, figs 9a-g, 10. Diagnosis. — This is the smallest (TL 3.5-6 mm) and one of the most common wolf spi- ders at the South Australian artesian springs. It can easily be distinguished by its body col- oration. The carapace is black, with distinct white marginal bands caused by a dense layer of white setae. The abdomen is dark grey to black, an indistinct lighter heart mark may be visible. The patella and tibia of the first leg of males are bright yellow. The tibia and basal half of the cymbium of the male pedipalp bear a dense cover of white setae. The median apophysis of the male pedipalp bears an apical triangular lobe (Fig. 59). The female epigyn- um is a simple, laterally sclerotized posterior atrium (Fig. 61). Distribution and habitat preferences. — In addition to being widespread and common within the South Australian artesian springs (Table 1), this species has been found in low- land floodplains of rivers and other moist hab- itats in Victoria and South Australia, including Kangaroo Island (Framenau 2002; D. Hirst, pers. comm.) (Fig. 62). Within the artesian springs, Artoria howquaensis prefers C. lae- vigatus wetlands and can be found foraging on top of dense mats of vegetation. It is active mainly during the day and retreats to a silk shelter at the base of C. iaevigatus during the night. Artoria victoriensis new species Figs. 63-70 Types examined. — Holotype male Austra- lia, Victoria: Melbourne, 37°49'S, 144°58'E, 8 October 1956, A Neboiss (MV K7742). Par- atype female, Kilsyth, 37°48'S 145°19'E, 11 October 1981, on fence, ME Roberts (MV K7741). Other material examined. — AUSTRA- LIA: South Australia: 1 d, 1 $, 1 ? with eggsac, 4 juv., Adelaide foothills nr. Waite Campus (SAM NN131 13-15); 1 $, 1 juv., Adelaide Parklands between Adelaide Zoo and Hackney Bridge (SAM NN13132); 4 6, 4 $, Belair (SAM NN13094-101); 1 $, Bo- livar (SAM NN13139); 1 6, 3 9, Bridgewa- ter, Mt Lofty Ranges (SAM NN 13086-9); 1 9, Calpatanna Waterhole Conservation Park, Wedina Well (SAM NN13163); 2 6, 1 9, Cape St Albans Lighthouse, 5.25 km WNW, Kangaroo Island (SAM NN13068-70); 1 9, Carrieton Township, E side (SAM NN13161); 8 (3, 4 9, 1 juv., Francis Swamp, mound spring near Leonard Bore (SAM NN1236- 47); 1 9, Ceres, Burner (SAM NN13042); 1 (3, Charleston Conservation Park (SAM NN13112); 2 (3, 4 9, 2 juv., Chowilla, Nil Nil Bend (SAM NN617-9, NN13157-9); 2 c3, 7 9, Cleland Conservation Park, cnr Wine Shanty & Pit Box tracks (SAM NN13102- 11); 3 9, Conmurra, 4.4 km N telephone ex- change (SAM NN13036-8); 1 (3, Coorong area, 5 km ENE Tilley Swamp telephone ex- change (SAM NN 13047); 3 c3, Coorong area, 1.5 km ENE Tilley Swamp telephone ex- change (SAM NNl 3048-50); 3 (3, Coroman- del Valley (SAM NN1309-2); 1 (3, Cox Scrub Conservation Park, 2 km S South tip (SAM NN13083); 1 9, Custon, 1.2 km S (SAM NNl 3056); 1 9, Flinders Ranges National Park, 1.7 km SW Wilpena Chalet (SAM NNl 3 160); 2 9, Frances, 1.1 km NNE (SAM NNl 3054-5); 1 9 with eggsac, Greenfield Wetlands, Salisbury (SAM NN13140); 1 (3, Inglewood Homestead, 1.3 km SSE (SAM NNl 3065); 1 9, King Fisher Spring, Dalhou- sie Springs (SAM); 1 9 with eggsac, Klemzig (SAM NN13131); 1 9, Kongorong Forest Re- serve, 16.4 km WNN Headquarter (SAM NNl 3026); 1 9, Kongorong Forest Reserve, 17.3 km WNN Headquarter (SAM NN13025); 14 9, 1 juv., Kongorong, 14.5 km W tele- phone exchange (SAM NN13011-24); 1 9, Lake Malata South (SAM NN13162); 2 c3, 4 9, Largs North (SAM NN 13 133-8); 1 c3, Lu- cindale (SAM NNl 3043); 1 (3, 1 9, 1 juv., Magill CAE, Adelaide (SAM NN13126-7); 1 c3, 1 9, Magrath Flat, 9 km NW (SAM FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 29 Figures 63-69. — Artoria victoriensis new species: Male holotype from Melbourne, Vic (MV K7742): 63. habitus; 64. eye arrangement; 65. left pedipalp, ventral; 66. left pedipalp, retrolateral; 67. apical part of bulb (MV K7774, from near Baxter, Vic). Female: 68. ventral view of epigynum (MV 7741, from Kilsyth, Vic); 69. dorsal view of epigynum (MV 7740, from Yarra Valley Park, Vic). Scale bar: (63) 1.18 mm, (64) 1.01 mm, (65, 66) 0.38 mm, (67) 0.27 mm, (68, 69) 0.33 mm. NN13 148-9); 1 cJ, Melville Gully, Belair Na- tional Park (SAM NN13093); 1 $, Millewa Road, NE Hahndorf (SAM NN13085); 2 $, 1 juv., Millicent Airport, 11 km SW (SAM NN13027-8); 1 9, Millicent Airport, 14 km SW (SAM NN13029); 3 9, Minecrow trip point, NNE (SAM NN13044-6); 4 9,29 with spiderlings, Mitcham (SAM NN13116- 21); 12 9, Monarto Zoo (SAM NN14200- 11); 1 9, Mt Benson telephone exchange, 8.7 km ENE (SAM NN13039); 1 d, 2 juv., Mt Compass, 21 km ESE (SAM NN13080); 1 9, Mt Compass, 22 km ESE (SAM NN13082); 1 d, Mt Compass, 9 km E (SAM NN13081); 3 d, Mt Rough, 12.1 km NNE (SAM NN 1305 1-3); 1 d, Muston, Kangaroo Island (SAM NN13067); 1 d, 1 juv., Muston, Kan- garoo Island, in midchannel of Pelican La- goon (SAM NN 13066); 2 9, Myponga, Mt Lofty Ranges (SAM NN13154-5); 1 9, Nap- pyalla (SAM NN13146); 1 d, 1 juv., nr. Pyap (SAM NN13151); 19,1 juv., Old Kings Sta- tion, 2 km W (SAM NN13147); 1 d, Parawa, 2 km WNW (SAM NN13078-9); 5 c3, 1 9, Parawa, 5 km ENE (SAM NN13072-77); 1 9 , Penola Forest Reserve, 19.7 km NW Head- quarter (SAM NN13030); 5 9,1 juv., Penola Forest Reserve, 5 km NNE Headquarters (SAM NN13031-5); 1 9, Point Sturt, Lake Alexandrina (SAM NN13150); 1 d, Poogin- agorie, 3.7 km NE (SAM NN13057); 2 9,1 juv., Pyap, 2 km S (SAM NN13152-3); 2 9, Robe substation, 5.3 km S (SAM NN13040- 1); 1 d, Scott Creek Conservation Park, MacKreath Creek (SAM NN13084); 5 d, Scott Creek, S of Morgan, near River Murray (SAM NN13141-5); 1 9, Tarkeerip, 6.1 km NE (SAM NN13064); 3 c5, 3 9, Teatrick, 0.4 km WSW (SAM NN13058-63); 1 d, Tindale East Cave (AM KS52385); 1 9 with eggsac, Torrens Park, Magill CAE (SAM NN13122); 1 9, Torrens Park (SAM NN13123); 19,1 9 with eggsac, Tusmore (SAM NN 13 124-5); 1 d, Victor Harbor (SAM NN13071); 3 S, Windsor Gardens (SAM NN13128-30). New South Wales: 30 d, 28 9, 1 juv., Coleambally 30 THE JOURNAL OF ARACHNOLOGY irrigation area (AM KS58090, KS58127, KS58164, KS58183, KS58235, KS58311, KS67076, KS67152, KS67342, KS67348, KS67354, KS67412, KS67506, KS67674, KS67678, KS67684, KS68649, KS68654, KS68662, KS67764, KS71271); 2 c?, Crown residency, corner of New England Highway and Old Tamworth Road (AM KS 82846, KS82854); 16,2 9, Gilgandra, 39 km NNW, turnoff to Warrumbungles National Park (AM KS76597~8, KS76600); 1 9, Gin Gin, 2.5 km NW, on road to Riverview Station (AM KS76601); 1 6, Kwiambal National Park, East side, 150m South of Road (AM KS82858); 1 6, 1 9, McIntyre River, 2.8 km S of Boggabilla on Bruxner Highway (AM KS76603, KS76605); 1 9, More^e (AM KS32588); 16,6 9, Wambianna Station, 7.5 km NW Gin Gin (AM KS76599, KS76602, KS76604, KS76606, KS76704); 1 9, Wee- melah, S of, 150m North of bridge over Ging- ham Watercourse (AM KS76706). TASMAN- IA: 1 9 , Bird Island, George Rocks (QVMAG 13:44297); 1 9, 1 9 with eggsac, Launceston (QVMAG 13:42995^6); 2 9 with eggsac, Launceston, 43 High St (QVMAG 13:44298); 1 9 with spiderlings, Launceston, Kings Meadows (QVMAG 13:42070); 1 6, Mt Cha- pell Island, Bass Strait (QVMAG 13:44299); 1 6, Waterhouse, South Croppies Point (QVMAG 13:43254). Victoria: 1 9, no exact location, 1923 (MV K7654); 1 9, no exact location (‘Teacher's Training College’) (MV K7649); 4 6, 3 9, Barmah Forest (WAM T53795, T55467); 1 6, near Baxter (MV K7774); 4 6,4 9, Bendigo, LaTrobe Univer- sity (CVIC); 1 9, 1 juv., Bendigo (MV K7658); 1 6, 2 9, 1 juv., Camberwell (MV K7653); 16 6,4 9, Cohuna, Kervies Rd, Barr Ck (MV K81 16); 2 9,2 juv., Dalyenong Flo- ra Reserve, Plantation Tk, 900 m S Gum Flat Tk (MV K9247, K9249); 2 9,6 juv., Deep Ck, 7 km SSE Barmah (MV K8724); 16,1 9, 2 juv.. Deep Lead Flora Reserve, Deep Lead Rd, 800 m NE of Western Hwy (MV K9226); 1 6, 1 9, 3 juv., Deep Lead Flora Reserve, 800 m SW Garnard Park/Deep Lead Rd along Deep Lead Rd (MV K9045, K9232); 1 6, 1 9,1 9 with spiderlings, 11 juv.. Deep Ck, 7 km SSE Barmah (MV K8719, K9057); 2 6, East St Kilda (MV K7650); 3 9, Eynes- bury Estate, Werribee (MV K9111, K9116, K9143); 1 9, Glen Waverley (MV K7651); 2 6, 1 juv., Glen Waverley, Watsons Rd (MV K7735); 4 9 , Goulburn River, 12 km SSE Na- thalia (MV K9029, K9041); 3 6, 1 9, Gray- town, 200 m N of Heathcote/Nagambie Rd, 80 m W on drive to abandoned house (MV K9238, K9262); 1 9, Hamilton (MV K7657); 2 9, Kilsyth, 38 Mountfield Rd (MV K7644^ 5); 1 9 with spiderlings, 1 9 with 16 spider- lings, 2 juveniles, Kotupna Barmah Rd at El- lingtons Bridge (MV K8748, K9053); 19,1 juv., Lerderderg Gorge, 9 km NNW of Bac- chus Marsh (MV K7655); 6 6,1 9,2 juv., Maldoe State Forest, 1.7 km along Red White and Blue Tk from Pullens Rd (MV K9246, K9250, K9253); 1 9 with spiderlings, Mel- bourne, in museum (MV K7656); 1 9, Mer- beie (AM KS32465); 1 6, Mitchell Link Tk, 200 m W Mitchell Tk (MV K9219); 1 9, Morrisons (MV K7648); 1 9, Murray Valley Hwy, Deep Ck Crossing (MV8775); 9 6, 6 9, 5 juv., Mt Bolangum Forest Reserve, 5.7 km N Aedersoris Rd, thee 200 m on minor Tk (MV K9209, K9229, K9233); 1 6, Mt Ida Flora Reserve, 2.3 km NW along Rodney Tk from Dargie Tk (MV K9244); 1 9, Nangiloc (AM KS86408); 1 9, Natimuk (MV K7646); 1 9 with spiderlings. North Melbourne (MV K7647); 2 6, 2 9, Point Cook (MV K9113- 4, K9135); 1 6, 1 9, Point Cook, opposite carpark 1 (MV K9106, K9108); 1 9, Point Cook, 100 m E of Recreation Beach area (MV K9109); 1 9, Point Cook, lower sanddune (MV K9112); 2 9, Point Cook, lower edge (MV K9115); 6 6,6 9, 10 juv., Pomfrets Rd, 0.6 km S Picola-Katuega Rd (MV K8767, K9034, K9050); 7 6, 4 9, Potter Creek, L7 km S of Western Highway (MV K7652); 3 9, Rathbones Rd, 3.0 km E Booths Rd (MV K8727, K8746); 3 6, 5 Juv., Reedy Lake Wildlife Reserve, 600 m W Goreys Rd along Reedy Lake Rd (MV K9259); 1 6, 1 juv., Reedy Lake Wildlife Reserve, 2. 1 km S along Reedy Lake Rd from Davies Rd (MV K9297); 19,19 with eggsac, Spring Gully (CVIC); 2 9, State Forest, 3.5 km NE Yambuna (MV K8708, 8759); 1 6, Upper Lurg (CVIC JStl04); 1 9, Werribee morticaie Saltmarsh (MV K9110); 2 9, Werribee Treatment farm (MV K9117); 1 9 with eggsac, West Bruns- wick (MV K8119); 1 6, Western Railway Rd (MV K9133); 2 9, Williamstown (MV K9107); 2 9, Yarra Valley Park (MV K7740). Etymology, — The species name is an ad- jective in apposition and refers to the Austra- lian state Victoria, which represents the center FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 31 of the distribution and the state where the ho- lotype was found. Diagnosis* — Males of A. victoriensis can be distinguished from all other Australian Artoria by the unique shape of the median apophysis that resembles an upside-down sock in ventral view. The female epigyeum is uniquely oval- shaped, with a white center and a sclerotized posterior rim reaching medially into this cen- ter. Description*-— Carapace (Fig. 63): Brown, with distinct light brown median and submarginal bands; head region black; dark grey radial pattern; carapace covered with white setae, particularly dense in median and submarginal bands and between PE; four brown bristles in median band anteriorly of fovea; two rows of black bristles between PME; few black bristles posterolateral of PME. Sternum: Light brown with dense, dark grey pigmentation; sparsely covered with brown bristles mainly towards margins and frontal border. Labium: Brown; front end trun- cate and white. Chelicerae: Uniformly brown; covered with long white setae and few brown bristles; three retrornarginal teeth, with the median largest; three promarginal teeth, with the basal smallest. Pedipalp (Figs. 65-67): Cymbium tip with approx, eight macrosetae; median apophysis shaped like an upside-down sock in ventral view; embolus stout and bluet, terminal apophysis a pointy hook (Fig. 67). Abdomen: Dark grey; yellowish-brown lan- ceolate heart mark in anterior half; irregular yellow patches lateral of heart mark; three yel- low chevrons in posterior half; covered with white setae and few longer, brown bristles; venter yellow-brown with irregular dark grey patches; covered with white and fewer brown setae; anterior spinnerets black, posterior spin- nerets yellow-brown. Legs: Leg formula IV > I > III > II; light brown, with distinct dark grey aeeulatioe which is in particular distinct on lighter ventral side of legs; spieation of leg 1 (based on holotype MV K7742): Femur: 3 dorsal, 1 apicoprolateral; tibia: 3 ventral pairs, 2 prolateral; metatarsus: 3 ventral pairs, 2 pro- lateral, 1 apicoventral, 1 apicoretrolateral. Female: Carapace: As male, more brown bristles in median band anteriorly of fovea. Sternum: Yellowish-brown with dense, dark grey pigmentation; sparsely covered with long brown bristles and few brown setae. Labium: as male. Chelicerae: Uniformly dark brown; covered with long white setae and few brown bristles; dentition as male. Epigynum (Figs. 68-69): Ventral view: oval shaped, wide scler- otizatioe reaching from posterior margin into center (Fig, 68); dorsal view: large oval sper- mathecae, copulatory ducts connected later- ally; small spermathecal organs (Fig. 69). Ab- domen: As male, light pattern less distinct; venter yellow-brown few dark grey patches in particular posterior of epigastric furrow; cov- ered with white and fewer brown setae; all spinnerets light brown. Legs: Leg formula IV > I > III > II; coloration as male, aeeulations less distinct; spination of leg I (based on par- atype MV K7741): Femur: 3 dorsal, 1 apico- prolateral; tibia: 3 ventral pairs, 1 prolateral; metatarsus: 3 ventral pairs, 3 prolateral, 1 ap- ico ventral. Measurements: Male holotype MV K7742 (female paratype MV K7741): TL 5.55 (6.5), CL 3.0 (3.0), CW 2.2 (2.2). Eyes: AME 0.08 (0.10), ALE 0.08 (0.09), PME 0.30 (0.30), PLE 0.20 (0.22). Row of eyes: AE 0.50 (0.54), PME 0.76 (0.80), PLE 0.98 (LOO). Sternum (length/width) 1.40/1.20 (1.45/1.2). Labium (leegth/'width) 0.42/0.4 (0.44/0.42). AL 2.4 (3.1), AW 1.9 (2.1). Legs: Lengths of seg- ments (femur + patella/tibia + metatarsus + tarsus — total length): Pedipalp 1.05 + 1.05 + — + 1.05 = 3.15, I 1.75 + 2.45 + 1.55 + 0.85 - 6.60, II 1.7 + 2.15 + 1.30 + 0.7 = .5.85, III 1.6 + 2.0 + 1.6 + 0.7 =.5.9, IV 2.1 + 2.75 + 2.4 + 1.0 = 8.25 (Pedipalp 1.05 + LO + — + 0.75 = 2.8, I 1.8 + 2.35 + 1.3 + 0.75 = 6.2, II 1.7 + 2.15 + 1.2 + 0.75 = 5.8, III 1.65 + 1.85 + 1.5 + 0.7 = 5.9, IV 2.2 + 2.75 + 2.45 + 0.95 = 8.35). Variation: Males (females) (range, mean ± s.d.): TL 3.5-6.3, 4.6 ± 1.0; n =10; CL 1.9- 3.3, 2.6 ± 0.5; n =11; CW L4-2.3, 1.9 ± 0.3; n =11 (TL 5.2-8.4, 6.4 ± 0.7, n = 19; CL 2.2-3.6, 3.0 ± 0.4, n =20; CW 1. 6-2.8, 2.2 ± 0.3; n =20). Distribution and habitat preferences* — This species is most common in temperate Victoria, South Australia and New South Wales (Fig. 70), where it can typically be found in open, moderately moist habitats. It is also common in suburban Adelaide and Mel- bourne. Within the South Australian artesian springs, A. victoriensis has been found at Kingfisher Springs in the Dalhousie Springs complex and at Big Depot Springs in the Francis Swamp complex (Table 1), where it 32 THE JOURNAL OF ARACHNOLOGY Figure 70. — Records of Artoria victoriensis new species in Australia. inhabits low open vegetation on saturated sub- strates. Remarks. — It is possible that the artesian springs populations were established by hu- man introduction as this species is most com- mon in highly populated suburban regions in South East Australia and has been found at artesian springs that are frequented by tourists. Females with eggsac or spiderlings have been found mainly in November and December, but also in September, February, and March. DISCUSSION The wolf spider fauna at South Australian artesian springs comprises a number of wet- land dependent species that have broad distri- butions, as well as others that appear to be more closely associated with the springs. Of the nine lycosids recorded during this study, seven occur in other Australian and overseas wetland habitats, such as river floodplains and lake shores (A. howquaensis, A. victoriensis, H. crispipes, H. diyari, H. kuyani, V. arenaris, and V. goyderi). In contrast, V. fontis and T. arabanae appear to be largely restricted to ar- tesian springs and have only rarely been found in other wetland habitats. The biology and habitat preferences of the artesian spring species are poorly understood. Venatrix goyderi and H. crispipes are mostly found at bore drains and it is possible that these species are recent arrivals to the region as they are rarely found at undisturbed arte- sian springs. The high dispersal capability of both species is also supported by their wide distribution, that includes offshore islands and island states including New Zealand. Venatrix arenaris and V fontis are both nocturnal spe- cies and are found mainly in the permanent wetland areas of the spring vent and tail, while the remaining species appear to be diurnal with some inhabitating the margins of the springs. Artesian spring wolf spiders in South Aus- tralia, like many lycosids, are dependent on a constant supply of water. For example, some lycosids are exclusively found near rivers (e.g. Manderbach & Framenau 2001, Framenau et al. 2002), lakes (Greenstone 1983), or coastal shore lines (e.g. McKay 1974; Dobel et al. 1990; Morse 1997, 2002). Certain behavioral adaptations may facilitate colonization of hab- itats near water bodies, such as mobile brood care, the capability to walk on the surface of the water (e.g. Ehlers 1939), and the ability to use polarized light for orientation (Papi 1955; Papi & Tongiorgi 1963; Ortega-Escobar & Munoz-Cuevas 1999). Most riparian species have been recorded to use water bodies as re- treats when predators are present and they can stay under water for a considerable period of time by trapping air in the dense cover of se- tae surrounding their body (V.W. Framenau & T.B. Gotch pers. obs.). However, an intriguing aspect of the lycosids associated with artesian springs is how they colonize these tiny, iso- lated habitats in an otherwise inhospitable en- vironment, given that several species are known to be extremely susceptible to even short periods of hot, dry conditions. Short distance dispersal by ballooning with- in spring groups (i.e., where springs are sep- arated by 50-1000 m) has been observed after monsoonal storms (T.B. Gotch pers. obs.), and dispersal may also occur during infrequent lo- calized floods when spiders could float be- tween springs. However, it is unknown how spiders move between more distant spring complexes (i.e., over lO's to of lOO's km), as these species are unable to survive for more than a few hours away from free water during summer (T.B. Gotch pers. obs.). It is possible that long distance ballooning may occur after summer rainfall when climatic conditions are optimal for ballooning, and/or that water- borne spider movement occurs as a result of extensive regional flood events. However, these events occur very rarely in the case of regional floods, while the chance of successful long distance ballooning must be considered very small, given that springs represent tiny targets, and the prevailing wind directions are largely west to east. Current research assess- ing the genetic differences among artesian FRAMENAU ET AL.— AUSTRALIAN ARTESIAN SPRING WOLF SPIDERS 33 spring lycosid populations along flood chan- nels in comparison to more remote popula- tions is aimed at testing indirectly whether one of these dispersal methods is more likely than the other. ACKNOWLEDGMENTS This study would not have been possible without the kind and generous support from the following individuals and their institu- tions: Graham Milledge and Mike Gray (AM), Bruce Halliday (ANIC), Jennifer Shields (CVIC), David Hirst (SAM), Ken Walker, Pe- ter Lillywhite, and Richard Marchant (MV), Jason Dunlop and Shahin Nawai (ZMB), Christine Rollard (MNHP), Janet Beccaloni (BMNH), Hieronymus Dastych (ZMH), Owen Seeman and Rob Raven (QM), Lisa Boutin (QVMAG), Liz Turner (TMAG), Peter Jager and Julia Altmann (SMF), Mark Harvey and Julianne Waldock (WAM), Kelli-Jo Kovac and Darren Niejalke (Western Mining Cor- poration), and Reg and Ronny Dodd (Marree Arabana Community). Jurgen Gruber (Natur- historisches Museum, Vienna) assisted in the (unfortunately unsuccessful) search for types of Trochosa candicans. Mark Elgar provided excellent laboratory facilities for VWF at the University of Melbourne during the initial stages of this study. TBG wishes to thank his intrepid field assistants Bruce Gotch, Paul Fitzpatrick, Darryl Fitzgerald and Sylvia Clarke. Thanks to Chris Wilcox and Hugh Possingham (University of Queensland) for their support, mentally and financially; the people of the outback for their hospitality, in particular the Clarke family, the Crozier fam- ily, the Sheahan family, the Sims family, and the Williams family. Melissa Thomas, Julian- ne Waldock, Mark Harvey, Torbjorn Krones- tedt and Cor Vink provided comments on ear- lier drafts of the manuscript. Funding for this project was provided by the Australian Bio- logical Resources Study (to Mark Harvey and ADA), Collex Flinders-Baudin Scholarship (to TBG), Western Mining Corporation (WMC) (to TBG) and the Department of En- vironment and Heritage, SA through a Wild- life Conservation Fund Grant (to TBG). We are particularly grateful to the elders of the Arabana, Diyari and Kuyani tribes for the per- mission to name several species after them. 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Zyuzin, A. A. 1993. Studies on the wolf spiders (Ar- aneae: Lycosidae). I. A new genus and new spe- cies from Kazakhstan, with comments on the Ly- cosinae. Memoirs of the Queensland Museum 33:693-700. Manuscript received 23 December 2003, revised 21 April 2004. 2006. The Journal of Arachnology 34:37-45 THE PREY OF A LITHOPHILOUS CRAB SPIDER XYSTICUS LOEFFLERI (ARANEAE, THOMISIDAE) Elchin Fizuli oglu Guseinov: Institute of Zoology of Academy of Sciences of Azerbaijan, kvartal 504, proyezd 1128, Baku 370073, Azerbaijan E-mail: elchin-f@ artel, net. az ABSTRACT, The natural prey of the crab spider Xysticus loejfleri Roewer 1955, living under stones, was studied. The percentage of feeding specimens in the population studied was low (1.4-4. 6%), and it declined with the beginning of the breeding period. Investigation has shown that X. loeffleri is a polyph- agous predator. Representatives of twelve arthropod orders were found in its diet. Arachnids (opilionids and spiders) formed the major food component constituting ca. 70% of prey captured. No insect order was present in any considerable percentage. Several worker ants were observed as prey suggesting that X. loejfleri is a myrmecophagic spider. Seven incidences of cannibalism were recorded, which all involved predation on adult conspecifics (two males and five females). The length of prey killed by X. loejfleri ranged between 1.25 and 15.00 mm (mean 4.68 mm) and constituted from 14.3-187.5% (mean 64.2%) of length of their captors. The most frequently captured prey were small arthropods not exceeding half the size of the spiders. Keywords; Crab spiders, lithophilic, prey, opilionids, cannibalism Thomisidae Sundevall 1833 (the true crab spiders) is one of the largest families of spi- ders including about 2000 species (Codding- ton & Levi 1991). Most crab spiders are typ- ical cursorial hunters which do not use silk for prey capture; instead, they lie in ambush and wait until prey comes within reach of their long forelimbs and seize it. In many terrestrial communities thomisids are among the domi- nant invertebrate predators, exerting signifi- cant pressure on prey populations (Young & Edwards 1990; Bogya & Mols 1996; Jennings & Cutler 1996; Nyffeler 1999). Despite the common occurrence and predatory signifi- cance of crab spiders, few studies have ad- dressed their natural diet. A survey of the spi- der literature revealed only eleven works that included quantitative data on the prey of Thomisidae (Broekhuysen 1948; Nyffeler & Benz 1979; Morse 1979, 1981, 1983; Ricek 1982; Lubin 1983; Dean et al. 1987; Agnew & Smith 1989; Castanho & Oliveira 1997; Romero & Vasconcellos-Neto 2003). All crab spiders studied in these works inhabit vege- tation or ground litter strata. However, many thomisids are known to live under stones. Un- like most cursorial spiders, which use spaces under rocks only as temporary shelters, a va- riety of thomisid species spend their entire life span here. Physical and microclimatic condi- tions of this microhabitat differ strongly from those of the surrounding environment (Cloud- sley-Thompson 1955). These conditions influ- ence the composition of local invertebrate fau- na and thereby the prey available to crab spiders. What is the prey spectrum of the Thomisidae living underneath rocks? How much does it differ from the diets of spiders occurring in other microhabitats? To answer these questions, I conducted an investigation of the natural prey of the crab spider Xysticus loejfleri Roewer 1955, which is among the commonest spiders found under stones in Azerbaijan. The range of this species also includes Turkey, Iran, Middle Asian re- publics of the former Soviet Union, and Af- ghanistan (Marusik & Logunov 1994). This is a pronounced lithophilous spider. Over the last several years I observed thousands of individ- uals of X. loejfleri under rocks, while none was seen on the open surface. Like most thomisids, X. loeffleri are typical ambushers which spend most of their time sitting im- mobile on the underside of rocks awaiting prey. These spiders have an annual life cycle (Guseinov, unpubl. data). Adult females are present from September through May, while males are found only in autumn, which ap- 37 38 THE JOURNAL OF ARACHNOLOGY pears to be the mating season (one mating pair was observed). Oviposition usually begins in early spring and continues to the end of May. Females spin hemispherical egg sacs on the underside of rocks which they guard until the young emerge (Guseinov, unpubl. data). Some females were observed guarding a second egg sac near the first empty cocoon. So, X. loejfleri seems to be an iteroparous spider. METHODS Investigation was carried out at “Bailov Park” in Baku City, Azerbaijan (40°38'N; 49°83'E). This habitat was characterized by pines Pinus eldaricus Medw., with an under- growth of short ephemeral grasses, predomi- nantly of Calendula persica C.A.M., Senecio vernalis W. & K., Medicago denticulata W., Carduus arabicus Jaqu., Hirschfeldia incana (L.), Erodium cicutarium (L.), Hedypnois cre- tica W., Pterotheca marschalliana (Rchb.), Torularia contortuplicata (Stapf.), Ornitho- galum gossonei Ten., Gagea tenuifolia (Boiss.), Poa bulbosa L., Anisanthea rubens (L.), Aegilops biuncialis Vis., Hordeum lepor- inum Link., Koeleria phleoides (VilL), Brom- opsis sp. Stones were prevalent on the ground in the study area, with X. loeffleri being among the commonest spider species under these stones. Two consecutive generations of X. loeffleri were observed throughout the study period. Spiders of the first generation were studied from 14 February-2 April 1997. Seven sur- veys were conducted during this time (ap- proximately once a week), which took about 13 hours. Spiders of the second generation were investigated from 9 September 1997-21 May 1998. Thirty-eight surveys were made during this period (on average one per week, but numbers of surveys varied greatly be- tween different months: from one in Septem- ber and October, when spiders were rare under stones, to six-seven in winter months, which were the peak of spider abundance). Over 56 hours were spent on these surveys. All surveys were done in daylight hours be- tween 1 1:00 and 17:00. During surveys, rocks in the study area were overturned and the mouthparts of each individual X. loejfleri found were inspected with a 4 power lupe to prevent small prey being overlooked. Stones were chosen randomly, but because the study area was not large (ca. 2500 m^) about 60- 70% of all appropriate sized stones (15-80 cm in diameter) in the study area were examined during each survey. Considering the low mo- bility of X. loejfleri it is highly likely that most spiders were observed repeatedly throughout the study period. Specimens with prey in their chelicerae were placed in separate vials con- taining 75% ethyl alcohol and brought back to the laboratory for measurement and prey identification. Spiders without prey were left in the field. At the same time, all spiders ob- served were classified into the following groups: (1) males; (2) solitary females; and (3) females guarding their egg sacs. During every survey, the numbers of spiders with and without prey were counted separately for each of these groups. A few additional prey items were collected during occasional observations in the spring and autumn of 1999 and the spring of 2000. Voucher specimens of X. loef- jleri and their prey items were deposited at the Institute of Zoology of the Academy of Sci- ences of Azerbaijan. A chi square test was used for statistical treatment of the data. Only raw numbers (count data), not proportions, were used for analysis throughout the paper. RESULTS Feeding percent.— Only 16 X. loeffleri- males were seen throughout the study period (none with prey). Thus, they are omitted in the following consideration and all subsequent references are to females. Of 2023 female observations made during the study period, only 80 (4.0%) included spi- ders with prey in their chelicerae. Females of first generation were observed feeding signif- icantly less frequently (6 prey records of 423 observations [1.4%]) than females of second generation (74 prey records of 1600 obser- vations [4.6%]) (x^ = 8.232; df = I, P < 0.001). Among females of the second gener- ation, spiders observed in winter months (De- cember-February) had the lowest feeding per- centages compared to spiders observed in autumn (September-November) and spring (March-May) (Table 1); the difference is sig- nificant (x^ = 4.168; df = 1; R < 0.05). Al- though these winter-feeding females exhibited higher percent of prey capture compared to solitary females of the first generation, the dif- ference is not significant (x^ = 1.857; df = 1; P > 0.1). The percentage of feeding speci- GUSEINOV— PREY OF XYSTICUS LOEFFLERI 39 Table 1. — Monthly variation in the number of spiders observed feeding in second generation female Xysticus loejfleri. Females found attending egg sacs are referred to as guarding. Females found without egg sacs are referred to as solitary. Month Number of sur- veys Number of spiders observed Number of spiders feeding Percentage of spiders feeding Solitary Guarding 9 9 9 9 S Solitary Guarding 9 9 9 9 S Solitary Guarding 9 9 9 9 S Sep./Oct. 2 25 — 25 2 — 2 8.0 — 8.0 Nov. 3 118 — 118 10 — 10 8.5 — 8.5 Dec. 7 357 — 357 18 — 18 5.0 — 5.0 Jan. 6 324 — 324 10 — 10 3.1 — 3.1 Feb. 7 284 — 284 19 — 19 6.7 — 6.7 Mar. 4 117 — 117 10 — 10 8.5 — 8.5 Apr. 5 27 213 240 2 2 4 7.4 0.9 1.7 May 4 4 131 135 — 1 1 — 0.8 0.7 mens among guarding females of second gen- eration (3 prey records of 344 observations [0.9%]) was significantly lower than among their solitary counterparts (71 prey records of 1256 observations [5.7%]) (x^ ~ 12.929; df = l\ P < 0.001). Guarding females of first gen- eration were also observed feeding less fre- quently (none found with prey in 89 obser- vations) than solitary females (6 prey records of 334 observations [1.8%]). However, these data are not sufficient for statistical analysis. Prey composition. — Altogether 88 prey items were taken from X. loejfleri. These were distributed among twelve arthropod orders (Table 2). Arachnids formed the major com- ponent in the food of X. loejfleri (ca. 70%). Opiliones (Phalangiidae, Opilio spp.) was the dominant prey order constituting 40.9% of the total prey. Spiders represented by 10 species from 5 families accounted for 28.4% of all prey caught (Table 3). Thomisidae were most abundant (52% of all spiders killed), followed by Theridiidae Sundevall 1833 (28%), Gna- phosidae Pocock 1898 (12%), and Oecobiidae Blackwall 1862 (8%). Seven conspecifics were captured by X. loejfleri, including five females and two males. Insects comprised 29.5% of all prey re- cords. However, no insect order was present in any considerable percentage (more than 10%). Most abundant were Coleoptera and Hymenoptera, 9% and 8% respectively. Co- leoptera consisted of four adult beetles (Car- abidae, Curculionidae, Histeridae) and four larvae (Carabidae). Hymenoptera included two parasitic wasps (Ichneumonidae) and five worker ants (Formicidae) represented by four Messor denticulatus Lepeletier and one Lep- tothorax sp. The remaining insects comprised three Hemiptera (Lygaeidae, Nabidae, Pyrrho- coridae), two Thysanura (Machilidae, Lepis- matidae), two Collembola (Sminthuridae), one Psocoptera (unidentified), one Homoptera (Aphididae), one Embiomorpha (Oligotomi- dae: Haploembia solieri Ramb.) and one Lep- idoptera larvae (Noctuidae). The only centi- pede captured by X. loejfleri was a lithobiid. Feeding phenology. — The study period covered the entire life span of adult females of the second generation and allowed me to consider seasonal changes in their diet. As seen from Table 4, there are differences in monthly distribution of some prey taxa cap- tured. While most arthropods were primarily caught in winter (December-February), adult beetles and ants were captured only in autumn (September-November) and spring (March- May). This difference becomes more striking if we examine the distribution of prey taxa captured between periods reflecting changes in the temperature regime. Harvestmen and most other arthropods were caught only dur- ing the cool season (late autumn-early spring). In contrast, Formicidae and adult Co- leoptera were captured only during the warm periods (early autumn and late spring). Spi- ders were caught throughout the course of the study. Length of prey. — Eighty-two prey items were measured. Their lengths varied from 1.25-15.00 mm (mean ± SD: 4.68 ± 3.10 mm) and constituted from 14.3-187.5% (64.2 40 THE JOURNAL OF ARACHNOLOGY Table 2. — Prey taken by Xysticus loeffleri under stones. The larvae of holometabolous insects are marked with an asterisk. Otherwise, holometabo- lous insects are adults. Prey taxa N % Insecta Collembola Sminthuridae 2 2.3 Thysanura Machilidae 1 1.1 Lepismatidae 1 1.1 Embiomorpha Oligotomidae 1 1.1 Psocoptera Unidentified 1 1.1 Homoptera Aphididae 1 1.1 Hemiptera Lygaeidae 1 1.1 Nabidae 1 1.1 Pyrrhocoridae 1 1.1 Coleoptera Carabidae 1 1.1 Carabidae* 4 4.5 Curculionidae 2 2.3 Histeridae 1 1.1 Hymenoptera Ichneumonidae 2 2.3 Formicidae 5 5.7 Lepidoptera Noctuidae* 1 1.1 Arachnida Opiliones Phalangiidae 36 40.9 Araneae Oecobiidae 2 2.3 Theridiidae 7 8.0 Gnaphosidae 3 3.4 Thomisidae 13 14.8 Chilopoda Lithobiomorpha Lithobiidae 1 1.1 Total 88 100.0 ± 42.2%) of the size of their captors which ranged from 4.75-9.00 mm (7.37 ± 0.95 mm). Size distribution of prey is shown in Fig. 1. The most abundant were small arthro- pods not exceeding half the size of the spiders, which accounted for 53.7% of the total prey measured. To this group belonged collembo- lans, an aphid, opilionids, a Leptothorax ant, oecobiids, theridiid spiders, and conspecific males. Medium- sized prey (from 50-100% of spider body length) constituted 25.6% and in- cluded a psocopteran, curculionid beetles, a lygaeid bug, Ozyptila, Xysticus sp., gnaphosid spiders, some Messor ants and some conspe- cific females. One fifth of the prey of X. loef- fleri (20.7%) consisted of large arthropods ex- ceeding the length of their captors. These were thysanurans, an embiomorpha, nabid and pyr- rhocorid bugs, carabid beetles, lepidopteran larvae, ichneumonid wasps, a lithobiid centi- pede, some Messor ants and some conspecific females. DISCUSSION As is typical of cursorial spiders (Nentwig 1986; Nyffeler et al. 1994), the percentage of feeding specimens in the population of X. loeffleri was low. The difference in percentage of feeding specimens between two generations is probably due to the fact that most of the observations of first generation females were made in February and early March, character- ized by low temperatures, which probably re- sulted in inhibited prey activity and, as a con- sequence, low prey capture by spiders. This assumption is confirmed by the data on sea- sonal changes in the feeding percent of soli- tary females of the second generation. Spiders observed in winter months were found feed- ing significantly less frequently than spiders observed in autumn and spring. In contrast, the difference between these winter-feeding females and solitary females of first genera- tion was not significant. Despite the fact that egg-guarding females occurred only in warm period (late spring), in both years the per- centage of feeding specimens among them was lower than among solitary females. Un- like females of an anthophilous thomisid, Mis- umena vatia (Clerck 1757), which build their reproductive nests on leaves, far away from their typical hunting site, flowers (Morse 1985), X. loeffleri females construct their egg sacs on the underside of rocks i.e. at the same site where they usually forage. This enables the spiders to catch prey during egg guarding period. However, most thomisids are pro- nounced ambushers, and the choice of prey- rich foraging sites is an important trait of their GUSEINOV—PREY OF XYSTICUS LOEFFLERI 41 Table 3. — Spiders captured by Xysticus loeffleri. Spider species N Sex-age stage Oecobiidae Oecobius maculatus Simon 2 1 submale, 1 female Theridiidae Enoploghatha gemina Bosmans et Van Keer 5 1 male, 3 females, 1 subfemale Enoplognatha quadripunctata Simon 1 1 female Theridion melanurum Hahn 1 1 female Gnaphosidae Drassodes lapidosus (Walckenaer) 1 1 juvenile Haplodrassus dalmatensis (L. Koch) 2 2 juveniles Thomisidae Ozyptila tricoloripes Strand 3 1 submale, 2 females Xysticus loeffleri Roewer 7 2 males, 5 females Xysticus sp. 3 3 females feeding strategy (Morse & Fritz 1982; Beck & Connor 1992). While guarding their eggs, female X loeffleri have no opportunity to change their locations apparently resulting in the declined percent of prey capture compared to solitary females. Investigation has shown that X loeffleri is a polyphagous predator feeding on a wide range of prey. The predominance of opilionids in its diet is unusual. To my knowledge no spiders are known to feed on harvestmen in any considerable percentage. Thus it might be suspected that X loeffleri specializes on opi- lionids as an unusual less available prey to spiders. However, this fact is more likely due to the abundance of harvestmen in the envi- ronment of X loeffleri. The density of poten- tial prey has not been quantified, but, subjec- tively, opilionids appeared to be one of the most numerous arthropods inhabiting spaces under stones. Furthermore, some other hunt- ing spiders, such as Philaeus chrysops (Poda 1761), Ozyptila tricoloripes Strand 1913, Thanatus kitabensis Charitonov 1946 and Drassodes lapidosus (Walckenaer 1802), were repeatedly seen feeding on harvestmen in this microhabitat. In contrast, only two opilionids were found among about 1500 prey organisms Table 4. — Monthly distribution of prey taxa captured by second generation female Xysticus loeffleri. In round brackets are the mean monthly temperatures (°C). In square brackets are the numbers of spider observations made during a given month. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May (18.8) (17.5) (10.8) (6.2) (3.9) (3.0) (6.6) (12.9) (14.2) Prey taxa [8] [17] [118] [357] [324] [284] [117] [240] [135] Opiliones 3 6 6 13 3 Araneae 1 3 6 2 2 5 1 Formicidae 1 2 1 Coleoptera (adult) Coleoptera (larvae) Ichneumonidae 1 1 3 1 2 1 Hemiptera Homoptera 1 1 Thysanura Collembola 1 2 1 Psocoptera Lepidoptera 1 1 Lithobiomorpha 1 42 THE JOURNAL OF ARACHNOLOGY 20 o o O O O o o o o o V“ CO IT) N- O) CO iO r^ 1 o o O 1 o 1 o 1 1 -T” 1 1 1 CN CD 00 o o o o o o CM CD 00 Size categories Figure 1. — Distribution of prey in different size categories. taken from various species of cursorial spiders frequenting other microhabitats (bare ground, herbaceous vegetation, ground litter, bark of trees, stone walls etc.) in the vicinity of Baku City in years 1997-1999 (Guseinov 1999). Thus, opilionids do not seem to be invulner- able prey to cursorial spiders, which probably take them in proportion to their abundance. Which, in turn, apparently depends on the type of microhabitat occupied by the spiders. It is remarkable that some insects (Thysanura, Embiomorpha, Coleoptera larvae) as well as a lithobiid centipede captured by X. loeffleri are also characteristic inhabitants of spaces under rocks, but usually lacking among the prey of spiders from other microhabitats (Bristowe 1941; Nentwig 1987; Nyffeler 1999). On the other hand, winged insects (es- pecially Diptera), constituting a substantial part of the food of most cursorial spiders (Nentwig 1986; Guseinov 1997), are almost entirely missing in the diet of X. loeffieri. The tight, constricted spaces under rocks are not favorable environment for winged insects and, as a consequence, crawling arthropods prevail among the prey of X. loeffleri. The high proportion of spiders in the diet of X. loeffleri is also probably due to their abundance in its habitat. Many spiders are known to occur under stones and during the cool season their number may even increase in this microhabitat. Although X. loeffleri cap- tured mainly cursorial spiders, several individ- uals of the family Theridiidae most of which are typical web builders were also eaten. The webs of spiders serve not only for prey cap- ture, but are also as efficient defensive devic- es. Thus only a small minority of spiders are able to invade alien webs and prey upon their residents (Jackson 1992). Most theridiids spin large three-dimensional space-webs. However, the habits of theridiids captured by X. loeffleri are apparently different from this common pattern of life style. These spiders were fre- GUSEINOV— PREY OF XYSTICUS LOEFFLERl 43 quently found on the underside of rocks with= out any silk or with several short threads laid down over the substrate. Therefore, they do not appear to be a more '‘difficult” prey for predators than typical cursorial spiders. Xysticus loeffleri is a cannibalistic spider with conspecifi.cs constituting 8% of its prey. Such a high rate of cannibalism is unusual for crab spiders which generally do not hunt con- specifics (Bristowe 1941; Broekhuysen 1948; Morse 1981, 1983; Ricek 1982), but similar to rates of cannibalism of other cursorial spi- ders from families Salticidae Blackwall 1841 (Jackson 1977), Lycosidae Sundevall 1833 (Schaefer 1974; Frameeau et ah 1996), Oxy- opidae Thorell 1870 (Turner 1979; Nyffeler et al. 1987a, 1987b, 1992) and Sparassidae Bert- kau 1872 (Henschel 1994). Moreover, it should be emphasized that most conspecifics killed by X. loeffleri were mature females (71.4%), with size comparable to that of their captors, whereas in cannibalistic lycosid spi- ders larger individuals usually catch smaller ones (Edgar 1969; Hallander 1970; Yeargan 1975). But such a situation is excluded in the case of X. loeffleri because the population consisted of individuals of the same age (Gus- einov, uepubl. data). Despite the fact that worker ants are not acceptable prey to most cursorial spiders (Nentwig 1986), they were found in the diet of X. loeffleri, though in low proportion (5%). At the same time, it is known that worker ants compose a considerable portion (30-35%) of the food of some species of the genus Xysticus (Nyffeler & Benz 1979; Guseinov 1997). It should be clarified, therefore, whether X. loef- fleri is a poor predator of ants or if ants are simply underrepresented in the species' habi- tat. Some data correspond to the second as- sumption, All ants were caught in early au- tumn and late spring. Yet, the number of prey records at that time was significantly lower than that in the cool season because the fre- quency of surveys conducted was low in early autumn and most females had oviposited in late spring resulting in a low prey capture among those females. Thus one can suppose that if the number of prey records in warm periods was greater, then the proportion of ob- servations of ants in the diet of X. loeffleri might be larger. Although small arthropods predominated in the diet of X loeffleri, it does not signify that spiders prefer prey of this size category. This fact is more likely due to the abundance of small prey in the spiders' habitat, since the dominant prey type, opilionids, consisted pri- marily of small specimens. Probably the ap- propriate prey size range for X loeffleri is within. 20-120% of spiders' body size, since larger or smaller organisms were rare in its diet (see Fig. 1). Earlier students of crab spiders have point- ed out that thomisids often catch very large prey (Lovell 1915; Hobby 1931, 1940; Turner 1946). In feeding expiriments, most cursorial spiders preferred prey not exceeding their own size, whereas the crab spiders, Xysticus cm- tatus (Clerck 1757), readily accepted insects two times larger than themselves (Nentwig & Wissel 1986). Although X. loeffleri sometimes captured very large arthropods, most of its prey (ca. 80%) were not exceeding spider length. This is similar to prey size spectra of “typical” cursorial hunters (Salticidae, Lycos- idae, Oxyopidae, Sparassidae) (Nentwig & Wissel 1986; Hayes & Lockley 1990; Nyffeler et al. 1992; Henschel 1994), but in striking contrast to flower-dwelling Thomisidae, which commonly feed on prey significantly larger than themselves (Nentwig & Wissel 1986; Guseinov 1999). Experimental investigation is required to clarify whether this difference is due to the difference in size of prey available on flowers and under stones or anthophilous crab spiders are superior toward X. loeffleri in catching large prey. LITERATURE CITED Agnew, C.W. & J.W. Smith. 1989. Ecology of spi- ders (Araneae) in a peanut agroecosystem. En- vironmental Entomology 18:30-42. Beck, M.W. & E.F. Connor. 1992. Factors affecting the reproductive success of the crab spider Mis- umenoides formosipes: the covariance between juvenile and adult traits. Oecologia 92:287-295. Bogya, S. & P.J.M. Mols. 1996. The role of spiders as predators of insect pests with particular ref- erence to orchards: a review. Acta Phytopathol- ogica et Entomologica Hungarica 31:83-159. Bristowe W. 1941. The Comity of Spiders, voL 2, The Ray Society, London. Broekhuysen, G.J. 1948. The behaviour and the life history of a Javanese spider, Thomisus sp. Jour- nal of the Entomological Society of South Africa 10:135-164. Castanho, L.M. & P.S. Oliveira. 1997. Biology and behaviour of the neotropical ant-mimickieg spi- der Aphantochilus rogersi (Araneae: Aphanto- 44 THE JOURNAL OF ARACHNOLOGY chilidae): nesting, maternal care and ontogeny of ant-hunting techniques. Journal of Zoology 242: 643-650. Cloudsley-Thompson, J.L. 1955. The effect of rock cover on the diurnal range of microclimatic con- ditions. Entomologist 89:1120. Coddington, J.A. & H.W. Levi. 1991. Systematics and evolution of spiders (Araneae). Annual Re- view of Ecology and Systematics 22:565-592. Dean, D.A., W.L. Sterling, M. Nyffeler & R.G. Breene. 1987. Foraging by selected spider pred- ators on the cotton fleahopper and other prey. Southwestern Entomologist 12:263-270. Edgar, W.D. 1969. Prey and predators of the wolf spider Lycosa lugubris. Journal of Zoology 159: 405-411. Framenau, V., M. Reich & H. Plachter. 1996. Zum Wander verhalten und zur Nahrungsokologie von Arctosa cinerea (Fabricius, 1777) (Araneae: Ly- cosidae) in einer alpinen WildfluBlandschaft. Verhandlungen der Gesellschaft fiir Okologie 26: 369-376. Guseinov, E.E 1997. Preliminary data on prey com- position of some species of cursorial spiders (Ar- aneae) inhabiting Apsheron Peninsula, Azerbai- jan. Deposited in AZNIINTI Baku, No 2500- Az, 5p. (In Russian). Guseinov, E.E 1999. Spiders of Lenkoran nature area and Apsheron Peninsula, Azerbaijan. Auto- referate of the Thesis of Candidate (Ph.D.) of Biological Sciences Degree. Baku. 29p. (In Rus- sian). Hallander, H. 1970. Prey, cannibalism and micro- habitat selection in the wolf spiders Pardosa che- lata O.E Muller and P. pullata Clerck. Oikos 21: 337-340. Hayes, J.L & TC. Lockley. 1990. Prey and noctur- nal activity of wolf spiders (Araneae: Lycosidae) in cotton fields in the Delta Region of Mississip- pi. Environmental Entomology 19:1512-1518. Henschel, J.R. 1994. Diet and foraging behaviour of huntsman spiders in the Namib dunes (Ara- neae: Heteropodidae). Journal of Zoology 234: 239-251. Hobby, B.M. 1931. Spiders and their insect prey. Proceedings of the Royal Entomological Society of London 5:107-110. Hobby, B.M. 1940. Spiders and their prey. Ento- mologist’s Monthly Magazine 76:258-259. Jackson, R.R. 1977. Prey of the jumping spider F/i/- dippus johnsoni (Araneae: Salticidae). Journal of Arachnology 5:145-149. Jackson, R.R. 1992. Eight-legged tricksters. Spiders that specialize in catching other spiders. Bio- Science 42:590-598. Jennings, D. & B. Cutler. 1996. Crab spiders (Ar- aneae: Philodromidae, Thomisidae) of Ramsey County, Minnesota. Forest Service General Tech- nical Report NC-185. 35p. Lovell, J.H. 1915. Insects captured by the Thom- isidae. Canadian Entomologist 47:115-116. Lubin, Y.D. 1983. An ant eating crab spider from the Galapagos. Noticias de Galapagos 37:18-19. Marusik, YM. & D.V. Logunov. 1994. The crab spiders of Middle Asia (Aranei, Thomisidae), 2. Beitrage zur Araneologie 4:133-175. Morse, D.H. 1979. Prey capture by the crab spider Misumena calycina (Araneae: Thomisidae). Oec- ologia 39:309-319. Morse, D.H. 1981. Prey capture by the crab spider Misumena vatia (Clerck) (Thomisidae) on three common native flowers. American Midland Nat- uralist 105:358-367. Morse, D.H. 1983. Foraging patterns and time bud- gets of the crab spiders Xysticus emertoni Key- serling and Misumena vatia (Clerck) (Araneae: Thomisidae) on flowers. Journal of Arachnology 11:87-94. Morse, D.H. 1985. Nests and nest-site selection of the crab spider Misumena vatia (Araneae, Thom- isidae) on milkweed. Journal of Arachnology 13: 383-390. Morse, D.H. & R.S. Fritz. 1982. Experimental and observational studies of patch choice at different scales by the crab spider Misumena vatia. Ecol- ogy 63:172-182. Nentwig W. 1986. Non- webbuilding spiders: prey specialists or generalists? Oecologia 69:571-576. Nentwig, W. 1987. The prey of spiders. Pp. 249- 263. In Ecophysiology of spiders. (W. Nentwig, ed.). Springer- Verlag, Berlin. Nentwig, W. & C. Wissel. 1986. A comparison of prey lengths among spiders. Oecologia 68:595- 600. Nyffeler, M. 1999. Prey selection of spiders in the field. Journal of Arachnology 27:317-324. Nyffeler, M. & G. Benz. 1979. Nischeniiberlappung beziiglich der Raum-und Nahrungskomponenten bei Krabbenspinnen (Araneae: Thomisidae) und Wolfspinnen (Araneae: Lycosidae) in Mahwie- sen. Revue suisse de Zoologie 86:855-865. Nyffeler, M., D.A. Dean. & W.L. Sterling. 1987a. Predation by green lynx spider, Peucetia viridans (Araneae: Oxyopidae), inhabiting cotton and woolly croton plants in east Texas. Environmen- tal Entomology 16:355-359. Nyffeler, M., D.A. Dean. & W.L. Sterling. 1987b. Evaluation of the importance of the striped lynx spider, Oxyopes salticus (Araneae: Oxyopidae), as a predator in Texas cotton. Environmental En- tomology 16:1114-1123. Nyffeler, M., D.A. Dean. & W.L. Sterling. 1992. Diets, feeding specialization, and predatory role of two lynx spiders, Oxyopes salticus and Peu- cetia viridans (Araneae; Oxyopidae), in a Texas cotton agroecosystem. Environmental Entomol- ogy 21:1457-1465. Nyffeler, M., W.L. Sterling. & D.A. Dean. 1994. GUSEINOV—^PREY OF XYSTICUS LOEFFLERI 45 How spiders make a living. Environmental En- tomology 23:1357-1367. Ricek, E.W. 1982. Die Lauerposten der Krabben- spinne Xysticus bifasciatus C. L. Koch. Linzer biologische Beitrage 14:15-22. Romero, Q.R. & J. Vasconcellos-Neto. 2003. Nat- ural history of Misumenops argenteus (Thomis- idae): seasonality and diet on Trichogoniopsis adenantha (Asteraceae). Journal of Arachnoiogy 31:297-304. Schaefer, M. 1974. Experimentelle Untersuchungen zur Bedeutung der interspecifischen Konkurrenz bei 3 Wolfspinnen-Arten (Araneida: Lycosidae) einer Salzwiese. Zoologische Jahrbiicher. Abtei- liing fur Systematyk, Okologie und Geographie der Tiere 101:213-235. Turner, A.H. 1946. The prey of Misumena calycina (Arachn., Thomisidae). Entomologists 's Record 58:113-114. Turner, M. 1979. Diet and feeding phenology of the green lynx spider, Peucetia viridans (Araneae: Oxyopidae). Journal of Arachnoiogy 7:149-154. Young, O.P. & G.B. Edwards. 1990. Spiders in United States field crops and their potential effect on crop pests. Journal of Arachnoiogy 18:1-27. Yeargan, K.V. 1975. Prey and periodicity of Par- dosa ramulosa (McCook) in alfalfa. Environ- mental Entomology 4:137-141. Manuscript received 25 March 2002, revised 1 July 2004. 2006. The Journal of Arachnology 34:46-50 FIRST SPECIES OF HESPEROPILIO (OPILIONES, CADDOIDEA, CADDIDAE) FROM SOUTH AMERICA Jeffrey W* Shultz: Department of Entomology, University of Maryland, College Park, MD 20742 U.S.A. E-mail: jshultz@umd.edu Tomas Cekalovic: Casilla 764, Concepcion, Chile ABSTRACT, This paper describes the first South American species of Hesperopilio Shear 1996, a genus previously known from a single species, H. mainae Shear 1996, from Western Australia. The new species is known from a single adult female and is one of the largest and most colorful species of the superfamily Caddoidea. The generic diagnosis of Hesperopilio is emended to accommodate information from the new species. RESUMEN. En el presente articulo se describe la primera especie sudamericana de Hesperopilio Shear 1996, un genero previamente representado por una sola especie, H. mainae Shear 1996, de Western Australia. La nueva especie se conoce de una sola hembra adulta y es de las mas grandes y mas coloridas de la superfamilia Caddoidea. La diagnosis generica de Hesperopilio se enmienda para acomodar la in- formacion de esta nueva especie. Keywords: Hesperopilio, Opiliones, harvestman, Chile, systematics, South America, new species Until recently, the acropsopilionine faunas of the Australian Region (Acropsopilio SiL vestri 1904, Austropsopilio Forster 1955, Hes- peropilio Shear 1996, Tasmanopiiio Hickman 1957), South Africa (Caddella Hirst 1925) and the New World (Acropsopilio) displayed little generic overlap, with the principal ex- ception being the presence of Acropsopilio in the Australian Region, the Americas and Ja- pan, However, a new species from the “Aus- tralian” genus Austropsopilio was recently discovered in southern South America (Co- kendolpher & Maury 1990; Shultz & Ceka- lovic 2003), thus further increasing the simi- larity of acropsopilionine faunas of the two continents. This paper continues this trend by describing a new South American species of Hesperopilio, a genus formerly known from one species from Western Australia, H. mad nae Shear 1996. The new species is substan- tially larger and more colorful than the Aus- tralian species, and its discovery further highlights the close biogeographic connection between the Australian and South American opilion faunas. The material examined for this study is lodged in the American Museum of Natural History, New York (AMNH). SYSTEMATICS Hesperopilio Shear 1996 Hesperopilio Shear, 1996: 456. Type species.— Hesperopilio mainae Shear 1996, by original designation. Emended generic diagnosis.— Caddidae with ocularium large, broad, not projecting beyond anterior margin of carapace; ocula- rium with broad median furrow, each carina with longitudinal row of five variably devel- oped protuberances. Female palpal femora without ventral apophyses but with midventral row of stout spines; patella with prolateral field of glandular spines, tibia inflated and prolaterally spieose. Ovipositor with terminal apparatus composed of three bilaterally paired apical segments and shaft composed of 10 or 12 unpaired segments. Hesperopilio magnificus new species Figs. 1-7 Type data. — Holotype female: CHILE: Provincia Chiloe: Isla Grande de Chiloe, Che- pu (42°00'S, 73°58'W), 14 January 2002, To- mas Cekalovic (AMNH). Etymology. — The species is named for its magnificent coloration and comparatively large size. 46 SHULTZ & CEKAWWIC—HESPEROPILIO FROM SOUTH AMERICA 47 Figures 1-7. — Hesperopilio magnificus, adult female, holotype: 1. Right pedipalp, prolateral aspect; 2. Right leg I, retrolateral aspect; 3. Body, left lateral aspect; 4. Body, dorsal aspect; 5. Body, ventral aspect; 6. Ovipositor, ventral aspect; 7. Right leg III, retrolateral aspect. Abbreviations: OZ = ozopore; b = brown; db = dark brown; lb = light brown; Irb = light reddish brown; rb = reddish brown; vdb = very dark brown; w = white, yb = yellowish brown. Scale bar applies to all figures. 48 THE JOURNAL OF ARACHNOLOGY Diagnosis* — Hesperopilio magnificus is the first species of its genus known from South America. At. 23 mm long, the holotype is substantially larger than female H, mainae (1.6 mm) (Shear 1996) and has a much more complex coloration (Figs. 2-5, 7), including an asymmetrical white “hour glass” dorsal figure bordered by dark brown bands; dorsal transverse rows of white tubercles; and white and reddish-brown striped legs. The prolateral spine field of the palpal patella is inflated in H. magnificus (Fig. 1); the patella-tibia joint operates so as to bring the prolateral surfaces of the patella and tibia in apposition rather than the ventral surfaces (Fig. 1); the tibia is greatly inflated (Figs. 1, 3); the tarsus is pro- portionally larger in H. magnificus (subequal to tibia) (Figs. 1, 3); the palpal claw is greatly reduced (Fig. 3). The carinal tubercles of the ocularium vary greatly in size, the second be- ing the largest, followed by the third; the re- mainder are reduced to low bumps. The ovi- positor shaft has more unpaired segments (12 instead of 10 in H. mainae) and there are more setae (12 or 8) per seta-bearing segment than in H. mainae (6). Description of female holotype.- — Dorsal surface.* Anterior margin with shallow median emarginatioe (Fig. 4). Ozopores located at level of coxa I, open laterally, indicated dor- sally by wide, shallow emarginatioe. Surfaces of carapace slope upward steeply to form dor- sal “peak” on which ocularium is mounted (Fig. 3). Ocularium large; width, including lenses, over one-half width of carapace; wide, shallow mediae furrow; carinae with two pairs of large transverse ridges terminating laterally with dorsolaterally projecting, bluet-ended processes; smaller ridges between and poste- rior to large ridges (Figs. 3, 4). Metapeltidium with one transverse row of tubercles and dor- sal surface of opisthosoma with seven seg- mentally arranged, transverse rows of tuber- cles (Fig. 4). First (metapeltidial) and second (first opisthosomal) rows with five tubercles (one median, two medial, two lateral), rows 3 and 4 with four tubercles (two medial, two lateral), row 5 with three tubercles (one me- dian, two medial) and two lateral white spots may correspond to lateral tubercles, row 6 with three (one median, two medial), and rows 7 and 8 with two tubercles (medial only). First four tergal somites of the opisthosoma not de- marcated externally except by patterns of tu- bercles and coloration, tergite 5 distinguished anteriorly by incomplete transverse groove, tergites 6, 7, 8 + 9 and anal operculum de- marcated by transverse grooves. Ventral surface: Epistomal lobe (“labrum”) short, blunt (Fig. 5). Coxapophysis I with white, transverse lateral portion and brown m.edial portion with row of six long setae. La- bium with thin, transparent distal portion and sclerotized brown basal portion terminating in pair of setae. Coxapophysis II with white, transversely oriented basal portion terminating in brown lobe with ‘crown' of six setae. Coxa III without coxapophysis, extending medially to base of distal lobe of coxopophysis 11. Coxa IV without coxapophysis, extending antero- medially to level of coxa III; margin adjacent to operculum with one large seta; coxa ter- minating anteriorly under the genital opercu- lum with small, medially projecting lobe. There is a pair of ventrally projecting rectan- gular processes between the labium and an- terior margin of genital operculum that ap- pears to represent a portion of the arculi genitales. Coxa I with about 20 setae, coxae II-IV with eight or fewer setae. Genital oper- culum narrowing gradually toward slightly rounded anterior margin; surface with about 15 scattered setae. Ventral surface of opist- hosoma with a few scattered setae, otherwise smooth; segmentation indicated by rows of si- gilla. Preanal sternite with posterior median notch. Chelicera: Proximal segment mottled brown, red-brown and white, with a few scat- tered setae. Second segment mottled brown and white with about 20 setae, most arranged in an imperfect longitudinal row on the ante- rior surface. Cheliceral fingers dark brown, in- ner margins toothed. Palp: The palp is illustrated in Figs. 1^2, 5. Trochanter with large, blunt process pro- jecting from distal ventral surface, terminating in crown of about seven macrosetae. Femur: cylindrical, expanded slightly at distal end; proximal ventral end with rounded promi- nence; proximal two-thirds of ventral surface with imperfect longitudinal row of about 12 stout macrosetae; distal prolateral surface with blunt-ended process terminating in crown of about nine macrosetae; otherwise surface with a few scattered setae. Patella: subequal to fe- mur; middle third of prolateral surface greatly expanded to form sub-hemispherical promi- SHULTZ & CEKALOVIC—HESPEROPILIO FROM SOUTH AMERICA 49 nence with numerous, evenly spaced, glan- dular macrosetae; distal half of ventral surface with imperfect line of four stout, tubercle- based macrosetae; otherwise with a few scat- tered setae, especially on dorsal and distal re- trolateral surfaces. Tibia: proximally narrower than adjacent patella, makes sharp dorsal bend at one-quarter of length and expands in di- ameter distally; distal three-quarters of prola- teral surface expanded to form large hemi- spherical prominence covered with numerous, evenly spaced glandular macrosetae; other- wise surface with a few scattered setae. Tar- sus: narrow proximally but expanded distally; distal half of prolateral surface expanded into a rounded prominence with about 40 glandu- lar macrosetae; distal half of retrolateral sur- face with numerous, distally projecting micro- setae and a few scattered larger setae. Legs: Only leg I and leg III from the right side were attached to the holotype specimen (Figs. 2, 7); all other legs missing. Coloration: Dorsal surface with prominent flat-white central figure beginning anteriorly as median stripe on ocularium and continuing posteriorly to anal operculum (Fig. 4). Central figure broad (about one-third width of body) posterior to ocularium, gradually narrows pos- teriorly reaching narrowest point (about one- fifth width of body) at third row of tubercles; figure broadens posteriorly to almost full width of body at posterior margin of tergite 5; figure substantially narrower on tergite 6 and gradually narrows to the anal operculum. Cen- tral figure with asymmetrically shaped, me- dian islands of light reddish brown, associated with median and medial white tubercles. Cen- tral figure bordered laterally by dark brown longitudinal bands with irregular margins. Dark bands begin anteriorly along an irregular line that begins medially at the base of the ocularium and extends posterolaterally to a level near the anterior margin of coxa III; dark bands narrow posteriorly as their lateral bor- ders move progressively away from the lateral margins of the body. Dark bands bordered lat- erally by an ill-defined band of mixed light brown and reddish brown, with reddish brown dominating at the lateral periphery. White lat- eral tubercles form a curved longitudinal row along the border of the dark and mixed brown-reddish brown bands. A narrow strip of reddish brown continues anteriorly around the margin of the carapace. An irregular trans- verse, opalescent-white band crosses the car- apace anterior to the ocularium, two thin fin- gers of which project anteriorly to either side of the anterior emargination separated by a dark brown median line and bordered laterally by two dark brown islands. The opalescent- white band contains dark brown islands which appear to indicate sites of muscle attachment. A large black band encircles each lens; the band projects slightly anteriorly. A white area borders the black band posteriorly, remaining lateral portions of ocularium light reddish brown; mid-dorsal portion of ocularium is white with irregular, light-reddish-brown is- land. Ventral surface of opisthosoma, including genital operculum, mostly flat white, inter- rupted laterally by islands of reddish brown and medially by a very irregular light reddish- brown central figure and transverse rows of sigilla. Preanal sternite uniformly reddish brown. Coxae with broad dark brown bands separating proximal and distal white regions. Medial surfaces of stomotheca white, except for basal plate of labium, terminus of coxa- pophysis of leg II and region of coxapophysis of leg 1 bearing row of long setae. “Lips” of coxapophysis of palp and leg I and distal por- tion of labium translucent. Ovipositor: Long, dorsoventrally flattened (Fig. 6), dorsal and ventral surfaces similar, terminal apparatus with three bilaterally paired segments, shaft with 12 unpaired seg- ments. Ultimate paired segment elongate, heavily pigmented (brown), with well-devel- oped sensory organ inserted on distolateral concavity; each segment with one basal ring of six socketed setae, one distal ring of six socketed setae and three medial apical sock- eted setae. Each sensory organ domelike with about 20 seta-like projections. Penultimate paired segment heavily pigmented, each with ring of five socketed setae; segment divided medially by “lips” of ovipore. Antepenulti- mate paired segment similar to shaft segments but divided medially. Shaft segments decreas- ing in pigmentation proximally, thin longitu- dinal membranes along each lateral surface di- vides each segment into dorsal and ventral plates; dorsal and ventral plates of distal eight shaft segments and antepenultimate paired segment with two pairs of socketed setae. Shaft segments with setae also with reduced pigmentation along midline of dorsal and ven- 50 THE JOURNAL OF ARACHNOLOGY tral surfaces giving the superficial impression that the plates are divided. Apparent seminal receptacles present at level of proximal mar- gin of terminal shaft segment; obscured by dark cuticle of segment but appearing as trans- verse dark band, associated with a pair of thin- walled sacs that project proximally to the level of shaft segment 3. Adult male and immatures.' Unknown. Distribution. — Known only from the type locality. ACKNOWLEDGMENTS We thank James Cokendolpher for facilitat- ing this collaboration and for comments on the manuscript, Mark Harvey for loaning speci- mens of Hesperopilio mainae, and Charyn Micheli for the Spanish translation of the ab- stract. The work was supported by the Mary- land Agricultural Experiment Station. LITERATURE CITED Cokendolpher, J.C. & E.A. Maury. 1990. Austrop- sopilio harvestmen (Opiliones, Cyphopalpatores, Caddidae) discovered in South America. Boletin de la Sociedad de Biologia de Concepcion 61: 59-62. Forster, R.R. 1955. Further Australian harvestmen (Arachnida: Opiliones). Australian Journal of Zoology 3:354-411. Hickman, V.V. 1957. Some Tasmanian harvestmen of the sub-order Palpatores. Papers and Proceed- ings of the Royal Society of Tasmania 91:65-79. Hirst, S. 1925. On some new genera and species of Arachnida. Proceedings of the Zoological Soci- ety of London 1925:1271-1280. Shear, W.A. 1996. Hesperopilio mainae, a new ge- nus and species of harvestman from Western Australia (Opiliones: Caddidae: Acropsopilioni- nae). Records of the Western Australian Museum 17:455-460. Shultz, J.W & T. Cekalovic. 2003. First species of Austropsopilio (Opiliones: Caddoidea: Caddidae) from South America. Journal of Arachnology 3 1 : 20-27. Silvestri, E 1904. Descrizione di un nuovo genere di Opilionidi del Chile. Redia 2:254-256. Manuscript received 17 September 2004, revised 10 June 2005. 2006. The Journal of Arachnology 34:51-61 ROLE OF THE ANTERIOR LATERAL EYES OF THE WOLF SPIDER LYCOSA TARENTULA (ARANEAE, LYCOSIDAE) DURING PATH INTEGRATION Joaquin Ortega-Escobar: Department of Biological Psychology, Faculty of Psychology, University Autonoma of Madrid, 28049“Madrid, Spain. E-mail: joaquin. ortega@uam.es ABSTRACT. Spiders of the species Lycosa tarentula (Linnaeus 1758) (Araneae, Lycosidae) use a vector navigation system while homing under natural conditions. Under laboratory conditions, in the absence of information relative to the sun’s position or any pattern of polarized light, L. tarentula uses a path inte- gration system which consists of turning at a fixed angle similar to one that could carry it to its burrow. In the absence of light, the angle is random. In this study we ask whether the spiders acquire the infor- mation about the angle turned during the outward journey through the anterior lateral eyes (ALEs), whose visual fields are directed towards the ground. To answer this question, two groups of animals were studied: one group with only the ALEs covered and another group with all eyes except ALEs covered. Our results show that ALE information alone is adequate to obtain the angle at which the animal should turn when homing. Keywords: Direction estimation, spiders, optical flow Animals that are central foragers move from a central point (nest, burrow) to find food or mates. After this displacement, these animals must be able to reach that central point. Path integration (PI) is one of the most frequently used mechanisms to get it (Papi 1992). While moving, the animal measures and integrates the angles (rotations) as well as distances travelled to obtain a vector whose orientation indicates home direction and whose length indicates the distance, so that it can always take a direct path back to its start- ing point. That means that the animal does not retrace its outward journey. Information about changes of direction can be obtained in arthropods through exo skeletal sense organs (Seyfarth et al. 1982; MitteL staedt 1983; Corner & Claas 1985; Durier & Rivault 1999) or by the use of biological com- passes based on the sun or the pattern of ce- lestial polarized light (Wehner 1997; Homberg 2004; Mappes & Homberg 2004). In several insects, it has been shown that they use trans- lational image motion (optic flow) to estimate flight- or running distances (review: Sriniva- san & Zhang 2004) In particular, several stud- ies made with honeybees (Srinivasan et al. 1997) demonstrated that honeybees integrate over time the image velocity that is experi- enced during the flight and that this measure- ment is independent of image structure (Si et al. 2003). In another experiment, Ugolini (1987) displaced wasps from their nests to various sites, released them, and observed their homing trajectories. He found that they headed accurately towards their nests if they had been displaced in a transparent container but not when they had been displaced in an opaque container. In spiders, homing has been thoroughly studied in the funnel web spider Agelena la- byrinthica (Clerck 1757), which can use vi- sual cues together with tactile and propriocep- tive ones (Corner & Claas 1985). Homing has also been studied in the nocturnal ctenid spi- der Cupiennius salei (Keyserling 1877) (Sey- farth & Barth 1972; Seyfarth et al. 1982; Barth 2002). It was demonstrated that C. salei needs proprioceptive information for homing because animals that have been surgically al- tered (e.g., spiders with the lyriform slit sense organs of the femur and tibia destroyed me- chanically) returned with less success to the site from which they had been chased. In C. salei, Schmid (1997) noted differences in the kind of locomotion depending upon whether they were in bright light (normal walking movements with eight legs) or complete dark- ness (first pair of legs used as antennae). 51 52 THE JOURNAL OF ARACHNOLOGY In Lycosidae, the first studies about homing were realized in the European species Arctosa perita (Latreille 1799) (Papi 1955; Papi & Tongiorgi 1963). This species displays so- called “zonal orientation” or “orientation to Y axis,” which means that after an active or passive displacement away from the shore, they orient and move perpendicular to the shore until they reach it. Papi (1955) demon- strated that A. perita could find the shore from which it had been displaced only if the sky was not heavily overcast. Later, the contribu- tions of innate and learned components to as- tronomical orientation were analyzed by Papi & Tongiorgi (1963). Magni et al. (1964) showed that the anterior median eyes (AMEs) were primarily responsible for homing behav- ior by using celestial polarized light in A. var- iana Koch 1847. However, the structural basis for polarization sensitivity in AMEs was not found (Bacetti & Bedini 1964). The first study that discovered the structural basis for polar- ization sensitivity in Lycosidae was by Me- lamed & Trujillo-Cenoz (1966) in Lycosa er- ythrognatha Lucas 1836 (= L. raptoria Walckenaer 1837) followed by the research on L. tarentula (Linnaeus 1758) (Kovoor et al. 1993). Recently, Dacke et al. (2001) have found the same structural basis in Lycosa god- ejfroyi L. Koch 1865 and other lycosids. Lycosa tarentula is a circum-Mediterranean wolf spider that typically lives in a burrow in which the superior part is delimited by little twigs held together by silk (Ortega-Escobar 1986). The depth and diameter of the burrow is correlated with the spider’s size (Ortega-Es- cobar 1986). The prosoma of L. tarentula fe- males is variable in size: it can measure from 6. 0-9. 5 mm in width (unpub. data). The visual system of the lycosid spider Ly- cosa tarentula has been studied both from the behavioral (orientation to nest: Ortega-Esco- bar & Munoz-Cuevas 1999; Ortega-Escobar 2002a; locomotor activity rhythms: Ortega- Escobar 2002b; Ortega-Escobar et al. 1992) and structural aspects (Kovoor et al. 1992, 1993, 1999, 2005a, b). In the study by Ortega- Escobar & Munoz-Cuevas (1999), when spi- ders were under an overcast sky, they did not orient homewards; instead, they turned an al- most constant angle for PI or path integration. In an indoor study (Ortega-Escobar 2002a), individuals of L. tarentula were displaced by moving them along a two-leg trajectory with a 90° angle between legs; at the end of the outbound path, the spider was lifted and placed in an arena with its body axis oriented at random. When this procedure was carried out under illumination, the spiders showed PI by turning a constant angle and walking in search of the burrow, while in darkness (really under red light to which they are insensitive) they also showed PI but in this case they turned a random angle. Thus, it is possible that L. tarantula needs visual information about their movement (optic flow) and given the visual fields of their eyes (Land 1985), the eyes that could give more precise information about optic flow would be the anterior lateral eyes (ALEs) which look towards the ground. The aim of the present study was to check what eyes provide to L. tarentula the most reliable information about directional changes in PI in the laboratory in the absence of ce- lestial cues. In a first approach, I have ana- lyzed the contributions of anterior lateral eyes (ALEs) versus the rest of the eyes. METHODS Experimental animals.— Twelve lab- reared adult females of L. tarentula were used. They were maintained in individual containers measuring 17 X 13 X 8 cm, big enough for them to move around to dig burrows. They were fed blow flies (Calliphora vomitoria) and given water twice a week. These animals had been captured from a wild population in Madrid (central Spain; N 40° 32' W 3° 42') and went through the final 2-3 molts in the laboratory; all were close to the same age and all trials were conducted after maturation. Experimental procedure. — To begin the study of homing orientation, animals were placed in a terrarium measuring 60 X 30 X 35 cm. This terrarium had a 15 cm deep sub- stratum of soil similar to the natural substrate (Fig. 1 right); in the middle of one long side of the terrarium, an artificial burrow was built, similar to that which the spider digs in the field. After 5 days of habituation to the ter- rarium, the experiment began. During these 5 days, spiders were mostly in the burrow dur- ing the daytime and moved about during some hours at night. To displace the spiders, they were gently removed from the burrow and pushed along the edge of the terrarium on a path traversing half the length and the full width of the terrarium. When the spider ar- ORTEGA-ESCOBAR— ANTERIOR LATERAL EYES AND PATH INTEGRATION 53 Figure 1. — Left: Setup used to study homing in L. tarentula. Right, top view of terrarium in which the animal lived during the study; arrows indicate the outward path. Left, dorsal view of the arena in which the animal was left after being taken from the right corner opposite to the burrow. Burrow direction was at 350°. The big arrow indicates the transfer of the animal to the center of the arena (shown at half of its actual size in relation to the terrarium). To go to the burrow, the spider must turn an angle of 135° in its terrarium. Right: Aspect of the substratum of the terrarium. rived at the end of the path, it was placed into a transparent open glass container and trans- ferred to the center of an arena 90 cm in di- ameter (wall height, 48 cm; visual angle, 47°) (Fig. 1 left). There, if the animal turned at an angle of 135° towards the left, it would be oriented to its burrow. Both the terrarium and the arena were in a room without natural light- ing. The room was lit in the daytime (0800- 2000 h) with white light by two SYLVA- NIA® Standard F36W fluorescent tubes producing 200 lux at the floor level of the are- na. Each animal was used in 8 trials (eight control trials and eight experimental trials; see below) and placed in one of the following compass directions at random: 0°, 45°, 90°, 135°, 180°, 225°, 270°, 315°. The spider’s ori- entation was recorded when it was at a dis- tance of 20 cm from the center of the arena. If the spider had not moved during 20 minutes it was returned to the terrarium. The floor of the arena was thoroughly cleaned with ethanol before each test. All the trials were run be- tween 1 1 and 1 8 h with lights on. All spiders (ji = 12) were observed first with all eyes uncovered (control test; eight tri- als for each animal). Afterwards, spiders were assigned at random to one of two groups. One set of spiders {n — 6) had all eyes but the ALEs covered (uncovered ALEs group, ex- perimental test), while the other set {n = 6) had only the ALEs covered (covered ALEs group, experimental test). Therefore, we had two groups: “uncovered ALEs group” and “covered ALEs group” that were observed without eye covering (control test) and with eye covering (experimental test). To cover the eyes, the animals were anesthetized with ether, their legs restrained with adhesive plaster and their eyes covered by first applying a layer of collodion over the anterior region of the pro- soma; then by applying two layers of water- soluble black paint (Van Gogh); and, finally, by applying another layer of collodion. Eye occlusion was checked in each case after the completion of runs using a stereo microscope. Automated video tracking. — The image of the arena was captured by an Ikegami ICD- 42B BAV CCD video camera and displayed on a Sony® Trinitron color video monitor. Si- multaneously, the video signal was digitized by a Targa 1 frame grabber that was interfaced with a personal computer supporting an object video-tracking system (Etho-Vision, Noldus Information Technology, Wageningen, The Netherlands). The paths supplied by Etho- Vision were later digitized. The best-fitting line to a trajectory was computed by the meth- od of principal axes (Sokal & Rolf 1995). The following parameters were determined in both conditions (covered or experimental and uncovered or control eyes) in both groups: ( 1 ) topographic bearing of the digitized home- ward path when the spider crossed a virtual circle 20 cm in diameter from the starting point of the return; (2) angle (a angle) of the 54 THE JOURNAL OF ARACHNOLOGY Figure 2. — Example of a homeward path in the arena. The black square in the center represents the point where the spider is placed, and its orientation is indicated by the arrow; the black circle represents the burrow compass direction; a is the angle be- tween the initial orientation and final bearing (where homing path crosses the circle). body axis when the spider crossed the virtual circle with respect to the starting position of the body axis; as the animal could turn either clockwise or counterclockwise, the a angle was always taken counterclockwise, which was the expected direction for the animal to turn in the terrarium (Fig. 2); (3) turning di- rection (clockwise or counterclockwise). Statistical analyses. — The directions fol- lowed by the animals were analyzed as cir- cular variables according to Batschelet (1981). For first-order statistics, the Rayleigh test was used to determine whether the observed hom- ing directions from particular individuals were significantly oriented. To see if the deviation between individual significant vectors and the angle of home direction (Fig. 2) was signifi- cant we used the confidence interval for the mean angle (P < 0.05; Batschelet 1981). On the second-order level, Moore’s and Mardia- Watson-Wheeler’s tests (Batschelet 1981) were used to test directionality significance and differences in the orientation of the sub- jects between control and experimental tests respectively. The percentage of turning in the correct di- rection (counterclockwise) was analyzed by a two X two repeated measure analysis of var- iance with the type of eyes (PMEs/PLEs/ AMEs, and ALEs) and covering (control - without eye covering-, and experimental -with eye covering-) as factors. Voucher specimens have been deposited in the Museo Nacional de Ciencias Naturales (Madrid, Spain). RESULTS Homing in uncovered ALEs group (con- trol test). — Homing paths followed by the spiders in the arena were either roughly straight, finishing with a sudden turn either to the right or to the left, followed by a turn in the opposite direction, as described previously (Ortega-Escobar 2002a) or they walked until they touched the arena wall. This series of turns has also been observed when the animal is taken from the burrow without having been displaced and transferred to the center of the arena. This type of behavior, called “system- atic search” (Wehner & Wehner 1986), was not analyzed in this study. Topographic bearings: None of the six spi- ders oriented themselves towards the burrow position or towards another point of the room in a constant way in the eight trials (Fig. 3 top, left). a angle: a was non-randomly oriented in all six animals (Table 1 and Fig. 4 top, left). The mean vectors of the six animals were not randomly distributed (Moore’s test: D = 1.277, P < 0.05). Table 1 shows the mean angle and length of the vector for each animal. In two animals the turn that the spider should have made to go to the burrow in the terrarium (135°) was included in the confidence interval of the mean, but their mean vectors were sta- tistically significant (Fig. 4 top, left). Homing in uncovered ALEs group (ex- perimental test). — The homing paths in the experimental test were similar to those shown by animals when all eyes were uncovered. Topographic bearings: Only one of the six spiders oriented itself towards one point of the room in a consistent way in the eight trials. (Fig, 3 bottom, left). The other five animals oriented themselves at random. OL angle: a was non-randomly oriented in all six animals (Table 1 and Fig. 4 bottom, left). The mean vectors of the six animals were not randomly distributed (Moore’s test: D = 1.353, P < 0.05), Table 1 shows the ORTEGA-ESCOBAR— ANTERIOR LATERAL EYES AND PATH INTEGRATION 55 ALEs uncovered group ALEs covered group Experimental test Experimental test Figure 3. — Top left: Mean vectors (control test) of topographical bearings of the uncovered ALEs group. The dashed circle indicates the critical r-value of E = 0.05. 0° indicates the magnetic North. Top right: Mean vectors (control test) of topographical bearings of the covered ALEs group. Bottom left: Mean vectors (experimental test) of topographical bearings of the uncovered ALEs group. Bottom right: Mean vectors (experimental test) of topographic bearings of the covered ALEs group. mean angle and length of the vector for each animal. In four animals, the turn that the spi- der had to make to go to the burrow in the terrarium (135°) was included in the confi- dence interval of the mean. Homing in covered ALEs group (control test). — As expected, the homeward paths of these animals were very similar to those ob- served for the other group (ALEs uncovered group). Topographic bearings: None of the six spi- ders oriented itself towards the burrow position or towards another point of the room in a con- sistent way in the eight trials (Fig. 3 top, right). 56 THE JOURNAL OF ARACHNOLOGY Table 1. — a angle (mean a angle (6) and vector length (r)) in the controls and tests of both groups. Asterisks indicate the degree of significance of the first and second order data; *, R < 0.05. Covered ALEs group Uncovered ALEs group All eyes uncovered ALEs covered All eyes uncovered ALEs uncovered Individual 0 r 0 r Individual 0 r 0 r 1 159° 0.87* 61° 0.39 1 154° 0.81* 146° 0.80* 2 38 0.71* 42 0.56 2 164 0.83* 172 0.80* 3 134 0.65* 318 0.14 3 169 0.74* 158 0.91* 4 165 0.66* 51 0.27 4 165 0.95* 154 0.93* 5 161 0.87* 204 0.69* 5 195 0.91* 187 0.75* 6 60 0.68* 55 0.55 6 122 0.94* 137 0.96* Group means Mardia test 126 0.64 Rd = 14.91, 42 P < 0.05 0.51 162 0.93* Rd = 159 0.27, NS 0.96* a angle: The a angle (Fig. 4 top, right) was non-randomly oriented in all the animals of this group. The mean vectors of the six ani- mals were randomly distributed (Moore’s test: D = 0.923, P > 0.05). Table 1 shows the mean angle and length of the vector for each animal. In four animals the turn that the spider had to make to go to the burrow in the ter- rarium (135°) was included in the confidence interval of the mean. In the other two animals, the 135° value was not included in the confi- dence interval of the mean, but their mean vectors were statistically significant (Fig. 4 top, right). Homing in covered ALEs group (experi- mental test). — The homeward paths of the animals with covered ALEs were very similar to those observed when no eye was covered. However, in several animals, circular path- ways were observed (Fig. 5). These circular pathways were not used for the analysis. Topographic bearings: Only one of the six spiders oriented itself towards a point of the room in a consistent way in the eight trials (Fig. 3 bottom, right). a angle: The a angle (Fig. 4 bottom, right) was randomly oriented in all but one of the animals of this group. In this animal, this an- gle (Table 1) has a value of 204°, very differ- ent from 135°; this value was not included in its confidence interval. Comparison of ol angle in control tests between both groups. — To test if both groups have the same mean orientation in control tests we have used the Mardia- Watson- Wheel- er test. In this case, Rf = 1, NS, therefore there is no difference in the a angle between both groups. Percentage of turning in the expected di- rection (see Methods), — Turning in the ex- pected direction by the six animals of the un- covered ALEs group (control test) was 70.8 ± 6.5 % (mean ± SD), while among the six in the covered ALEs group (control test) it was 87.5 ± 19.4 % (Fig. 6). In experimental tests, 75 ± 22.4% of the animals with only the ALEs uncovered turned in the expected direction while 50 ± 22.4% of those animals with only ALEs covered turned in the expected direction. The ANOVA of the percentage of turning in the expected direction showed no effects for the two factors and significant effects for the interaction (Fi lo — 15.625, P = 0.003). DISCUSSION As in a previous study (Ortega-Escobar 2002a), the present results show that during the day L. tarentula does not orient itself to- wards the topographic burrow position in the absence of tacto-chemical information and the presence of distant visual landmarks of the laboratory. The results agree with what has been observed when the animals could use neither the sun nor the polarized light pattern for homing (Ortega-Escobar & Mufioz-Cue- vas 1999). With all the eyes uncovered, this study shows that L. tarentula tries to return home by turning a fixed angle, a, near to 135°. The turn near to 135° would let the animal walk to a point near the burrow if its orientation had not been changed in the arena. If L. tarentula used only proprioceptive in- formation for homing, it should be able to turn an a angle near to 135° when it was displaced ORTEGA-ESCOBAR— ANTERIOR LATERAL EYES AND PATH INTEGRATION 57 ALEs uncovered group ALEs covered group Control test Control test Experimental test Experimental test Figure 4. — Top left: Mean vectors (control test) of a of the uncovered ALEs group. The dashed circle indicates the critical revalue of F = 0.05. 0° indicates that the animal walks in the same direction that it has been placed in the arena. The external arrow represents the angle the spider has to turn in the terrarium to go back to the burrow (a= 135°). Top right: Mean vectors (control test) of a of the covered ALEs group. Bottom left: Mean vectors (experimental test) of a of the uncovered ALEs group. Bottom right: Mean vectors (experimental test) of a of the covered ALEs group. in the darkness. The previous study (Ortega- Escobar 2002a) had shown that this is not the case. Therefore, there must be some kind of visual information that the spider must use for homing. The present results exclude the pos- sibility of using distant visual landmarks giv- en that topographic bearings are not constant. There is another possibility to estimate the an- gle a: the use of the self-induced optic flow through some eyes. The visual field of ALEs is disposed in such a way that optic flow through them is more constant than optic flow through the other eyes (Fig. 7). As the animal walks, the distance to the ground is rather con- 58 THE JOURNAL OF ARACHNOLOGY Figure 5. — Two examples of homeward paths of two different spiders with only ALEs covered. Black square represents the initial position of the spider; arrow represents the initial body direction; black circle represents burrow direction. stant and they mainly image the ground on which the animal walks. On the other hand, the other eyes image different objects and the optic flow is more complex in relation to the distance to the eyes. It is proposed that L. tar- entula is able to perceive the optic flow of the natural soil of the terrarium where it is dis- placed, and afterwards it uses this information to turn in the correct direction and angle even if it is placed over an unstructured substrate such as the white substratum of the arena. Are the ALEs able to discriminate the small peb- bles of the terrarium soil? One measure of the resolution capacity of a simple eye is the sam- pling frequency, Ug (Land & Nilsson 2002) such that Ug — 1/ (2AT>) where A# is the inter- receptor angle. Using the values obtained by Kovoor & Mufioz-Cuevas (1996/1997) for L. tarentula female, for the ALEs “ 0.30 cy- cles/degree which means that a grating con- sisting of two stripes, one black and one white, will occupy 3.33° of the visual field. p--DZ Glenogmtha Pachygnatha Argiope Epe'mtypus UnypMa Pima Steatoda 1 /55 L_.WesffCIIS Phonogmtha Clitaeto Nephiia 3 1 8S I — We/eflw'a /esi— Nephilengys Azilia 2/-L___.OofeliogMia Meta €h!ysometa Metellm S. Uineatus S. acofeemis Tetmgmtha Glenogmtha 3 /S4L— Pachygnatha Figure 16.= — Strict consensus of the six shortest cladograms (unweighted parsimony analysis). Family and subfamily names follow Hormiga et al. (1995). Sancus bilineatus and S. acoreensis form a clade sister to tetragnathines. Figure 17,- — ^Preferred tetragnathid phylogeny with Sancus (a single successively weighted cladogram identical to one of the six fundamental cladograms). Branch support values given as Bremer/bootstrap, reported for values 1 and more for Bremer and for 50% and more for bootstrap. apophysis (Fig. 14) and the tibial length, which is 1.7 times as long as wide. Description.— (paratype): Total length 3.9. Cephalothorax 1.56 long, 1.14 wide, 0.55 high; dark brown. Sternum 0.79 long, 0.78 wide; light brown, darker at mar- gins. Abdomen 1.36 long, 1.66 wide, 1.7 high; dark gray covered with silver and golden spots (Fig. 5). AME diameter 0.10. PME 0.09, ALE 0.08, PLE 0.08. AME separation 0.10, PME separation 0.10. AME- ALE separation 0.08. PME-PLE separation 0.12. Clypeus height 0.10. Legs light brown. Leg I length 8.7, Leg II 7.0, Leg III 3.8, Leg IV 5.6, pedipalp length 1.9. Epigyeum as in Figs. 8, 9: Anterior de- pression 0.8 times as wide as the transverse bar. Epigynal caudal plate wide and deeply notched (Fig. 9). Male (paratype): Total length 2.98. Ceph- alothorax 1.29 long, 0.90 wide, 0.44 high; col- or as in S. bilineatus. Sternum 0.7 long, 0.64 wide; color as in female. Abdomen 1.72 long, 1,0 wide, 1.04 high; dark gray covered with silvery spots and two longitudinal white-gold- en lines. AME diameter 0.09. PME 0.07, ALE 0.07, PLE 0.08. AME separation 0.09, PME separation 0.1. AME- ALE separation 0.12. PME-PLE separation 0.11. Clypeus height 0.09. Cheliceral teeth and leg pigmentation as in female. Leg I length 8.7, Leg II 6.7, Leg III 3.4, Leg IV 5.4, pedipalp length 1.6. Ped- ipalp as in S. bilineatus except for the diag- nostic characters (see above). Variation (from Wunderlich 1992). — Fe- male total length ranges from 3. 8-4. 3, ceph- alothorax length from 1.45-1.55. Male total length from 2. 5-3. 2, cephalothorax length from L2-L5. Distribution.— Azores (Fig. 15): Sao Mi- guel, Santa Maria, Fajal, Pico, Terceira, Flores (Wunderlich 1992). Ecology. — In Azores the spiders live in 122 THE JOURNAL OF ARACHNOLOGY ACCTRAN Meta — Chiysometa Metellina 25(1) - Leucauge 7(0) 61(1) 62(1) 63(1) 7(1) 21(1) 25(1) 40(1) 41(1) Sancus bilineatus Sancus acoreensis Tetragnatha Glenognatha Pachygnatha DELTRAN 25(2) Meta Chiysometa Meteliina 7(0) 25(1) - Leucauge 21(1) 25(1) 61(1)62(1)63(1) S. bilineatus S. acoreensis 7(0) 40(1) 41(1) Tetragnatha — — Glenognatha — — Pachygnatha Figure 18. — Alternative optimizations of Sancus synapomorphies and other relevant characters and states (in parentheses) on the preferred phylogeny. Bolded are non-homoplasious characters. Delayed transfor- mation (DELTRAN) is the preferred and more logical optimization (see text for details). sunny to shaded upper vegetation layers near lakes (Wunderlich 1992:360). They were col- lected by beating vegetation and thus their webs are not known (Wunderlich in litt.). PHYLOGENETICS Heuristic searches in NONA produced two trees of minimal length under amb-, repre- senting a subset of six minimal length trees in NONA under amb=^ and in PAUR All mini- mal length topologies (length 136, Cl = 0.56, RI = 0.72) have in common the placement of Sancus within Tetragnathidae, as well as the monophyly of Sancus and of Tetragnathidae. These results are congruent with those of Hor- miga et al. (1995). Figure 16 shows the strict consensus of the six shortest trees. Sancus is recovered as sister to the tetragnathine clade (as defined by Hormiga et al. 1995) contain- ing the genera Tetragnatha Latreille 1804, Glenognatha Simon 1887 and Pachygnatha Sundevall 1823. The six trees conflict in the placement of Leucauge White 1841, which is sister either to Sancus + tetragnathines (Fig. 17) or sister to Meta C.L. Koch 1836, MeteT Una Chamberlin & Ivie 1941 and Chrysometa Simon 1894, as well as in the position of Epeirotypus O. P.-Cambridge 1894 relative to the sheet- web builders. Successive weighting resulted in one stable topology after a single iteration, identical to one of the most parsi- monious cladograms under equal weights (Fig. 17). Bremer support and bootstrap sup- port values are mapped on this preferred phy- logeny. The diagnostic characters and synapomor- phies (Fig. 18) of the genus Sancus (under DELTRAN optimization; see justification be- low) are the CBP apical denticles (61/1), the epigynal transverse bar (62/1) and the epigyn- al ventral depression (63/1). Another diagnos- tic character, lack of dorsal femoral tricho- bothria (7/1) serves as synapomorphy under ACCTRAN (Fig. 18). In addition, unambig- uous synapomorphies of Sancus (homopla- sioLis on the cladogram) are mesal cymbium KUNTNER AND ALVAREZ-PADILLA— SYSTEMATICS OF SANCUS 123 orientation (21/1) and procurved CBP (25/1) (see Character analysis). DISCUSSION Monotypic genera are problematic because they contain no grouping information and therefore are not phylogenetic hypotheses (Zujko^Miller 1999). Platnick (1976, 1977) also argued that monophyly cannot apply to monotypic genera. In some cases, taxonomists have no choice but to retain monotypic genera (e.g. Kunteer 2002), for example if the sister species or clade is unknown or unresolved. By synonymizing Leucognatha with Sancus we rid tetragnathid systematics of two monotypic genera. Alternative optimizations and synapo- morpMes. — Delayed transformation (BEL- TRAN) optimizes three out of four diagnostic Sancus characters as syeapomorphies for the genus (Fig. 18). The accelerated transforma- tion alternative (ACCTRAN) optimizes the CBP denticles (character 61) as a synapomor- phy for Leucauge + (Sancus + tetragnathines) (Fig, 18). However, the CBP itself (character 25) is primitively absent at this node. Since the CBP denticles are an attribute of the CBP, any optimization in which the denticles arise before the process is illogical, an artifact re- sulting from, the inapplicable coding of the CBP denticles (character 61) in Leucauge and tetragnathines, which lack the CBP. Since the presence of the CBP is an unambiguous syn- apomorphy of Sancus, the DELTRAN alter- native is more reasonable, implying the evo- lution of the CBP denticles (along with the CBP) in the common ancestor of Sancus. ACCTRAN optimizes the two new epigyn- al characters (62, 63) as synapomorphies of Sancus + Tetragnathinae. However, tetragna- thines are haplogyne, meaning they lack the epigynum. (40/1) and fertilization ducts (41/1), both unambiguous syeapomorphies of the clade (Fig. 18). Thus, for tetragnathines, the two new epigynal characters are inapplicable. The ACCTRAN optimization implicitly as- sumes tetragnathine ancestor had the Sancus epigynal characters but lost them (along with the epigynum itself), an unwarranted pre- sumption. In this case, DELTRAN is a simpler explanation of the data. The presence of dorsal femoral trichoboth- ria (character 7/0) served as a synapomorphy for Leucauge + tetragnathines in Hormiga et al. (1995). In this analysis the optimization of this homoplasious character is ambiguous (Fig. 18), ACCTRAN resolves the presence of trichobothria as a synapomorphy for Leucauge + (Sancus + tetragnathines) and the absence as a synapomorphy of Sancus. On the other hand, DELTRAN favors two separate origins (Fig. 18) and thus implies that trichobothria in Leucauge may not be homologous to the ones in tetragnathines. Phylogenetic placement with comments on tetragnathid relationships. — This paper establishes the phylogenetic placement of Sancus, not new phylogenetic relationships of the tetragnathid genera. The preferred phytog- eny (Fig. 17) agrees with the phytogeny found by Hormiga et al. (1995), and Sancus groups with tetragnathines. Of course, we basically re-ran the Hormiga et al. (1995) data, so such congruence is not surprising, even though we think some homology statements should be re- assessed. We will present these new hypoth- eses in future papers on nephiline and metine systematics. Three unambiguous synapomorphies sup- port the group Sancus + Tetragnathinae: 1) long and finger-like paracymbium (Figs. 11- 14); 2) presence of an anterior paracymbial apophysis (Figs, 13, 14); 3) spiraled reservoir course (homoplasious). One unambiguous but weak synapomorphy supports Leucauge + (Sancus + tetragnathines): posterior gut caeca (character 11 of Hormiga et al. 1995), but we did not score the feature for Sancus because specimens are too rare to dissect. The tetragnathid phylogeny, as currently understood (Fig. 17), must be considered pre- liminary and interpreted cautiously. Hormiga et al. (1995) did not present branch support statistics, but most nodes are poorly supported (Fig. 17; Bremer = 1, bootstrap < 50%). Bootstrapping collapsed 11 out of 20 nodes and tetragnathid monophyly collapsed. Ne- philinae, especially distal nephilines (Clitaetra (Nephila (Herennia + Nephilengys), and Te- tragnathinae are well supported (also in Hor- miga et al. 1995). On the other hand, current work disputes the placement of the nephiline clade as tetragnathids (Kuntner 2003; Kuntner 2005, 2006a & b) and some genera, tradition- ally classified as nephilines, have been trans- ferred to Araneidae (Kuntner 2002; Kuntner & Hormiga 2002). Wunderlich (1992:359) placed Leucognatha 124 THE JOURNAL OF ARACHNOLOGY (= Sancus) in Leucauginae, but did not pro- vide synapomorphies for the subfamily. All Leucauge species possess characteristic rows of fourth femoral trichobothria (Levi 1980: figs. 50, 51, 67), Similar condition can be found in tetragnathid genera Opadometa Ar- cher 1951, Tylorida Simon 1894, Mesida Kul- czynski 1911 and Orsinome Thorell 1890 (none of them placed phylogenetically), but not in Sancus. Femoral trichobothria of tetrag- nathines, though present, are not in rows and may not be homologous to the Leucauge con- dition (Fig. 18; see above). We will test and discuss homology of femoral trichobothria in Leucauge and tetragnathines and possible monophyly of Teucaugines' and ‘metines’ elsewhere. Behavior. — Sancus behavior and web ar- chitecture are unknown. Our prediction based on the phylogenetic outcome is that Sancus builds orb webs with an open hub and few radii, which are more horizontal than vertical. Leucauge and most Tetragnatha species build such webs (e.g. Levi 1980; own data). Sancus acoreensis was collected adjacent to bodies of water (Wunderlich 1992), which is typical for Tetragnatha. Biogeography. — Sancus is now known from the Azores in the Atlantic Ocean and two mountain peaks (Kilimanjaro and Mt. Kenya) in equatorial eastern Africa (Fig. 15). The two areas are more than 7,500 km apart, in very different climatic regimes, latitudes and ele- vations, and are habitat islands. The Azores are 1,370 km from Europe and 1,530 km from Africa. The type series of S. bilineatus says 3,000 m on Kilimanjaro; the other collection simply says Mt. Kenya. We are not aware of any other comparable taxon distribution. This unusual distribution is probably an un- dersampling artifact. However, we tried but failed to find more Sancus material in African collections. Sancus (= Leucognatha) is ap- parently also absent from Madeira and the Ca- nary Islands (Wunderlich 1992: 359; see also Fig. 15), which lie between the Azores and the mainland Africa; nor does the genus occur in the Mediterranean. If not artifactual, the distribution might be explained either by extinction of Sancus in in- tervening Africa or dispersal and divergence into the two clearly diagnosable species we see today. An undiscovered African popula- tion of S. acoreensis might also exist and have been introduced to the Azores. We expect f more records of Sancus in the future from Af- : rica, Macaronesia, and perhaps from the Med- iterranean, and hope this paper will facilitate such discoveries. ACKNOWLEDGMENTS We thank Jonathan Coddington, Gustavo 1 Hormiga, Jeremy Miller and Ingi Agnarsson for valuable help and comments to an early draft. Mark Harvey, Peter Cranston, Rudy ; Jocque, Lara Lopardo and three anonymous ; reviewers also much improved our paper. Jorg Wunderlich kindly shared unpublished data ; and provided useful comments. Jeremy Miller helped with the distribution map illustrations. Torbjorn Kronestedt (SMNH) and Jorg Wun- derlich kindly loaned or donated the speci- mens for this study. Further curatorial help came from Jonathan Coddington and Dana deRoche at USNM, Charles Griswold, Darrel Ubick and Diana Silva at CAS, Norman Plat- nick, Randy Mercurio and Lou Sorkin at AMNH, Jason Dunlop at ZMB, Rudy Jocque at RMCA, Janet Beccaloni at BMNH and Christine Rollard at MNHN. This project was supported by U.S. National Science Founda- tion grants DEB-9712353 and DEB-0328644, and collection study grants from CAS and AMNH. Alvarez-Padilla has been supported by a doctoral fellowship from CONACYT (Consejo Nacional de Ciencia y Tecnologia, Mexico). We further acknowledge the finan- cial and logistical support of the George Washington University and the Smithsonian Institution. LITERATURE CITED Boeck, A. 1872. 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Manuscript received 11 May 2004, revised 16 July 2004. 2006. The Journal of Arachnology 34:126-134 THREE NEW SPECIES OF PHOLCUS (ARANEAE, PHOLCIDAE) FROM THE CANARY ISLANDS WITH NOTES ON THE GENUS PHOLCUS IN THE ARCHIPELAGO Dimitar Dimitrov* and Carles Ribera: Departament de Biologia Animal, Universitat de Barcelona, Av. Diagonal, 645, Barceiona-08028, Spain. E-mail: ddimitrov@ub.edu ABSTRACT. Over the last decade, numerous papers focusing on the fauna of the Canary Islands have reported that many spectacular species radiations have taken place, leading to a very high level of ende- micity in this archipelago. The species of the genus Pholcus are a very good example of such a fascinating process. The Canary Islands harbor the highest number of endemic species of this genus. Therefore, in order to obtain a detailed picture of the diversity and the phylogeny of the Canarian Pholcus, a complete taxonomic revision is required. The present work is the second contribution to achieve this goal. Three new species of Pholcus are described: Pholcus bimbache, P. anachoreta and P. corniger. The first endemic species of Pholcus from El Hierro {P. bimbache) is reported; P. anachoreta is the only Pholcus species found on the Montana Clara Islet; and P. corniger is the second and most troglomorphic species known from Tenerife. Keywords: Araneae, Pholcidae, Pholcus new species, taxonomy, Canary Islands The Canary Islands are situated about 100 km off the northwestern coast of Africa. This volcanic archipelago was formed during var- ious volcanic episodes and is nowadays com- posed of seven main islands and several islets. All of them are situated almost on a straight line with an east-west orientation, with the age of the islands decreasing towards the east. The estimated ages of the islands are: Fuerteven- tura 20-22 My, Lanzarote 15-19 My, Gran Canaria 14-16 My, Tenerife 11.6-14 My, La Gomera 10-12 My, La Palma 1.6-2 My and El Hierro 0.8-1 My (Anguita & Hernan 1975; Ancochea et al. 1990; Coello et ai. 1992). The older islands, Fuerteventura and Lan- zarote, are lower in elevation due to the ef- fects of erosion. As a result of their low height they receive less moisture from the northeast trade winds than the other, higher islands. This, and the proximity of the Sahara Desert, renders them the driest islands in the archi- pelago, with most of their habitats being dry lowlands. The remainder of the Canary Is- lands have higher mountains, reaching an el- evation of 37 1 7 m (Teide, Tenerife). This high ' Current address: George Washington University, Department of Biological Sciences, 2023 G. Street, NW, Washington, DC, 20052. E-mail: dimitard@ gww.edu elevation combined with the trade winds (hu- mid from the northeast and dry from the northwest), causes a thermic inversion that forms a cloud belt between 600 and 1000 m. These clouds are almost permanent on the northern slopes, favoring the growth of a char- acteristic subtropical forest named laurel for- est. Differences in humidity and elevation be- tween and within islands are the main reasons for the development of a large variety of hab- itats. The so-called hypogean environment also contributes to changes in the diversity of habitats. In the case of the Canaries it is formed by lava tubes and the MSS (mesocav- ereous shallow stratum) (Oromi et al. 1986; Medina 1991). This high diversity of ecolog- ical niches and the initial emptiness of habitats provide the best conditions for species radia- tions. The spider genus Pholcus Walckeeaer 1 805 is a good example of this process. The 114 species that it comprises are distributed almost all around the world. However, it is interesting to note that there are no indigenous Pholcus species in Central and South America and only a few are known from North America. Before the present study, eighteen species of Pholcus had been reported from the Canary 126 DIMITROV & RIBERA— THREE NEW CANARIAN PHOLCUS SPECIES 127 Islands. This represents more than 15% of the total number of species of this genus. If we add to them the five species from Madeira (another Macaronesian island), this ratio reaches more than 19%. At the same time, the area of these islands represents an extremely small part of the total area of the generic dis- tribution, providing clear evidence of a spe- cies radiation on the Canary Islands. Apart from the cosmopolitan P. phalan- gioides (Fuesslin 1775), the remaining species of Pholcus recorded from the Canary Islands are endemic to this archipelago. Two of them have been collected from more than one is- land: P. ornatus Boesenberg 1895 has the broadest distribution, occurring on all the is- lands except Lanzarote and Fuerteveetura; and P. fuerteventurensis Wunderlich 1991 is found on Fuerteventura and Gran Canaria. Pholcus knoeseli Wunderlich 1991, P. malpaisensis Wunderlich 1991, P. mascaensis Wunderlich 1987, P. baldiosensis Wunderlich 1991, R ro- quensis Wunderlich 1991, P. intricatus Dim- itrov & Ribera 2003 and P. tenerifensis Wun- derlich 1987 are endemic to the island of Tenerife. Pholcus multidentatus Wunderlich 1987, P. calcar Wunderlich 1987, R corcho Wunderlich 1987, R edentatus Campos & Wunderlich 1995 and R helenae Wunderlich 1987 are known only from the island of Gran Canaria, while R gomerae Wunderlich 1980, R gomeroides Wunderlich 1987 and R sveni Wunderlich 1987 are endemic to La Gomera. In the present work, three new endemic species of Pholcus are described. Pholcus bimbache is the first endemic Phoclus species from El Hierro; R anachoreta is endemic to the Montana Clara islet and R corniger is the second and most troglomorphic species of this genus known from this archipelago. With these three new species the number of Can- arian endemic species of Pholcus reaches twenty, eighteen of which are mono-insular endemics, indicating that the genus has a higher diversity in the Canary Islands than previously suspected. Pholcus corniger is the most troglomorphic species of Pholcus known from the Canaries. While in R baldiosensis, the other troglo- morphic species, the reduction of the eyes is incomplete, in R corniger they are totally ab- sent. This species, unfortunately, may be ex- tinct due to the destruction of its habitat in Cueva de San Miguel, where residual waters are thrown out of houses nearby. This seems to be still a common practice in the Canaries, affecting numerous volcanic tubes and small caves. Estimating how many species suffer from this particular activity and how many are brought to extinction will be a difficult task. Taking into account the high vulnerability of both cave faunas and island ecosystems, Can- arian authorities and the Spanish government should implement more active and efficient measures to eliminate these type of activities. METHODS Specimens were examined under a Wild Heerbrugg (12-lOOX) stereomicroscope. The female vulva was removed and treated with 50% solution of lactic acid in order to render the remaining soft tissues transparent. After observation and drawing the vulva were washed in distilled water and stored in 70% ethyl alcohol. All measurements are in milli- meters. The total body length is the sum of the prosoma and the opisthosoma omitting the pedicel. The specimens are deposited in the Departament de Biologia Animal, Universitat de Barcelona (CCRUB). The numbers of the collection are given in brackets. TAXONOMY Family Phoicidae C.L. Koch 1851 Genus Pholcus Walckenaer 1805 Pholcus bimbache new species Figs. 1-9 Material examined.— Holotype male, Cue- va del Juaclo de las Moleras, Froetera, El Hierro, Canary Islands, 27°43'N, 18°08'W, 7 November 1991, C. Ribera (CCRUB 3523- 140). Paratypes: Canary Islands: 1 female, same locality and date as holotype, C. Ribera (CCRUB 3524-140) (drawings and descrip- tion of the female are based on this specimen); 1 male, 2 females and 8 juveniles (CCRUB 3522, 3525 to 3527-140) same locality and date as holotype; 1 male and 1 juvenile, same locality, 4 February 2000, N. Mercader & E. Munoz (CCRUB 4505-170). Etymology. — The species is named after the original inhabitants of El Hierro island, the so-called “Bimbaches”. Diagnosis. — Pholcus bimbache can be dis- tinguished from similar Canarian species (R sveni and R gomerae) by the less pronounced callosity of the procursus, the narrower base 128 THE JOURNAL OF ARACHNOLOGY Figures 1-9. — Pholcus bimbache new species: 1. Male palp, prolateral view; 2. Male palp, retrolateral view; 3. Male palp, frontal view; 4. Male chelicerae; 5. Uncus; 6. Trochanter of the male palp; 7. Epi- gynum, ventral view; 8. Vulva, dorsal view; 9. Epigynum, lateral view. Scale 0.2 mm. of the uncus (Fig. 5), the longer claw=-shaped apophysis of the appendix and the long, al- most straight trochanteral apophysis (Fig. 6) of the male palp (Figs. 1-3); also, by the shape of the apophyses of the male chelicerae (Fig. 4). The diagnostic characters of the fe- male are the shape of the epigynum and the large oval pore plates of the vulva (Figs. 7- 9). Description. — Male (holotype): Prosoma yellowish with well marked cephalothoracic junction and fovea. Ocular area elevated. Tho- rax with brown marking, wider than long, which starts at the fovea and extends to the posterior margin of the prosoma. It has three lighter zones dividing it into four darker radial lobes. Sternum brown-yellowish with borders slightly darker brown. Distance between AME equal to their diameter. Distance AME-ALE slightly more than two times the diameter of AME; AME-PME three times the diameter of AME. Anterior eye line frontal view slightly i DIMITROV & RIBERA— THREE NEW CANARIAN PHOLCUS SPECIES 129 recurved. Posterior eye line dorsal view re- curved. Clypeus high with yellowish color. Chelicerae (Fig. 4) yellow-brownish; chelic- eral apophyses brownish with cylindrical shape finishing with small darker outgrowths; upper margin of the proximolateral apophyses does not reach the lower margin of the frontal prominence. A few dark bristles are placed near the base of the cheliceral apophyses. Palps (Figs. 1-3) with, yellow-brownish color, trochanter with long retrolateral apophysis (Fig. 6), femur large with ventral bulge, pro- cursus with dark process of the apical apoph- ysis. Opisthosoma elongated, almost cylindri- cal, whitish with small darker transversal zone in the genital area. Female (paratype): All characters as in male except: less elevated ocular area, dis- tance between AME slightly less than their diameter, distance AME-ALE slightly less than two times the diameter of the AME, AME-PME two and half times the diameter of the AME. Chelicerae without apophyses. Genital zone without pigmentation except the sclerotized zone of the epigynum. By trans- parence some parts of the vulva can be ob- served. Epigynum and vulva as in (Figs. 7- 9). Measurements. — Male (holotype): Proso- ma 1.2 (1.2) wide, 1.3 (1.3) long; opisthosoma 1.1 (1.5) wide and 2.5 (3.0) long. Total body length 3.8 (4.3). Legs: I, femur 8. 7(9. 2), pa- tella 0.5(0. 5), tibia 8. 1(9.5), metatarsus 13.2(14.3), tarsus 2.0(2.2), total 32.5(37.5); II 6.5(7.0), 0.5(0.5), 6.0(6.5), 9.0(8.5), 1.3(1.5), 23.3(24.0); III 5.0(5.0), 0.5(0.5), 4.0(4.2), 6.2(7.0), l.O(l.O), 16.7(17.7); IV 6.3(7.0), 0.5(0.5), 5. 8(6. 2), 8.0(9.0), 1.2(1. 1) 21.8(23.9). In brackets male paratype no. 3522-140. Palp: femur 0.60, patella 0.18, tibia 0.50, tarsus 0.20, total 1.48. Procursus 0.8. Female: Prosoma 1.3 wide, 1.2 long; opis- thosoma 1.5 wide, 3.5 long; total body length 4.7. Legs: I, femur 9.0, patella 0.7, tibia 8.2, metatarus 14.0, tarsus 1.2, total 33.1; II 6.5, 0.7, 8.6, 9.2, 1.2, 23.6; III 5.0, 0.7, 4.5, 6.5, 1.0, 17.7; IV 7.0,- 0.7, 5.0, 9.0, 1.2, 22.9. Palp femur 0.40, patella 0.14, tibia 0.19, tarsus 0.3, total 1.03. Distribution. — This species is endemic to El Hierro, and is only known from the type locality. Remarks. — Pholcus bimbache appears to be related to members of the so-called Tener- ifensis group (Wunderlich 1987, 1991; Dimi- trov & Ribera 2003) composed of ten species (seven in Tenerife and three in Gomera). Here we should note that the term “Tenerifensis group” is used as merely descriptive and does not imply any phylogenetic relationship. All these species are characterized by the claw- shaped apophysis of the appendix, and by the shape of both the uncus and the lamella of the procursus. Pholcus sveni Wunderlich 1987 is the most similar species. Pholcus bimbache can be distinguished from it by the longer and more curved claw-shaped apophysis, the shape of the procursus and the morphology of both the epigynum and the vulva. The pres- ence of this species on El Hierro Island em- phasizes the close faunistic relationships be- tween Tenerife, La Gomera and El Hierro. Pholcus anachoreta new species Figs. 10-18 Material examined. — Holotype male, Montana Clara islet, Canary Islands, 29°18'N, 13°3LW, 24 April 1994, C. Ribera (CCRUB 2459-99). Paratypes: Canary Islands: 1 male, 1 female, same locality, 23-27 November 2002, A.J. Perez (CCRUB 4502, 4503-170); 1 juvenile, from the same locality, 27 January 2002, P. Oromi (CCRUB 4504-170). Etymology. — The name comes from the Greek word “anachoretes” meaning person who lives in a lonely place dedicated to con- templation. This word was adopted in the Lat- in as anachoreta-ae keeping the male gender. Named after the remoteness and isolation of the type locality. Diagnosis. — Pholcus anachoreta is easily distinguished from canarian congeners by the serrated keel of the uncus (Fig. 15), the mor- phology of the apex of the procursus (number and shape of the apophyses and lamellas) and the cheliceral apophyses (Fig. 13). The female can be distinguished from the most similar canarian species (P. fuerteventurensis and P. edentatus) by the lower sclerotized plate of the epigynum (Figs. 16, 17) and the more arced ridges of the valve (Fig. 18). Description. — Male (holotype): Prosoma whitish without a clearly marked fovea and practically indistinguishable cephalothoracic junction. The prosoma does not carry hairs except for its borders and the intraocular area. Ocular area elevated. Sternum with a whitish coloring. Clypeus high and whitish. Chelic- 130 THE JOURNAL OF ARACHNOLOGY I i Figures 10-18. — Pholcus anachoreta new species: 10. Male palp, prolateral view; 11. Male palp, frontal view; 12. Male palp, retrolateral view; 13. Male chelicerae; 14. Trochanter of the male palp; 15. Uncus; 16. Epigynum, lateral view; 17. Epigynum, ventral view; 18. Vulva, dorsal view. Scale bar 0.2 mm. erae (Fig. 13) whitish; cheliceral apophyses darker with conical shape and group of 2-3 thick bristles placed near the base. The prox- imolateral apophyses (proximal teeth) and the frontal prominence show the same coloring as the rest of the chelicerae. The upper margin of the proximolateral apophyses is higher than the lower margin of the frontal prominence. Distance between AME less than their diam- eter. The rest of the eyes situated in two ele- vated triads. AME around 50% of the size of the other eyes. Anterior eye line frontal view slightly recurved. Posterior eye line dorsal view recurved. Palps as in Figs. 10-12. Tro- chanter (Fig. 14) with long curved retrolateral apophysis, femur with ventral bulge, procur- sus very complex with many apical lamellae and with three distal dorsal spines. Uncus (Fig. 15) very characteristic with serrated keel. Opisthosoma elongated and cylindrical with whitish color, dorsally darker. A longi- tudinal zone with darker pigmentation starting from the genital area and followed by two tear-shaped spots is observed ventrally. Female (paratype): Prosoma: all characters as in male except for the cheliceral apophyses, which are absent. Sizes and distribution of the eyes as in the male but the elevation of the DIMITROV & RIBERA-THREE NEW CANARIAN PHOLCUS SPECIES 131 ocular area is less conspicuous. Opisthosoma cylindrical with yellowish coloring. The gen- ital zone is darker, with brownish pigmenta- tion. Dorsally with two parallel lines of dark spots. The whole opisthosoma is covered with short and regularly distributed hairs. Epigyn- um and vulva as in Figs. 16-18. Measurements.— Male (type): Prosoma 1.2 wide, 1.0 long; opisthosoma 1.0 wide, 3.1 long; total body length 4.1. Legs: I, femur 9.5, patella 0.4, tibia 12.0, metatarsus 16.0, tarsus 2.0, total 39.9; II 6.2, 0.4, 6.0, 10.0, 1.0, 23.6; III 5.0, 0.4, 4.2, 7.0, 1.0, 17.6; IV 7.0, 0.4, 6.0, 6.5, 0.8, 20.7. Palp femur 0.8, patella 0.2, tibia 0.7, tarsus 0.3, total 2.0. Procursus 0.75 Female: Prosoma 1.9 wide, 1.5 long; opis- thosoma 1.8 wide, 4.0 long; total body length 5.5. Legs: I, femur 8.3, patella 0.5, tibia 8.0, metatarsus 12.9, tarsus 1.9, total 31.6; II 6.4, 0.5, 5.6, 9.6, 1.5, 23.6; III 4.9, 0.5, 4.9, 7.3, 0.9, 18.5; IV 6.8, 0.5, 5.6, 8.8, 1.7, 23.4. Palp femur 0.34, patella 0.10, tibia 0.24, tarsus 0.24, total 0.92. Distribution.— This species appears to be endemic to Montana Clara and is known only from type locality, although it might occur in the neighboring islets of Graciosa and Ale- graeza and in Lanzarote island considering their geographical vicinity. Remarks.' — -The structure of the procursus of this species is similar to that of P. edentatus Campos & Wunderlich 1995 and P. fuertev- enturensis Wunderlich 1992. Similar finger- like lamellae in the procursus allow the three species to be distinguished from the rest of the Canarian Pholcus. Despite of this remarkable similarity the procursus and the uncus are very different and therefore very useful for specific identification. Pholcus cormiger new species Figs. 19-32 Material examined.— Holotype male, Cue- va de San Miguel, San Miguel de Abona, Ten- erife, Canary Islands, 28°06'N, 16°36'W, 1 January 1991, P. Oromi (CCRUB 4500-170). Paratype: Canary Islands: 1 female from the same locality, 1 January 1991, P. Oromi (CCRUB 4501-170). Etymology,— The specific name refers to the shape of the elevated ocular area remind- ing horns. The word “corniger” in Latin means “with horns”. Diagnosis. — Pholcus corniger can be easily distinguished from the rest of canariae PhoF cus species by the simplified structure of the procursus with single membranous lamella on its apex, the presence of six teeth and the ab- sence of basal bristles on the cheliceral apoph- yses (Fig. 26). The female is differentiated by the elevated conically shaped epigyeum (Figs. 27-29). This species can be easily distin- guished by the total reduction of the eyes (both in male and female) and the shape of the elevated ocular area that reminds horns (Figs. 19-22). Description.— Male (holotype): Prosoma with an ochre-yellow coloring and well- marked fovea and cephalic furrow. Eye area elevated with characteristic shape and brown coloring (Figs. 19-20), with group of hairs be- tween the ocular elevations. Clypeus high, yellowish with brownish spot in the center and almost transparent at the edges. Sternum yel- lowish. Chelicerae yellow-brownish, with brown apophyses carrying six dark brown teeth without bristles at the base (Fig. 26). Frontal prominence small. The upper margin of the proximolateral apophyses roughly reaching the lower margin of the frontal prom- inence. Eyes completely missing. Palps as in Figs. 23-25. Uncus as in Fig. 31. Trochanter (Fig. 32) with retrolateral apophysis, femur with ventral bulge. The procursus is very characteristic, conspicuously different from those of the remaining species of Canariae Pholcus. While in all the other species the api- cal part of the procursus is very complex and carries one or various apophyses and lamellar processes, in P corniger it is much simpler and ends with a single membranous lamella. This lamella extends along all the apex of the procursus. Opisthosoma cylindrical with pale yellowish color. Female (paratype): Like the male, although the elevations of the ocular area are much smaller and colored like the rest of the pro- soma (Figs. 21, 22). Prosoma lighter. Clypeus almost transparent. The genital area is not pig- mented except for the sclerotized parts of the epigynum. Epigynum and vulva as in Figs. 27-30. As in the male, eyes are absent. Measurements.— Male (holotype): Proso- ma 1.1 wide and 1.0 long; opisthosoma 1.0 wide and 2.5 long. Total body length 3.5. Legs: I, femur 7.0, patella 0.3, tibia 7.0, meta- tarsus 11.5, tarsus 1.3, total 27.1; II 5.5, 0.3, 5.2, 8.0, 1.2, 20.2; III 4.0, 0.3, 3.9, 5.3, 1.0, Figures 19-26, — Pholcus corniger new species: 19. Male prosoma, frontal view; 20. Male prosoma, lateral view; 21. Female prosoma, frontal view; 22. Female prosoma, lateral view; 23. Male palp, frontal view; 24. Male palp, prolateral view; 25. Male palp, retrolateral view; 26. Male chelicerae. Scale bar 0.2 mm. 132 THE JOURNAL OF ARACHNOLOGY DIMITROV & RIBERA— THREE NEW CANARIAN PHOLCUS SPECIES 133 Figures 27—32. — Pholcus cornigern&w species: 27. Epigynum, lateral view; 28. Epigynum, caudal view; 29. Epigynum, ventral view; 30. Vulva, dorsal view; 31, Uncus; 32. Trochanter of the male palp. Scale bar 0.2 mm. 145; IV 6.5, 0.3, 5.0, 7.0, LO, 19.8. Palp fe- mur 0.50, patella 0.10, tibia 0.50, tarsus 0.28, total 1.38. Procurses 0.4 Female: Prosoma 1.1 wide, LO long; opis- thosoma 1.1 wide, 2,2 long; total body length 3.2. Legs: I, fem.er 7.0, patella 0.3, tibia 6.5, metatarsus 11.0, tarsus 1.8, total 26.6; II 6.2, 0.3, 5.0, 7.2, 1.1, 19.8; III 4.0, 0.3, 3.8, 5.0, 0.8, 13.9; IV 6.0, 0.3, 5.0, 7.0, 1.0, 19.3. Palp femur 0.24, patella 0.07, tibia 0.19, tarsus 0.30, total 0.80. Distribution.— Endemic to the island of Tenerife, only known from the type locality. Remarks.— Determining the closest rela- tives is difficult for this species. Taking into account the shape of the uncus, this species can be associated to the Teeerifeesis group (see above). On the other hand, though, the epigynum of the female is very different from those of the Tenerifensis group, and it looks more similar to that of P. baldiosensis Wun- derlich 1992 (the other troglomorphic spe- cies). Unfortunately, we cannot determine whether the male of P. baldiosensis is in fact similar to the Tenerifensis group since it still remains unknown. Taking in account the characteristic features of the epigynum and the procursus, which are remarkably different from those of the other Canarian Pholcus species, P. corniger could possibly be a member of a different group, or it could form a subgroup (with P. baldiosen- sis) of the Tenerifensis group. In order to yield an answer to this question, a detailed phylo- genetic study must be performed. ACKNOWLEDGMENTS We would like to thank to Dr. Pedro Oromi and the other members of the Department of Biology at the Universidad de La Laguna for their inestimable help and collaboration. We thank also to Salvador Carranza for critically reading manuscript. This research was sup- ported by BOS2002-00629 project from the Ministerio de Ciencia y Tecnologia of the Spanish Government. LITERATURE CITED Ancochea, E., J.M. Fuster, E. Ibarrola, A. Cender- ero, J. Coello, F. Heman, J.M. Cantagrel & C. Jamond. 1990. Volcanic evolution of the island of Tenerife (Canary Islands) in the light of the new K-Ar data. Journal of Volcanology and Geo- thermal Research 44:231-249. Anguita, F. & F. Heman. 1975. A propagating frac- ture model versus a hot spot origin for the Ca- 134 THE JOURNAL OF ARACHNOLOGY nary Islands. Earth and Planetary Science Letters 27:11-19. Bosenberg, W. 1895. Beitrag zur Kenntnis der Ar- achniden-Fauna von Madeira und den Canarisch= en Inseln. Abhandiungen und Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 13: 1-13. Campos, C.G. & J. Wunderlich. 1995. The distri- bution of the species of the genus Pholcus Wal- ckenaer on Gran Canaria — a first note, with the description of a new species. Beitrage zur Ara- neologie 4:293-299. Dimitrov, D. & C. Ribera. 2003. Pholcus intricatus (Araneae, Pholcidae) una nueva especie endem- ica de la isla de Tenerife (Islas Canarias). Revista Iberica de Aracnologia 8:7-11. Medina, A.L. 1991. El medio subterraneo superfi- cial en las Islas Canarias: Caracterizacion y con- sideraciones sobre su fauna. Tesis doctoral, Universidad de La Laguna, Tenerife. Oromi, P., A.L. Medina & M.L. Tejedor. 1986. On the existence of a superficial underground com- partment in the Canary Islands. Acta de IX Con- greso Internacional de Espeleologia Barcelona 2: 147-151. Wunderlich, J. 1980. Zur Kenntnis der Gattueg Pholcus Walckenaer, 1805 (Arachnida: Araneae: Pholcidae). Senckenbergiana Biologica 60:219- 227. Wunderlich, J. 1987. Die Spirinen der Kanarischen Inseln und Madeiras. Taxonomy and Ecology 1: 1-435. Wunderlich, J. 1991. Die Spinnen-fauna der Mak- aronesischen Inseln. Beitrage zur Araneologie 1 : 1-619. Manuscript received 17 February 2004, revised 10 September 2004. 2006. The Journal of Arachnology 34:135-141 A NEW SPECIES OF CUPIENNIUS (ARANEAE, CTENIDAE) COEXISTING WITH CUPIENNIUS SALEI IN A MEXICAN MANGROVE FOREST Francisco J. Medina Soriano: Laboratorio de Acarologia “Anita Hoffmann”, Facultad de Ciencias, UNAM, Av. Universidad # 3000, Coyoacan, Mexico, D.R 04510, Mexico ABSTRACT, The new species Cupiennius chapanensis is described from a mangrove forest in the coastal regions of Chiapas, Mexico. The most noticeable characteristic of the species is the bright red coloration of the chelicerae, given by a covering of long setae on the anterior surface; because of this coloration, it has been previously confused with Phoneutria fera Perty 1833. It is generally similar to Cupiennius getazi Simon 1891, but lacks the spotted pattern on the ventral surface of the femora, together with other differences in genitalic morphology. Cupiennius salei Keyserling 1877 was also found on the same forest during the wet season, while C. chiapanensis appeared in the dry season. Adults of both species were never collected at the same time. This is also the first record of C. salei at the sea level, being previously considered a highland species. RESUMEN. Se describe Cupiennius chiapanensis nueva especie, la decima del genero. Fue recolectada del manglar de la costa de Chiapas, Mexico. La caracterfstica mas notoria es la coloracion roja brillante de sus queliceros, dada por una cubierta de sedas largas en el frente del quelfcero. Esta coloracion ha provocado que se le confunda con Phoneutria fera Perty 1833 en tres referencias que se documentan. En general, es similar a Cupiennius getazi Simon 1891, pero no exhibe el patron moteado en la superficie ventral de los femures, ademas de otras diferencias en su estructura genital que se discuten. Se encontro tambien en el mismo bosque a Cupiennius salei Keyserling 1877 durante la epoca de Iluvias, mientras que C. chiapanensis aparecio durante la epoca de secas. Los adultos de ambas especies nunca fueron recolectados en la misma epoca. Este tambien es el primer registro de C. salei a nivel del mar, pues era considerada una especie de tierras altas. Keywords: Mangrove forest, Cupiennius, wandering spiders, taxonomy, new species The genus Cupiennius Simon 1891 is com- prised of nine species distributed throughout Mexico, Central America and Cuba, while Cupiennius celerrimus Simon 1891 is found from Venezuela to Brazil. The latest revision of the genus (Lachmuth et al. 1985) regards the terminal apophysis of the male bulb, and the internal epigynal structure as diagnostic characteristics among the species; the external coloration is often taken into account for some species. In a later work, C remedius Barth & Cordes 1998 was reported as the only species known to share habitat with C. salei Keyser- ling 1877 (Barth & Cordes 1998). During a study on the spider fauna of a mangrove forest in the reserve “La Encruci- jada”, Chiapas, Mexico, specimens of C salei and a different species of the same genus were collected in exactly the same localities, but in different seasons. This other species displays sufficient differences to be considered as a new species. Several references to this same spider have been found in several publications and web pages (Browning 1989; Alvarez del Toro 1992; Spider Homepage 2000) where it has been incorrectly referred to the genus Phoneutria Perty 1833. In the present paper, this new species is de- scribed, the tenth of the genus and the second besides C. salei to be reported from Mexico. Some aspects concerning the occurrence of both species in a mangrove forest are dis- cussed. METHODS The specimens were collected during three expeditions to the reserve “La Encrucijada”, in Chiapas, Mexico, in April 2002, September 2002 and April 2003, on the island “Solo Tu” (15°04'28"N, 92°45'49"W) and in the path 135 136 THE JOURNAL OF ARACHNOLOGY called “La Vida Sigue”, next to the monitor- ing station (15°04'06"N, 92°45'20"W). The plant coverage at the sites corresponds to a mangrove forest, where Rhizophora mangle (red mangrove) is the dominant species, as- sociated with Laguncularia racemosa (white mangrove). The uederstory is mainly com- posed of ferns (Acrosticum aureum) and “pi- nuela” Bromelia plumieri (Rico-Gray 1990). A total of 21 specimens were examined, 13 females and eight males. All measurements are given in millimeters, maximum and min- imum; the numbers within parentheses corre- spond to the mean. Body length was consid- ered to be the distance from the anterior edge of the carapace to the posterior edge of the opisthosoma, carapace length from its anterior to its posterior edge, as well as the opistho- soma length, including spinnerets. Carapace width was taken at the third leg pair; opist- hosoma width was taken at the middle. The general description was based upon dried specimens, since those wet in alcohol appear darker and the setae covering carapace and legs are not clearly visible; in over- dried specimens the chelicerae color may look paler. The holotype and paratype were those com- plete specimens closer to the mean. All type specimens are deposited in the National Arachnid Collection (CNAN), Laboratorio de Acarologia, Instituto de Biologfa, UNAM, Mexico. Only the most common abbreviations were used in the description: AME = anterior me- diae eyes, PME = posterior median eyes, PLE — posterior lateral eyes, RTA = retrolateral tibial apophysis. For male and female genita- lia the names used in Lachmuth et al. (1985) were kept as close as possible. For the male bulb: embolus = Em; terminal apophysis = TAp; embolar apophysis == EAp; conductor = Co; embolar base — StE, For the female epigynum: median plate (septum) = MP; lat- eral plate = LP; seminal receptacle = R; sem- inal duct = SD. TAXONOMY Family Ctenidae Keyserling 1877 Genus Cupiennius Simon 1891 Cupiennius chiapanensis new species Type material* — Mexico: Chiapas: Holo- type female, from the island “Solo Tu”, 15°04'28"N, 92°45'49"W, 17 April 2003, Y. Garcia Martinez (CNAN). Allotype male, I, from the path “La Vida Sigue”, next to the S monitoring station of the reserve “La Eecru- !' cijada”, 15°04'06"N, 92°45'20"W, 10 April ! 2003, E Medina (CNAN). Paratypes: 9 fe- j males from the path “La Vida Sigue” and 3 [ females from the island “Solo Tu”; 6 males collected from “La Vida Sigue” and 1 male [ from “Solo Tu”. ji Etymology, — -The specific epithet refers to ; the state where the species was found. | Diagnosis,^— chiapanensis is j distinguished from other species of the genus i by the characteristic color of the chelicerae j which are mostly covered by bright red setae I in females and pale red setae in males. The female has the epigyeal lateral plates curved on the inner edge; the septum is thinner in its i upper part and wider below, rounded in the middle and square at the end (Fig. 2); the sem- | ieal receptacles are spherical with a small dis- i tal hump and bits of cuticle adhered to the | surface (Fig. 3). The male pedipalp has the j RTA triangular viewed from the front, and square viewed from the side (Fig. 6); the bulb | is quite similar to that of Cupiennius getazi ' (Lachmuth et al. 1985, fig. 3), but the terminal | apophysis has the upper edge gently sloping rather than almost square; the embolar apoph- : ysis is longer and thinner; the conductor is larger, overlapping with the terminal apophy- sis and the embolar base has clearly visible keels through the upper and lower edges of its extension (Fig. 7). Description.- — -Female: Total length 21.9“ 27.0 mm (25.3); carapace length 9.4-13.1 mm (10.8), width 9.0-10.5 mm (9.7); opisthosoma length 11.0-15.0 mm (13.4), width 6.4-8.9 mm (8.2). Leg I: femur 10.9-16.8 (12.2), pa- tella 4.4-5.7 (5.3), tibia 10.2-12.7 (1L3), metatarsus 9.1-13,2 (11.2), tarsus 3. 0-5.0 (4.1). Leg II: femur 10.7-13.5 (1L9), patella 4. 3- 5.9 (5.2), tibia 9.9-13.1 (10.8), metatarsus 9.3- 13.8 (11.2), tarsus 3.2-4.3 (3.9). Leg III: femur 8.1-12.9 (9.9), patella 43-5.9 (5.2), tibia 6.0-10.3 (8.1), metatarsus 6.7-12.2 (8,8), tarsus 2. 5-4. 2 (3.4). Leg IV: femur 10.5-13.7 (11.6) , patella 3.7-5.5 (4.6), tibia 8.3-11.6 (9.6) , metatarsus 10.9-15.1 (12.5), tarsus 3.2- 4.8 (3,9). Pedipalp: femur 4. 1-5.7 (4.7), pa- tella L8-2.5 (2.1), tibia 3.3-4.0 (3.5), tarsus 3. 6-4. 6 (3.9). Prosoma: Carapace orange- brown with a darker brown median band that begins behind PME, reaching posterior edge; Figures 1-3. — Cupiennius chiapanensis new species, female. 1. Dorsal aspect of body, scale =1.0 cm; 2. Ventral view of epigynum, scale =1.0 mm; 3. Dorsal view of epigynum, scale = 1.0 mm. MP = median plate, LP = lateral plate, R = seminal receptacle, SD = seminal duct. 138 THE JOURNAL OF ARACHNOLOGY Figures 4-7. — Cupiennius chiapanensis new species, male. 4. Dorsal aspect of body, scale = 1.0 cm; 5. Ventral view of left pedipalp, scale = 1.0 mm; 6. Retrolateral view of left pedipalp tibia; 7. Embolar area, scale = 0.5 mm. Em = embolus, Co == conductor, MAp = median apophysis, RTA = retrolateral tibial apophysis, TAp = terminal apophysis, EAp = embolar apophysis, ESt = embolar base. MEDINA— Ct/F/£iViV/£/5 FROM MANGROVE FOREST 139 surface covered by short, white setae, except for small spots over ocular area and behind the PLE, where short black setae cover the spot. Long, white setae along the entire cara^ pace edge, those on the anterior lateral edge red colored. Ocular area slightly elevated, long black setae below AME and in between them v/ith long white setae between the PME and PLE and between each other. Fovea Ion- gitudinal with two long, erected setae on an- terior edge and divergent black lines of short setae leading up to the edge of the carapace. Endites black with long white setae along an- terior edge, with white border; long black se- tae curved downwards coming from the lateral endite edge. Labium trapezoidal, black col- ored with anterior border white and covered with long black setae. Sternum light brown with black border, covered with long black se- tae mixed with short white setae. Coxae the same color as sternum. Opisthosoma: General color brown-grayish, covered with long, thick white setae. Mediae black band with winding edge on dorsum in which there is a shorter white band that reaches up to less than half its length, dorsal pattern as illustrated (Fig. 1). Sides of opisthosoma dark and covered with long brown setae. Ventral surface light brown, with a median longitudinal black band, begin- ning in the epigastric furrow and ending at the spinneret circled area. Anterior spinnerets light brown with white apical band, posterior spinnerets darker and slightly longer and thin- ner. Epigyeum with the common structure in the genus, two lateral plates and a median plate or septum. Septum thin anteriorly and wider posteriorly, rounded in the middle, then constrained and ending in a square shape. Lat- eral plates curved on their inner side, linked to the septum by their upper lateral edge, on the dorsal side (Fig. 2). Seminal receptacles spherical, with a small anterior mound and dark bits of cuticle adhered to their surface, similar to the “porose area” described for oth- er lycosoid families by Griswold (1993), the diameter less than half of duct length. Ducts straight, elbowed on the last two thirds (Fig. 3). Chelicerae: General color black, covered with long bright setae on their front surface, up to three quarters of their length, with a few scattered black long setae. Interior border of chelicerae with long white setae, fangs black. Pedipalps: General color dark brown, covered with white setae. Legs: General color dark brown, covered with short white setae, in liv- ing spiders the legs look much darker than the rest of the body. Coxae with long black setae on ventral surface, all trochanters notched with a mediae transverse band of long black setae. Femora densely covered with long black setae with white tip, mixed with short white setae. Femur I with three dorsal spines, 3 prolateral and 3 retrolateral, and dorsum covered with short white setae concentrated in two bands on anterior half. Femur II with two bands of white setae. Femur III without com- plete bands, only spots visible on dorsum. Fe- mur IV without bands. Patellae dark brown covered with white short setae, mixed with some long black ones, without spines. Tibiae dorsally darker and devoid of setae, with one prolateral spine, one retrolateral and three pairs of ventral spines; ventral surface covered with short white setae. Metatarsi without setae on dorsum, with an irregular row of tricho- bothria, two pairs of dorsal spines, 3 prolate- ral, 3 retrolateral and two pairs of ventral spines. Scopula covering the whole ventral surface of metatarsi, except the posterior edge. Tarsi dorsally covered with dense scopula, mixed with tarsal tufts, three tarsal claws, dor- sum covered with short white setae and one or two trichobothria. Male: Total length 7.6™23.6 mm (20.4); carapace length 8,3-“ 10. 8 mm (9.5), width 7.8- 18.9 mm (9.8), opisthosoma length 8.6- 11.9 mm (10.3), width 4.5-16.0 mm (7.2). Leg I: femur 10.9-12.3 (1L7), patella 4.2-5.5 (4.8), tibia 10.6-13.1 (11.8), metatarsus 11.3- 14.2 (12.8), tarsus 4. 1-5.0 (4.6). Leg II: femur 11.3-12.7 (12.1), patella 4.5-5.9 (5.1), tibia 10.8- 12.5 (11.4), metatarso 10.8-13.5 (12.6), tarsus 3.8-4.6 (4.3). Leg III: femur 9.7-11.5 (10.4), patella 3.4-4.5 (3.9), tibia 7.0-8.7 (8.0) , metatarsus 8.8-10.2 (9.5), tarsus 3.1- 4.2 (3.6). Leg IV: femur 10.8-1 L5 (11.3), pa- tella 3.8-5.2 (4.2), tibia 9.3-10.8 (10.0), meta- tarsus 11.4-13.4 (12.4), tarsus 3. 5-4. 5 (4.2). Pedipalp: femur 3. 1-5.0 (4.4), patella 1.7-2. 6 (2.0) , tibia 2.8-4.0 (3.5), tarsus 2.9-3.8 (3.2). Prosoma: Carapace color orange-brown, with a median darker band, beginning behind the PME and reaching the posterior edge. Sur- face entirely covered with short grayish setae that leave only two parallel black lines delin- eating the mediae band. Long white setae along the margin, those on the anterior lateral border orange colored. Ocular area slightly el- 140 THE JOURNAL OF ARACHNOLOGY evated, with white setae above the posterior eyes and between the median eyes. Fovea Ion™ gitudinal. Opisthosoma: General color dark gray, covered with light brown setae, except some small spots where the setae have come off, without any pattern but a pair of parallel black lines that follow from the carapace (Fig. 4). Ventral surface light gray with a median black band originating at the epigastric furrow and ending at the circled spinneret area. Che- licerae: General color black, covered with pale red setae on most of their surface, up to two thirds their length, with some scattered long black setae, interior border with long white setae, fangs black. Eedites and labium dark brown with long black setae. Sternum and coxae light brown, covered with grayish short setae. Pedipalps: General color same as the carapace, dorsally covered with short grayish setae, femur with one dorsal and three apical spines, patella with one apical prolate^ ral spine, tibia with two dorsal spines and long white setae on its lateral borders. RTA short with square shape in lateral view and triae™ gular in ventral view (Fig. 5). Cymbium dark brown, covered with short black and white se™ tae, bulb in agreement to the general structure of the genus and very similar to that of C getazi (Fig.5). Median apophysis with the up- per side rounded and the lateral process sharp and curved downwards. Embolus ending close to the conductor; embolar apophysis slender, hook shaped, and the tip covered with the ter- minal apophysis, which is slightly ovoid and the upper edge slopes down to the embolar base; conductor rounded and directed out- wards, bent downwards on its distal portion, covering part of the embolus (Fig. 7). Legs: Same color as the carapace covered with light grayish setae, with no visible bands, only dark spots on dorsum. Femur with spines similar to those of the female. All legs covered with light setae, except dorsal surfaces of tibiae and metatarsi. Dorsal irregular row of trichoboth- ria on metatarsi; tarsi with no visible tricho- bothria. Ventral surface of metatarsi covered with scopula, not as dense as in the female, and beginning at the second third of its length. Tarsi completely covered with scopula and claw tufts, three tarsal claws. Other structures as in the female. Distribution* — This species has only been collected from the mangrove forest of ‘'La Eecrucijada”, municipality of Acapetahua, Chiapas state, Mexico. DISCUSSION The first external characteristic indicating that Cupiennius chiapanensis could represent a distinct species was the striking red color of the chelicerae, which couldn’t have been over- looked in any other description and is an in- variable feature of every adult which is not lost when kept in alcohol at least after one year; therefore, it is herein regarded as a di- agnostic character. This red color probably led to the confusion of this spider with Phoneutria fera, in two photographic references. The first one is from a web site from Australia (Spider Homepage 2000), where some pictures were sent by a photographer and incorrectly idee- tified; the other is from a book on tarantulas where it is regarded as “the Brazilian hunts- man spider” (Browning 1989:67). A third ref- erence was found in the book “Aranas de Chiapas”, where it is stated that the spider probably belongs in Phoneutria due to the color of the chelicerae (Alvarez del Toro 1992, plate 63). Cupiennius salei sometimes shows red setae on the chelicerae too, as can be observed in color drawings (Cambridge 1900) and even in living spiders, however, they don’t fully cover the front of the chelic- erae, so the characteristic dark longitudinal bands of this species prevail. Barth & Cordes (1998) consider the color- ation of the sternum and coxae as a character to separate some of the species of the genus, pointing out that external body pattern could allow the identification of even subadult spec- imens. The ventral pattern of C. chiapanensis is similar to that of C valentinei Petrunke- vitch 1925, which is the only other species with a ventral black median band. The rest of the coloration, including the dorsal opistho- somal pattern, is very similar to C getazi, which however shows a distinctive spotted ventral surface of the femora, that is not pre- sent in this new species. Cupiennius chiapa- nensis is also similar to C. getazi in the shape of the seminal receptacles and ducts, but the small distal mound and the granules all over the surface of the receptacles, together with the external shape of the epigynum, set them apart. Before this study, only Cupiennius rerne- dius was known to share habitat with C salei. MEDINA— CUPIENNIUS FROM MANGROVE FOREST 141 in the highlands of Guatemala (Barth & Cor- des 1998). Here, C. chiapanensis is the second species to exist in the same place as C salei, but at different times. During the wet season, between September and October 2002, 35 adults of C. salei were collected at exactly the same sites where 18 of C. chiapanensis had been obtained in the dry season, including fe- males carrying egg sacs attached to their spin- nerets. Adults of both species at the same time were never found. Juveniles, on the other hand, are morphologically very similar, so their identification was not possible. In this study, the first report of C. salei at a lower altitude than 800 msnm is given, for it was formerly considered to be a highland species (Barth & Seyfarth 1979). ACKNOWLEDGMENTS This work was financed by the National University of Mexico (UNAM) PAPIIT IN2 15701. I want to thank Dr. Anita Hoff- mann for advice and revision of the manu- script. Thanks to Donaji Cid, who collaborat- ed with illustrations of the spider’s body. Also thanks to the personnel from reserve “La En- crucijada” and those involved in the field work. LITERATURE CITED Alvarez del Toro, M. 1992. Las aranas de Chiapas. Ed. Universidad Autonoma de Chiapas. Mexico. Barth, EG. & D. Cordes. 1998. Cupiennius reme- dius new species (Araneae, Ctenidae) and a key to the genus. Journal of Arachnology 26: ISS- UE Barth. EG. & E.A. Seyfarth. 1979. Cupiennius salei Keys (Araneae) in the highlands of central Gua- temala. Journal of Arachnology 7:255-263. Browning, J.G. 1989. Tarantulas. TEH. Publica- tions, Inc. Cambridge, EO.-R 1900. Arachnida — Araneida and Opiliones. In Biologia Centrali-Americana, Zo- ology. Taylor and Francis, London, vol. 2: SO- US. Lachmuth, U., M. Grasshoff & EG. Barth. 1985. Taxonomische revision der Gattung Cupiennius Simon 1891 (Arachnida: Araneae: Ctenidae). Senckenbergiana Biologica 65:329-372. Rico- Gray, V. 1990. Observaciones y comentarios preliminares al estado actual de la flora y vege- tacion de La Encrucijada municipio de Acape- tahua, Chiapas, Mexico. Informe del Programa Flora de Mexico. Proyecto “Flora Yucatanen- sis”. Simon, E. 1891. Descriptions de quelques arachni- des du Costa Rica communiques pa M. A. Getaz (de Geneve). Bulletin de la Societe Zoologique de France 16:109-112. Spider Homepage (2000). http://www.rochedalss. qld.edu.au/spider/wandering.htm. Manuscript received 9 September 2003, revised 25 October 2004. 2006. The Journal of Arachnology 34:142-158 HAVE YOU SEEN MY MATE? DESCRIPTIONS OF UNKNOWN SEXES OF SOME NORTH AMERICAN SPECIES OF LINYPHHDAE AND THERIDHDAE (ARANEAE) Nadine Dupere: 341 15 eme me, Laval, Quebec, H7N 1L5, Canada. E-mail: dupere.nadine @ videotrm.ca Pierre Paquinb Department of Biology, San Diego State University, San Diego, California, 92812-4614, U.S.A. E-mail: paquinp@mlmk.net Donald J. Buckle: 620 Albert Avenue, Saskatoon, Saskatchewan, S7N 1G7, Canada ABSTRACT. The previously unknown sexes of 13 species of Linyphiidae and Theridiidae are described and illustrated for the first time. These include the following members of the Linyphiidae: Centromerus furcatus (Emerton) female, Cheniseo sphagnicultor Bishop & Crosby female, Coloncus siou Chamberlin male, Dismodicus alticeps Chamberlin & Ivie female, Floricomus praedesignatus (Bishop & Crosby) female, Glyphesis idahoanus (Chamberlin) male, Gnathonaroides pedalis (Emerton) female, Lepthyphan- tes intricatus (Emerton) female, Scyletria inflata Bishop & Crosby female, Sisicus penifusifer Bishop & Crosby female, Walckenaeria clavipalpis Millidge male; and two members of the Theridiidae: Thymoites minnesota Levi female and Robertus crosbyi (Kaston) male. Synonymy, new records, and comments on distribution, habitat and taxonomy are also given. The generic placement of Glyphesis idahoanus (Cham- berlin) and Glyphesis scopulifer (Emerton) is confirmed. Keywords: taxonomy, Canada, U.S.A., undescribed sexes, distribution, museum collection Recently, Paquin & Duperre (2003) pub- lished an identification guide to the known and suspected spiders of Quebec. In the treat- ment of these species, we came across several for which only one sex was known. The other sex of these species is described below. We also give the synonymy, new distribution data and provide relevant taxonomic and ecologi- cal comments. The failure to recognize match- ing sexes may result in situations where a fe- male is known under a certain name and the male under another, therefore leaving the false impression that only one sex is known. Such cases, however possible, are not common and for most species in which only one sex is known, there may be rather simple and direct explanations. Firstly, the species is quite rare and it happens that only one sex was collect- ed. Secondly, differences in either microhab- itat selection or behavior of males and females may result in one sex being overlooked. For instance, approximately one hundred males are known for Maro amplus Dondale & Buck- le 2001, but so far, the female remains un- ' Corresponding author. known (Dondale & Buckle 2001). In other cases, such as Nesticus Thorell 1869 (Nestici- dae), the ratio of specimens collected in the field largely favors females and juveniles while mature males are rarely encountered (M. Hedin pers. comm., pers. obs.). Similar observations were made by Gertsch (1992) for the genus Cicurina Menge 1871 (Dictynidae). Thirdly, in most cases the undescribed sex has been col- lected and properly assigned to a species but awaits formal description. Most species treated in this paper belong to this last category. We have collected both sexes of some spe- cies [Gnathonaroides pedalis (Emerton 1923), Centromerus furcatus (Emerton 1882), Lep- thyphantes intricatus (Emerton 1911), Dis- modicus alticeps Chamberlin & Ivie 1947, Scyletria inflata Bishop & Crosby 1938, Co- loncus siou Chamberlin 1949, Sisicus penifus- cifer Bishop & Crosby 1938, Thymoites min- nesota Levi 1964], in the same pitfall sample, or together in the field, thus allowing the as- sociation. Most other records were sorted to- gether in vials belonging the Canadian Na- tional Collection. These associations were 142 DUPERRE ET AL.-^LINYPHIID AND THERIDIID DESCRIPTIONS 143 made over the years by C.D. Dondale and J.H. Redner from samples in which both sexes were collected together. METHODS Specimens were examined in 70% ethanol under a SMZ~U Nikon dissection microscope. A Nikon Coolpix 950 digital camera attached to the microscope was used to take a photo^ graph of the structure. The digital photo was then used to trace proportions and the illustra- tion was detailed and shaded by referring back to the structure under the microscope. Female genitalia were excised using a sharp entomo- logical needle and transferred to lactic acid to clear non-chitinous tissues. A temporary lactic acid mount was used to examine the genitalia under an Olympus BX40 microscope, and was photographed and illustrated as explained above. All measurements were made using a micrometric ruler fitted on the eyepiece of the microscope. When available, 5 specimens were measured for the description. Calcula- tion for the location of Tml follows Denis (1949). Most of the specimens studied were from the Canadian National Collection of Insects and Arachnids, Ottawa, Canada (CNC). In ad- dition, material from several other collections was examined. The collection is indicated in brackets and unless specified otherwise, the specimens are deposited in the CNC. Abbre- viations used: AG = Collection of Alice Gra- ham; CMB = Collection of C.M. Buddie; CPAD = Collection of Paquin-Duperre; DJB = Collection of D.J. Buckle; DSU ^ Dick- inson State University, North Dakota (cur- rently at Texas A&M International Universi- ty); HAC = Collection of H.A. Carcamo; MCZ = Museum of Comparative Zoology, Harvard University; MLC = Collection of Maxime Larivee; RF ^ Collection of Robert Fimbel; RGH = Collection of R.G. Holmberg; RPC ” Collection of Roger Pickavance; RSM = Royal Saskatchewan Museum; UASM = University of Alberta, Strickland Museum. Latitude and longitude given for each locality should be considered approximate. TAXONOMY Family Linyphiidae Blackwall 1859 Genus Centromerus Dahl 1886 Centromerus fur catus (Emertoe 1882) Figs. 1-3 Microneta furcata Emerton 1882:76, pi. 24 fig. 5. Centromerus furcatus (Emerton): van Helsdingen 1973:27, figs. 22-24; Jennings et al. 1988:61; Be- langer & Hutchinson 1992:50; Buckle et al. 2001: 105; Paquin et al. 2001:16; Paquin & LeSage 2001:96; Paquin & Duperre 2003:137, figs. 1503-1506. Material examined.— U.S. A.: Maine: Pis- cataquis County Soubunge Mountain [45°58'N, 69°12'W], Id, 1 9 (CNC); CAN- ADA: Newfoundland: Eastern Blue pond [50°27'N, 57°07'W], 16,19 (CNC); Crab- bes River [48°13'N, 58°52'W], 2 9 (CNC); Barachois Brook [48°27^N, 58°26'W], 1 9 (CNC); Lloyd’s Lake [48°23'N, 57°3rW], 2 9 (CNC); Highlands River [48°1UN, 58°53'W], 1 9 (CNC); Big Falls [47°05'N, 54°03'W], 1 9 (CNC); Pasadena [49°0rN, 57°36'W], 1 d, 6 9 (CNC); New Brunswick: Green River 30 mi N Edmunston [47°19'N, 65°27'W], 2 d (CNC); Quebec: Parc de la Gaspesie Mont Albert [48°56'N, 66°10'W], 1 9 (CNC); 24 mi S of Ste-Aene-des-Moets [48°52'N, 65°58'W], 3 d (CNC); Abitibi Lac Duparquet [48°30'N, 79°13'W], 1 d, 1 9 (CPAD). Description. — Female (n =5): Total length: 1.35 ± 0.08 mm; carapace length: 0.63 ± 0.03 mm; carapace width: 0.47 ± 0,02 mm; carapace smooth, shiny, light yellow to yellow with a tinge of orange, lightly shaded with gray along radiating lines; carapace margin more strongly shaded, 3-4 erect setae along midliee; sternum yellow, strongly shaded with gray, margin darker. Chelicerae yellow with a tinge of orange, promargie with 3 large teeth, retromargie with 5 denticles. Cheliceral strid- ulatory organ not visible with stereomicro- scope. Abdomen unicolor, off-white, lightly suffused with gray, densely covered with long semi-erect setae. Legs light yellow to yellow with a tinge of orange, tibia LIV with two dorsal macrosetae; metatarsus I with dorsal trichobothrium, Tml 0.28-0.33, TmlV absent. Epigynal plate flat, protruding, wider than long; scape short, broad, straight or widening slightly toward the tip, cochlear present (Figs. 1, 2); spermathecae bean- shaped (Fig. 3). Distribution.' — Eastern species, southern- most record from New Hampshire (Buckle et al. 2001). Habitat. — Collected in coniferous habitat, in moss and forest litter. 144 THE JOURNAL OF ARACHNOLOGY Genus Cheniseo Bishop & Crosby 1935 Cheniseo sphagnicultor Bishop & Crosby 1935 Figs. 4, 5 Cheniseo sphagnicultor Bishop & Crosby 1935a: 263, pL 21 figs. 64-69; Buckle et aL 2001:110; Paquin et al. 2001:16; Paquin & Duperre 2003: 96, figs. 883-886. Acartauchenius sphagnicultor (Bishop & Crosby): Aitchison-Benell & Dondale 1992:221; Dondale & Redner 1994:36; Belanger & Hutchinson 1992: 22. Material examined.— CANADA: Nova Scotia: Cape Breton National Park French Lake [46°44'N, 60°52^W], 1 $ (CNC); Cape Breton National Park North Mount [46°53'N, 60°35'W], 2 (5 (CNC); Quebec: Gatineau Park Hopkin’s Hole [45°34'N, 75°57'W], 1 (7 (CNC); Ontario: Alfred [45°33'N, 76°52'W], 1 (7, 1 ? (CNC); Mer Bleu 8 miles E. of Ottawa [45°24'N, 75°30'W], 8 (7, 3 $ (CNC); Upper Rock Lake 30 km N Kingston [44°30'N, 76°24'W], 1 <7 (CNC); Crieff Bog 3 km W Puslinch [43°26'N, 80°05'W], 1 c7 (CNC); Wylde Lake Bog 8 km E Arthur [43°50'N, 80°22'W], 7 (7, 1 9 (CNC); Brm cedale conservation area nr Port Elgin [44°26'N, 8r24'W], 1 9 (CNC); Manitoba: Riding Mountain National Park Swanson Spring [50°53'N, lOOMS'W], 6 d, 1 9 (CNC). Description. — -Female (n = 5): Total length: 0.98 ± 0.07 mm; carapace length: 0.43 ± 0.05 mm; carapace width: 0,33 ± 0.04 mm; carapace smooth, shiny, light brown to dark brown, radiating lines and carapace margin with diffuse gray pattern, cephalic region oc- casionally ornamented by a dark gray marking forming a trident (or psi, T"); 3 long erect setae along midline; sternum light brown to dark brown strongly shaded with gray. Chelicerae yellow to light brown, promargin with 1 large tooth and 5 small teeth, retromargin with 4-5 denticles. Cheliceral stridulatory organ not visible with stereomicroscope. Abdomen uni- color, light to dark gray, densely covered with short semi-erect setae. Legs light yellow with ; a tinge of orange, tibia I-IV with one dorsal I macroseta; metatarsus I with dorsal tricho- bothrium, Tml 0.38-0.47, TmlV absent. Epi- gyeum with plate resembling an hexagon; me- dian lobe, broad, pointed, extending in pale area; copuiatory openings small, round, situ- ated at anterior end of the median lobe (Fig. 4); spermathecae round, widely separated, flanking the median lobe (Figs. 4, 5). Distribution.— Species restricted to the eastern portion of North America, W to Man- j itoba. i Habitat. — ^This species has been collected | in coniferous forest litter but seems mainly as- | sociated with sphagnum bogs. Genus Coloncus Chamberlin 1949 Coloncus siou Chamberlin 1949 Figs. 6-8 Coloncus siou Chamberlin 1949:525 figs. 48, 49; Levi & Levi 1955:36; Belanger & Hutchinson 1992:27; Buckle et al. 2003.110; Paquin et al. 2001:16; Paquin & Duperre 2003:97, figs. 891- 893. Material examined, — U.S.A.: Massachu- setts: Barnstable County: Quisset [41°43'N, 70°39^W], 10 (7, 8 9 (CNC); North Dakota: Dunn County: Lake Ilo [47°20'N, 102°39'W], 1 S (DSU); Canada: Quebec: Gatineau Park King Mountain [45°29'N, 75°52'W], 1 c7 (CNC); Saskatchewan: 10 km S Cadillac [49°30'N, 107°50'W], 22 c7, 4 9 (RSM); Grasslands National Park West Block [49°07'N, 107°26'W], 55 c7, 27 9 (DJB); North Battleford [52°47'N, 108°17'W], 5 (7, 2 9 (DJB); Morse [50°30'N, 106°53'W], 15 (7, 2 9 (RSM); 22 km W Hazlet [50°24'N, 108°36'W], 1 d (RSM); 5 mi NE Saskatoon [52°irN, 106°34'W], 18 (7 (DJB); 21 km N Scotsguard [49°43^N, 108°09'W], 1 (7 (RSM); Figures 1-10. — Linyphiid genitalic structures: 1 — 3.Centromerus furcatus: 1. Epigynum, ventral view; 2. Epigynum, lateral view; 3. Spermathecae, dorsal view. 4, 5. Cheniseo sphagnicultor: 4, Epigynum, ventral view; 5. Spermathecae, dorsal view. 6-8. Coloncus siou: 6. Palpal tibia of male, dorsal view; 7. Palpal cymbium, lateral view; 8. Palpus of male, ventral view. 9, 10. Dismodicus alticeps: 9. Epigynum, ventral view; 10. spermathecae, dorsal view. Abbreviations used: CH = Cochlea, CO — Copuiatory Opening, CY = Cymbium, E = Embolus, ML = Median Lobe, MPr = Median Process, PTA = Palpal Tibia Apophysis, SC = Scape, S = Spermatheca. DUPERRE ET AL=— LINYPHIID AND THERIDIID DESCRIPTIONS 145 146 THE JOURNAL OF ARACHNOLOGY 24 km N Shaunavoe [49°53'N, 108°30'W], 7 (?, 1 $ (RSM); ~10 km NE Simmie [49°59'N, 108°00'W], 9 (7, 1 $ (RSM); AN berta: Lethbridge [49°42'N, 112°49'W], 2 c? (DJB); Suffield [50°12'N, IIUIO'W], 25 (3, 6 9 (DJB). Description. — Male (n =5): Total length; 1.58 ±0.11 mm; carapace length: 0.66 ± 0.03 mm; carapace width: 0.54 ± 0.04 mm; cara- pace smooth, shiny, light brown to brown with diffused gray markings along midline and ra- diating line, carapace margin with dark gray markings; 6 short erect setae along midline; sternum brown strongly shaded with gray, margin darker. Chelicerae yellow to light brown, paler basally and apically, promargin with 3 large teeth and 1 small tooth, retro- margin with 2 small teeth. Cheliceral stridu- latory organ easily visible with —20 ridges. Abdomen unicolor, dark gray, sparsely cov- ered with long erect setae. Legs light yellow with a tinge of brown, coxae lightly shaded with gray; tibia LIV with one dorsal macro- seta; metatarsus I with dorsal trichobothrium, Tml 0.40-0.48, TmlV absent. Palpal tibia with two apophyses (Fig. 6); cymbium with large, deep, longitudinal, retrolateral groove (Fig. 7); paracymbium concealed behind pal- pal tibia apophysis; embolus flat, ribbon like, curving twice at almost a right angle (Fig. 8). Distribution. — Apparently a northern spe- cies. It has been found from Alberta to Que- bec to the North and in North Dakota and Massachusetts (Buckle et al. 2001). Habitat, — This species appears to inhabit forest litter and moss in the east of its distri- bution, and prairie in the west. Remarks. — Five species are listed in the genus Coloncus (Buckle et al. 2001). One spe- cies, C. americanus, was described by Cham- berlin & Ivie (1944), the remaining by Cham- berlin (1949). Four of these species were described from females only and appear very similar based on available illustrations. Colon- cus cascadeus Chamberlin 1949 was briefly described from both male and female, but no illustrations of the genitalia were provided. The description and illustration of the male of Coloncus siou given here will hopefully bring attention to the genus and result in a re-ex- amination of the five species, which may prove to be synonyms. On the other hand, C. siou is associated with forest and moss in the East, but specimens collected in Saskatche- wan, Alberta, Montana and North Dakota are found in prairie habitats (Buckle unpub.). It is presently unclear whether this indicates a broad habitat selection for C siou, or that more than one species is present. A revision of the genus is necessary to clarify these ques- tions. The name Coloncus siou has been used for the species found in the East to remain consistent with Buckle et al. (2001). As mentioned in Buckle et al. (2001) and Paquie et al. (2001), the date of Chamberlin’s paper “On some American spiders of the Family Erigonidae” is erroneously cited as 1948. The paper was published in 1949 as stated on page 570 of the volume 41 of the Annals of the Entomological Society of America. Genus Dismodicus Simon 1884 Dismodicus alticeps Chamberlin & Ivie 1947 Figs. 9, 10 Dismodicus alticeps Chamberlin & Ivie 1947:34 figs. 29-31; Hackman 1954:28, figs. 69-71; West et al. 1984:86; Belanger & Hutchinson 1992:28; Aitchison-Benell & Dondale 1992:222; Mamsik et al. 1993:76; Hutchinson 1994:168; Dondale et al. 1997:83; Buckle et al. 2001:112. Paquin et al. 2001:17; Paquin & Duperre 2003:99, figs. 918- 921. Material examined.- — U.S.A.: North Da- kota: Benson County: Wood Lake [47°54'N, 98°53'W], 1 (3, 2 $ (DSU); Bottineau County [County record only], 1 (3, 2 $ (DSU); Ro- lette County [County record only], 1 ? (DSU); Rolette County: Fish Lake [48°06^N, 99°33'W], 1 (3 (DSU); CANADA: Newfound- land: Noel Pauls Brook [48°49'N, 56°18'W], 1 9 (CNC); Nova Scotia: Cape Breton High- lands National Park North of Paquet Lake [46°48'N, 60°4rW], 4 c3 (CNC); Cape Breton Highlands National Park New Ross Lunen- burg County [44°44'N, 64°27'W], 2 c3 (CNC); Cape Breton Highlands National Park Sweet’s Cove [44°44'N, 64°27'W], 1 (3, 12 9 (CNC); Hebbvilie [44°2rN, 64°32^W], 2 9 (CNC); Cape Blomidon [45°13'N, 64°22'W], 1 9 (CNC); Kentville [45°05'N, 64°30'W], 3 9 (CNC); New Brunswick: Fredericton Lincoln [45°54'N, 66°35'W], 2 9 (CNC); Green River 30 mi N Edmunston [47°19'N, 68°09'W], 1 9 (CNC); Kouchibouguac National Park [46°5rN, 64°58'W], 4 9 (CNC); Quebec: Lac Roddick [46°15'N, 75°53'W], ! 9 (CNC); La Riviere-du-Nord, Saint-Hippolyte, Station DUPERRE ET AL.— LINYPHIID AND THERIDIID DESCRIPTIONS 147 biologie Ueiversite de Montreal [45°59'N, 74°00^W], 4 d, 4 $ (CPAD); Ontario: Shir-^ leys Bay 15 km w of Ottawa [45°22'N, 75053 'W], 1 $ (CNC); Algonquin Provincial Park Lake Opeoego [45°42'N, 78°23'W], 3 d, 6 $ (CNC); Algonquin Provincial Park Lake Opeongo Deer Island [45°42'N, 78°23'W], 5 $ (CNC); Petawawa [45°54'N, 77°20'W], 2 d, 3 ? (CNC); Iroquois Falls [48°46'N, 80°4rW], 1 ? (CNC); Manitoba: Riverton [50°59'N, 96°59A¥], 14 <5, 22 ? (CNC); 15 km SW Swan River [SLSS'N lOlAV], 2 $ (DJB); South Indian Lake [56°47'N, 98"56'W], 1 ? (CNC); Seddoe’s Corner [50°03'N, 96"i7'W], 4 9 (CNC); Pine Falls [50°33^N, 96n3^W], 1 d, 5 9 (CNC); Rennie [49°5rN, 95°33'W], 2 d, 2 9 (CNC); Agas=» siz Provincial Park [49°59^N, 96°09'W], 3 9 (CNC); Telford [49'^50'N, 95°23'W], 1 d, 3 9 (CNC); Darwin [49°55'N, 95°49'W], 4 d, 4 9 (CNC); Gleelea [49°38'N, 97‘^08'W], 1 9 (CNC); Eardley Lake [52°3rN, 96WAV], 1 9 (CNC); Spuce Woods Provincial Forest [49°46'N, 99°2rW], 3 9 (CNC); Ninette [49°20'N, 99°33'W], 1 d (CNC); Riding Mountain National Park [50°39'N, 99°58'W], 1 9 (CNC); Saskatchewan: Lady Lake [52°02'N, 102°37'W], 4 d, 3 9 (DJB, RGH); Anglin Lake [53°44'N, 105°56'W], 3 d, 2 9 (DJB); Fort Carlton [52°52'N, 106°32'W], 1 9 (DJB); Besnard Lake [55°25'N, 106°00'W], 1 d, 2 9 (DJB); Alberta: Winfield [52°58'N, 114°26'W], 2 9; Fox Lake Reservation [58°26'N, 114°33'W], 1 d, 1 9 (DJB); Wee^ tzel Lake [59"02'N, 114°28'W1, 2 d, 1 9 (UASM); Steele Lake [54°40'N, 113°38'N], 2 d (DJB); Athabasca [54°43'N, 113°17'W], 1 9 (DJB); Baptiste Lake [54°45^N, 113°35'W], 1 d, 2 9 (DJB); Marguerite Crag and Tail Provincial Park [57°43'N, 110°20'W], 1 9 (UASM); 90 km NW Peace River [56°42^N, 118°29'W], 2 9 (DSU); British Columbia: Little Prairie Lake [54°57'N, 120°irW], 1 9 (CNC); Babirie Lake Johnson Bay [54°45'N, 126°00'W], 9 9 (CNC); Atlie [59°34'N, 133°42'W], 2 9 (CNC); Northwest Territory: Martin River [bUSS^N, 12r34'W], 1 9 (CNC); Maueoir Lake [67°29'N, 124°55'W], 1 9 (CNC); Wrigley [63M6'N, 123°36'W], 2 d, 2 9 (CNC); Yukon Territories: Kathleen Lake Kluane National Park [60°34'N, 137°17'W], 5 9 (CNC); Gravel Lake 58 mi E Dawson [63°48'N, 137°53'W], 2 9 (CNC); 13 mi E Dawson [64°03'N, 139°25'W], 2 9 (CNC); Old Crow [67°35'N, 137°53'W], 1 d, 2 9 (CNC). Description, — Female (n = 5): Total length: 2.14 ± 0.46 mm; carapace length: 0.86 ± 0.09 mm; carapace width: 0.67 ± 0.05 mm; carapace smooth, shiny, yellow to light or- ange, radiating lines light brown, cephalic re= gioe of the carapace occasionally ornamented by a gray marking forming a trident (or psi, 'T'); 5 short erect setae along midliee; sternum yellow to light orange with dusky gray mar- gins. Chelicerae yellow to light orange, pro- margin with 4 large teeth, retromargin with 3“- 4 large teeth; cheliceral stridulatory organ not visible with stereomicroscope. Abdomen uni- color, gray to dark gray, densely covered with semi-erect setae. Coxae, femora and patella yellow to light orange, tibia, metatarsi and tar- si light orange to dark brown, tibia LIV with one dorsal macroseta; metatarsus I with dorsal trichobothrium, Tml 0.68-0.82, TmlV pre- sent. Epigynum with plate distinctly wider than long, posterior end of plate rising and recurving into a median process, tapered to- ward midline (Fig. 9); spermathecae c-shaped, beanlike, widely separate, situated near the anterior end of the epigyeal plate (Figs. 9, 10). Distribution.' — Widespread species, appar- ently boreal. Habitat.' — This species has been recorded from several habitats, but mainly on conifer- ous vegetation and in forest litter. Genus Floricomus Bishop & Crosby 1925 Floricomus praedesignatus Bishop & Crosby 1935 Figs. 11, 12 Floricomus praedesignatus Bishop & Crosby 1935b:38, pL 6 figs. 22-24; Hormiga 1994:32 figs. 19e, f; Belanger & Hutchinson 1992:31; Buckle et al. 2001:119; Paquie et al. 2001:17; Paquin & Duperre 2003:105, figs. 1042-1045. Material examined. — U.S.A.: North Car- olina: Jackson County: Blue Ridge Park [36°53'N, 80°95'W], 1 d (CNC); CANADA: Quebec: Pink Lake Gatineau Park [45°28'N, 7"5°48'W], 1 9 (CNC). Description, — Female (n = 1): Total length: 1.39 mm; carapace length: 0.58 mm; carapace width: 0.50 mm; carapace smooth, shiny, cephalic region and cervical groove dark brown shaded with gray; thoracic region light brown with radiating line slightly shaded with gray; carapace margin strongly shaded; 148 THE JOURNAL OF ARACHNOLOGY Figures 11-19. — Linyphiid structures: 11, 12. Floricomus praedesignatus: 11. Epigynum, ventral view; 12. Spermathecae, dorsal view. 13, 14. Glyphesis idahoanus: 13. Palpal tibia of male, dorsal view; 14. palpus of male, ventral view. 15, 16. Gnathonaroides pedalis: 15. Epigynum, ventral view; 16. Sperma- thecae, dorsal view. 17-19. Lepthyphantes intricatus: 17. Abdomen, lateral view; 18. Epigynum, ventral view; 19. Spermathecae, dorsal view. Abbreviations used: CH = Cochlea, CDu = Copulatory Ducts, E = Embolus, EM = Embolic Membrane, L = Lobe, PTA = Palpal Tibia Apophysis, SC = Scape, S = Spermatheca, TP = Tail Piece. DUPERRE ET AL,— LINYPHIID AND THERIDIID DESCRIPTIONS 149 sternum dark brown suffused with gray, mar- gin darker. Chelicerae light brown, promargin with 1 large tooth and 5 small teeth, retro- margin with 5 denticles. Cheliceral stridula- tory organ not visible with stereomicroscope. Abdomen unicolor, dark gray, densely covered with semi-erect setae. Legs light orange with a tinge of brown, tibia I without dorsal ma- crosetae, tibia II-IV with one such seta; meta- tarsus I with dorsal trichobothrium, Tml 0.45; TmlV absent. Epigynum with plate inconspicuous, slightly convex, bearing two longitudinal fissures (Fig. 11); spermathecae rounded, and separated by less than half of their width (Figs. 11, 12). Distribution. — Based on the few records known, this is an eastern species. Habitat. — Floricomus praedesignatus has been collected in forest litter. Genus Glyphesis Simon 1926 Glyphesis idahoanus (Chamberlin 1949) Figs. 13, 14 Tapinocyba idahoana Chamberlin 1949:551 figs. 129, 130; West et al. 1988:82. Glyphesis idahoana (Chamberlin): Aitchison-Be- nell & Dondale 1992:222; Belanger & Hutchin- son 1992:31; Dondale & Redner 1994:37. Glyphesis idahoanus (Chamberlin): Buckle et al. 2001:119; Paquin et al. 2001:17; Paquin & Du- perre 2003:106, figs. 1050-1052. Material examined. — CANADA: Quebec: Les Buissons [49°06'N, 68°23'W], 1 $ (CNC); Lac Roddick [46°15'N, 75°53'W], 5 d, 2 $ (CNC); Ontario: Richmond [45°irN, 75°50'W], 1 $ (CNC); Schaffeys Locks [44°35'N, 76°19'W], 1 $ (CNC); ~10 km W Carleton Place [45°08'N, 76°09'W], 1 d, 2 $ (CNC); Alfred [45°33'N, 76°52'W], 1 $ (CNC); Manitoba: Dauphin [51°09'N, 100°03'W], 1 d (CNC); Saskatchewan: Grasslands National Park West Block [49°07'N, 107°26'W], 1 d (DJB); British Co^ lumbia: Oliver [49°1LN, 119°33'W], 2 $ (CNC). Description. — Male (n ^5): Total length: 1.32 ± 0.09 mm; carapace length: 0.51 ± 0.03 mm; carapace width: 0.47 ± 0.03 mm; cara- pace smooth, shiny, dark brown with radiating line and margin strongly shaded with gray, ce- phalic region ornamented by a dark gray in- verse pear shape marking, 3 long erect setae along midline; sternum dark brown strongly shaded with gray. Chelicerae yellow to light brown, promargin with 3 large and 2 small teeth, retromargin with 4 denticles. Cheliceral stridulatory organ not visible with stereomi- croscope. Abdomen unicolor, dark gray, densely covered with short semi-erect setae. Legs yellow, coxae lightly shaded with gray, tibia I-IV with one dorsal macroseta; metatar- sus I with dorsal trichobothrium, Tml 0.44- 0.48, TmlV absent. Palpal tibia with one cup- like apophysis, dark brown to black, long, broad with 3 thick serrate setae (Fig. 13); sec- ond apophysis black, short, half hidden behind the cuplike apophysis; tail piece rather small, rounded; embolus thick, stout, hidden behind embolic membrane (Fig. 14). Distribution. — Widespread species in Can- ada: from British to Quebec (Buckle et al. 2001). Habitat. — This species has been recorded from litter near ponds, sphagnum and salt marshes. Remarks. — This species was originally placed in the genus Tapinocyba Simon 1884 by Chamberlin (1949:551) based on the fe- male genitalia. In defining the genus Glyphes- is, Simon (1926:350) gave a diagnostic feature in the male palpal tibia bearing several strong setae (see Simon 1926, fig. 605). The male of T. idahoana has the same character as that illustrated by Simon for the type species G. servulus (Simon 1881). The species was placed in Glyphesis by Aitchison-Benell & Dondale (1992), without indication that this was a new combination, and subsequent au- thors have followed this placement. The ex- amination and illustration of the male palp of the species confirm the generic placement in Glyphesis, along with Glyphesis scopulifer (Emerton 1882), which has the same tibial character (see Paquin & Duperre 2003:106, fig. 1054). Holm (1968) proposed that G. sco- pulifer was a junior synonym of G. servulus (the type species), but this synonymy has been rejected (Buckle et al. 2001). Genus Gnathonaroides Bishop & Crosby 1938 Gnathonaroides pedalis (Emerton 1923) Figs. 15, 16 Araeoncus pedalis Emerton 1923:239, fig. 2. Gnathonaroides pedale (Emerton): Jennings et al. 1988:61, Peck 1988:1202, Belanger & Hutchin- son 1992:32. Gnathonaroides pedalis (Emerton): Bishop & Cros- 150 THE JOURNAL OF ARACHNOLOGY by 1938:84, pL 6 figs. 65, 66; Levi & Field 1954: 447; Buckle et al. 2001:120; Paquin et al. 2001: 17; Paquin & Duperre 2003:106, figs. 1056” 1058. Material examined. — U.S.A.: New Hamp- shire: Somerswort [43°15'N, 70°5rW], 1 (3 (CNC); Vermont: Mounts Mansfield [44°32'N, 72°48^W], 2 d (CNC); Maine: Pis^ cataquis County: Soubunge Mountain [45°58'N, 69°12'W], 1 <7 (CNC); CANADA: Nova Scotia: Cape Breton Highland National Park Lone Shieling [46°48'N, 60°57'W], 1 (CNC); Bridgewater [49°17'N, 122°54'W], 10 S (CNC); New Brunswick: Acadia forest 10 mi E of Fredericton [45°56^N, 66°40'W], 5 d (CNC); Kouchibouguac National Park [46°51'N, 64°58'W], 1 9 (CNC); Quebec: Lac Roddick [46°15'N, 75°53'W], 1 d (CNC); MonUAlbert, La Haute-Gaspesie, Parc de la Gaspesie, Ruisseau Cap Seize [48°59'N, 66°2rW], 1 d (CNC); MasMeonge, Sainte^ Angele^de^-Premoet [46°2rN, 73°03'W], 1 <3 (CNC); Drummondville [45°53'N, 72°29'W], 1 9 (CNC); Lac Duparquet Abitibi [48°30'N, 79°13'W], 3 d, 3 9 (CPAD); Ontario: 1 km W Carleton Place [45°08'N, 76°09'W], 4 d (CNC); Eastman Farm Chatterton [44°15'N, 77°29'W], 4 d (CNC); El Dorado Gold Mine [44°45'N, 78°06'W], 1 d (CNC); Guelph [43°33'N, 80°15'W], 1 d (CNC); Ancaster [43°13'N, 79°59'W], 2 d, 1 9 (CNC); Rait [48°50'N, 89°56'W], 1 d (CNC); Manitoba: Onanole [50°37'N, 99°58'W], 3 d, 1 9 (CNC); Riding Mountain National Park Clear Lake [50°40'N, 100°00'W], 1 d (CNC); Sas- katchewan: Lady Lake [52°02'N, 102°37'W], 1 d (DJB); Alberta: Edmonton [53°33'N, 113°20'W], 3 d (DJB); George Lake 16 km W Busby [53°57'N, 114°06'W], 1 d (CMB). Description, — Female (n = 3): Total length: 1.19 ± 0.13 mm; carapace length: 0.56 ± 0.05 mm; carapace width; 0.40 ± 0.05 mm; carapace smooth, shiny, light yellow, cephalic region light yellow with a tinge of orange, ra- diating lines and midline with diffused gray patterns; 5 long erect setae along midline; sternum yellow shaded lightly with gray, mar- gin darker. Chelicerae yellow to light brown, promargin with 4 large teeth and 1 small tooth, retromargin with 5 denticles. Cheliceral stridulatory organ not visible with stereomi- croscope. Abdomen unicolor, off-white, densely covered with long semi-erect setae. Legs yellow with a tinge of orange, tibia I-III with two dorsal macrosetae and tibia IV with one dorsal seta; metatarsus I with dorsal tri- chobothrium, Tml 035-0.38, TmlV absent. Epigyeal plate conspicuous, somewhat pen- tagonal, with two longitudinal dark bands con- verging below the middle of the plate, trans- versal band present near the posterior margin; posterior margin darker, more sclerotized (Fig. 15) ; spermathecae rounded, widely separated, situated at edge of lateral margin (Figs, 15, 16) .^ Distribution.-— pedalis oc- curs in northern North America east of the Rockies. Buckle et al. (2001) report the spe- cies from New York and Maine. Habitat. — -This species has been found in various habitats including fields and grass, but it is mainly associated with forest litter, spruce litter and duff. Specimens from Lac Dupar- quet (Abitibi, Quebec) have been collected under snow during winter. Remarks.— External characters of the epi- gynum of G. pedalis are quite subtle and dif- ficult to recognize. Thus, it is not surprising that the female has not been described as it has probably been classified in many collec- tions as ‘undet. Linyphiidaek Genus Lepthyphantes Menge 1866 Lepthyphantes intricatus (Emerton 1911) Figs. 17”19 Microneta complicata Banks 1892:47, pi. 2, fig. 50 (preoccupied by Lepthyphantes complicata Emer- ton 1911); Banks 1916:77, pL 10, fig. 14; Levi & Field 1954:446, figs. 22, 23 (male; not female, = Centromerus cornupalpis). Bathyphantes intricata Emerton 1911:397, pL 3 figs. 7, 7a--d. Centromerus intricatus (Emerton): Freitag et al. 1969:1329. Lepthyphantes intricatus (Emerton): Ivie 1969:6; van Helsdingen 1973:7 (synonymy with Micro- neta complicata Banks 1892); Koponen 1987: 285; West et al 1988:79; Jennings et al. 1988:61; Belanger & Hutchinson 1992:53; Aitchisoe & Sutherland 2000:638, 644; Buddie al 2000: 427-431; Buckle et al. 2001:128; Paquin et al. 2001:18; Paquin & LeSage 2001:98; Paquin & Dupenre 2003:141, figs. 1559-1561. Material examined,- — U.S.A.: Maine: Pis- cataquis County: Soubunge Mountain [45°58'N, 69°12'W], 1 (3 (CNC); Montana: 5 mi N Whitefish [48°24'N, 114°20^W], 1 <3, 2 9 (DJB); New Mexico: Los Alamos [35°5rN, DUPERRE ET AL.— LINYPHIID AND THERIDIID DESCRIPTIONS 151 106°18'W], 1 (? (DJB); New York: Hamilton County: —10 km ESE Brandreth [43°56'N, 74°5rW], 4 (5, 2 $ (RF); CANADA: New Brunswick: Green River 30 mi N Edmunstoe [47°19'N, 68°09'W], 1 d, 3 9 (CNC); New Scotia: Cape Breton Highlands National Park [46°48'N, 60°57'W], 3 (CNC); Cape Breton Highlands National Park Lone Shieling [46°48^N, 60°57'W], 2 d, 2 9 (CNC); Cape Breton Highlands National Park MacKenzie Mountain [46°46'N, 60°49'W], 3 d, 1 9 (CNC); Cape Breton Highlands National Park North Mountain [46°53'N, 60°35'W], 4 d, 14 9 (CNC); Cape Breton Highlands National Park Paqeet Lake [46°48'N, OOMLW], 3 d, 1 9 (CNC); Ontario: Fathom Five National Park Bear Rump Island [45°17'N, SCdO'W], 1 d (CMB); 30 mi E Dryden [49°47'N, 92°45'W], 1 d (CNC); Grundy provincial Park [45°56'N, 80°32'W], 1 9 (CNC); 56 mi N Hurket [49°20'N, 88°53'W], 1 9 (CNC); 20 mi E Kenora [49°49^N, 94°26'W], 2 d (CNC); Long Point Squires Ridge [42°34'N, 80H5'W], 1 9 (CNC); 75 mi W Marathon [48°52'N, 87°35'W], 1 d (CNC); 22 mi S Pickle Lake [5r28'N, 90°12'W], 1 9 (CNC); Raith [48°50'N, 89°56'W], 1 d (CNC); Spem cerville [44°5rN, 75°33'W], 1 9 (CNC); TilL sonburg [42°5rN, 80°44'W], 1 9 (CNC); Turkey Point [42°42'N, 80°19'W], 1 9 (CNC); Walsingham [42°4LN, 80°32'W], 1 d (CNC); Wawa [47°59'N, 84°47'W], 6 d, 25 9 (CNC); Quebec: Gatineau Park King Mountain [45°29'N, 75°52'W], 1 d, 3 9 (CNC); Lac Roddick [46°15'N, 75°53'W], 1 9 (CNC); 24 mi S Ste-Anne^des-Monts [48°52'N, 65°58'W], 1 d (CNC); Lac Dupar= quet Abitibi [48°30'N, 79°13'W], 3 d, 3 9 (CPAD); La Haute-Gaspesie, Parc de la Gas- pesie; Mines Madeleine [48°57'N, 66°0rW], 1 d (CNC); La Riviere-du-Nord, Saint-Hip- polyte, Station biologic Universite de Mon™ treal [45°59'N, 74°00'W], 1 d (CPAD); Am toine-Labelle, Lac Saguay, hwy 117 [46°32'N, 75°09'W], 1 d (CPAD); VaLd^Og ValleeMe- POr, Louvicourt, hwy 117, km. 491 [48°04'N, 77°23'W], 2 d, 1 9 (CPAD); Manitoba: Dau™ phin [5r09'N, 100°03'W], 1 d (CNC); RL ding Mountain National Park North Gate [50°53'N, 100°15'W], 1 d, 4 9 (CNC); RL ding Mountain National Park East Escarpment [50°53'N, 100°15'W], 1 d (CNC); Riverton [50°59'N, 96°59'W], 1 d, Wallace Lake [SrOO'N, 95°2rW], 1 d, 3 9 (CNC); Sas^ katchewan: Anglin Lake [53°44'N, 105°56'W], 8 d, 12 9 (DJB); Besnard Lake [55°25'N, 106°00'W], 4 d (DJB); Alberta: Blood Indian Reserve MSA [49°03'N, 113°42'W], 1 d (DJB); Watertoe National Park [49°04'N, 113°47'W], 7 d, 1 9 (DJB); Watertoe Lakes National Park Cameron Lake [49°0rN, 114°04'W], 2 d, 1 9 (CNC); Bap- tiste Lake [54°45'N, 113°35'W], 3 d, 3 9 (DJB); 19 km N of Calling Lake [55°15'N, 113°12'W], 2 d (DJB); Edmonton [53°33'N, 113°28'W], 1 d, 1 9 (DJB); 25 km sw Rocky Mountain House [52°22'N, 114°55'W], 6 d, 3 9 (HAC, DJB); -20 km s Slave Lake 3 d, 1 9, [55°23'N, 115°13'W], (CMB); British Columbia: Babine Lake [54°45'N, 126°00'W], 1 9 (CNC); Cougar Canyon Ecological Re- serve Vernon [50°09'N, 119°19'W], 1 d (CNC); 15 mi NE Kamloops [50°40'N, 126°19'W], 1 9 (CNC); Piekut Creek [54°27'N, 125°27'W], 1 9 (CNC); Lumby [50°15'N, 118°58'W], 1 d (CNC); Vance Creek Ecological Reserve Vernon [50°17'N, 118°57'W], 1 d (CNC). Description. — Female (n =5): Total length: 2.94 ± 0.23 mm; carapace length: 1.11 ± 0.09 mm; carapace width: 0.89 ± 0.06 mm; carapace smooth, shiny, light orange to or- ange-brown with radiating line and midline shaded with gray; carapace with diffuse gray margins; 2 long erect setae along midline; sternum light orange shaded with gray. Che- licerae light orange to orange-brown, promar- gin with 3 large teeth, retromargin with 4-7 denticles. Cheliceral stridulatory organ not visible with stereomicroscope. Abdomen uni- color, light to dark gray, sparsely covered with long erect setae. Legs light orange to orange- brown, tibia LIV with two dorsal macrosetae; metatarsus I with dorsal trichobothrium, Tml 0.26—0.35, TmlV absent. Epigynum with plate deeply notched, dividing into two protruding lobes; scape long, narrow, slightly widening toward the tip, cochlear present at tip (Figs. 17, 18); spermathecae small, copulatory ducts long, following the folding of the epigynum (Fig. 19). Distribution. — Widespread boreal species (see also Buckle et al. 2001). Habitat. — This common species has been collected in forested habitat, under rocks and logs, mainly in deciduous litter and occasion- ally in coniferous stands. Remarks. — Ivie (1969) was the first to 152 THE JOURNAL OF ARACHNOLOGY place M. intricatus in Lepthyphantes. In his 1973 paper, however, van Helsdingen over- looked Ivie’s paper and erroneously treated it as a new combination. While L. intricatus is similar to other Lep- thyphantes in its general morphology, the form of both palp and epigynum differ suffi- ciently from that of L. minutus, the type spe- cies of Lepthyphantes, and from other species of Lepthyphantes, sens lat., as to very likely justify its placement in a new genus. This transfer, however, is best left for a future re- visional study. Genus Scyletria Bishop & Crosby 1938 Scyletria inflata Bishop & Crosby 1938 Figs. 20, 21 Scyletria inflata Bishop & Crosby 1938:89, pL 7 figs. 72-74; Belanger & Hutchinson 1992:38; Aitchison-Benell & Dondale 1992:224; Buckle et al. 2001:141; Paquin et al. 2001:19; Paquin & Duperre 2003:118, figs. 1233-1235. Material examined. — CANADA: New- foundland: The Arches [50°06'N, 57°40'W], 1 9 (CNC); Nova Scotia: Cape Breton High- lands National Park N of Paquet Lake [46°48'N, 60°4UW], 1 9 (CNC); Cape Breton Highlands National Park Lone Shieling [46°48'N, 60°57'W], 1 d (CNC); Cape Breton Highlands National Park Pleasant Bay [49°49'N, 60°48'W], 1 d (CNC); New Bruns- wick: Priceville 12 mi NW Boiestown [46°3UN, 66°17'W], 1 9 (CNC); Green River 30 mi N Edmunston [47°19'N, 68°09'W], 4 d, 1 9 (CNC); 25 km SW Bathurst [47°37'N, 65°37'W], 1 9 (CNC); Fredericton [45°56'N, 66°40'W], 1 d (CNC); Quebec: Iles-de-la- Madeleine Grosse-Ile [47°37^N, 61°31'W], 1 9 (CNC); St-Methode [48°43'N, 72°24'W], 1 9 (CNC); St-Hippolyte [45°3rN, 73°4rW], 2 S (CNC); Baie-James; Jamesie [49°43'N, 79°17'W], 2 9 (CPAD); Ontario: Spruce Riv- er Sturgeon Lake 42 mi N of Hurkett [50°23'N, 92°30'W], 1 9 (CNC); Manitoba: Duck Mountain National Park Cowan Creek [52°0rN, 100°38'W], 1 9 (CNC); Riding Mountain National Park Swanson spring [50°53'N, 100°15'W], 1 d (CNC); Riding Mountain National Park Jackfish Creek [50°45'N, 100°14'W], 4 d, 2 9 (CNC); Fort Churchill [58°45'N, 94°04'W], 2 d (CNC); Saskatchewan: Lady Lake [52°02'N, 102°37'W], 13 d, 7 9 (DIB); Alberta: Wen- tzel Lake [59°02'N, 114°28'W], 2 c3, 1 9 (UASM); Cypress Hills Provincial Park Elk- water Lake [49°40'N, 110°17'W], 2 9 (CNC); Athabasca [54°43'N, 113°17'W], 2 c3 (DIB); Baptiste Lake [54°45'N, 113°35' W], 5 d, 5 9 (DJB); George Lake 16 km W Busby [53°57'N, 114°06'W], 1 S (AG); Winagami Provincial Park [55°36'N, 116°40'W], 1 9 (DJB); Northwest Territories: Harris River Fort Simpson [bUSUN, 121°20'W], 1 d (CNC). Description. — Female (n = 5): Total length: 1.65 ±0.16 mm; carapace length: 0.68 ± 0.05 mm; carapace width: 0.48 ± 0.05 mm; carapace smooth, shiny, light brown to dark brown with diffused gray patterns along ra- diating lines and midline; carapace margin darker strongly shaded with gray; 4-5 erect setae along midline; sternum dark brown to almost black, shaded with gray. Chelicerae yellow to light brown, promargin with 4-5 large teeth, retromargin with 4-5 denticles. Cheliceral stridulatory organ visible, weak, — 13 ridges. Abdomen unicolor, gray to dark gray, densely covered with short semi-erect setae. Legs light brown to brown, tibia I-III with two dorsal macrosetae and tibia IV with one dorsal seta; metatarsus I with dorsal tri- chobothrium, Tml 0.43-0.56, TmlV absent. Epigynum with plate extended posteriad over the epigastric furrow, wider than long, divided in two rounded blunt prominences (Fig. 20); spermathecae small, widely separated, situat- ed near the anterior margin of the epigynal plate (Figs. 20, 21). Distribution. — This species is widely dis- tributed in northern North America and is also found in New York and North Carolina in the east (Buckle et al. 2001). Habitat. — Scyletria inflata has been re- corded from a wide range of habitats: moss and litter in coniferous forest, moss and algal mats near beaches, riparian vegetation, in co- nifers and river banks. Remarks. — The female illustrated under the name Cephalethus birostrum Chamberlin & Ivie 1947: fig. 21 (now placed in Savignia) appears very similar to S. inflata. Genus Sisicus Bishop & Crosby 1938 Sisicus penifusifer Bishop & Crosby 1938 Figs. 22, 23 Sisicus penifusiferus Bishop & Crosby 1938:62, pi. 2 figs. 12, 13; Levi & Field 1954:448; Drew 1967:172; West et al. 1984:87; Jennings et al. DUPERRE ET AL.— LINYPHIID AND THERIDIID DESCRIPTIONS 153 Figures 20-30. — ^Linyphiid and theridiid structures: 20, 21. Scyletria inflata: 20. Epigynum, ventral view; 21. Spermathecae, dorsal view. 22, 23. Sisicus penifusifer 22. Epigynum, ventral view; 23. Spermathecae, dorsal view. 24-26. Walckenaeria clavipalpis: 24. Carapace of male, lateral view; 25. Palpal tibia of male, dorsal view; 26. Palpus of male, ventral view. 27, 28. Robertus crosbyi: 27. Palpus of male, cymbium and tibia, dorsal view; 28. Palpus of male, ventral view. 29, 30. Thymoites minnesota: 29. Epigynum, ventral view; 30. Spermathecae, dorsal view. Ab- breviations used: AT = Atrium, CD = Conductor, CDu = Copulatory Ducts, E = Embolus, EM = Embolic Mem- brane, ED = Fertilization Ducts, MA = Median Apophysis, MPr = Median Process, MS = Median Septum, PTA = Palpal Tibia Apophysis, RI = Rim, S = Spermatheca, TTA = Theridiid Terminal Apophysis, TP = Tail piece. 154 THE JOURNAL OF ARACHNOLOGY 1988:61; Aitchison-Benell & Dondale 1992:224; Belanger & Hutchinson 1992:38. Sisicus penifiisifer Bishop & Crosby: Buckle et al. 2001:142; Paquin et al. 2001:19; Paquin & Du- perre 2003:148, figs. 1639-1640. Material examined. — CANADA: New- foundland: Lloyds River [48°32'N, 57°13'W], 2 9 (CNC); Trout River w of Badger [48°59'N, 56°02'W], 2 d, 4 9 (CNC); Pasa- dena [49°0rN, 57°36'W], 3 9 (CNC); Nova Scotia: Cape Breton Highlands National Park Franey Mountain [46°4rN, 60°28'W], 2 d, 2 9 (CNC); Cape Breton Highlands National Park MacKenzie Mountain [46°46'N, 60°49'W], 3 9 (CNC); Cape Breton High- lands National Park Lone Shieling [46°48'N, 60°57'W], 1 d, 4 9 (CNC); Cape Breton Highlands National Park Beulack Ball Falls [46°44'N, 60°38'W], 1 d, 1 9 (CNC); Cape Breton Highlands National Park Black Brook [46°44'N, 60°38], 2 9 (CNC); Bridgewater [49°17'N, 122°54'W], 5 d (CNC); New Brunswick: Fredericton [45°56'N, 66°40'W], 2 d, 2 9 (CNC); Kouchibouguac National Park [46°5rN, 64°58'W], 7 9 (CNC); Que- bec: Forillon National Park [48°54'N, 64°2rW], 1 9 (CNC); Gatineau Park King Mountain [45°29'N, 75°52'W], 1 9 (CNC); Cedarville [45°01'N, 72°13'W], 1 9 (CNC); Maskinonge, Saiete-Angele-de-Premont [46°2rN, 73°03'W], 1 9 (CNC); BaieHames, Jamesie, Val Paradis [49°16^N, 79°08'W], 3 9 (CPAD); Pontiac, Les Colliees-de- rOutaouais, 2 km North of Eardley [45°34'N, 76°05'W], 1 d (CPAD); Abitibi-Ouest, Du- parquet [48°30'N, 79°14'W], 1 d, 1 9 (CPAD); Kazabazua, La Vallee-de-la-Gati- neau, Lac Danford [45°57'N, 76°08'W], 1 9 (CPAD); Ontario: Huntsville [45°20'N, 79°13'W], 1 9 (CNC); Ottawa [46°16'N, 75°45'W], 1 9 (CNC); Kinburn [43°23'N, 76°11'W], 5 9 (CNC); Christie Lake [44°49'N, 76°25'W], 1 9 (CNC); Gower [45°08'N, 75°43'W], 1 d (CNC); Manitoba: Turtle Mountain [49°03'N, 100°08'W], 5 9 (CNC); Saskatchewan: Beaver Creek 15 mi S Saskatoon [SCSS'N, 106°43'W], 1 d, 1 9 (CNC); Alberta: Edmonton [53°33'N, 113°28'W], 2 d, 1 9 (CNC); British Colum- bia: Gold Stream Park Vancouver Island [48°28'N, 123°33'W], 1 d, 1 9 (CNC); Burton [49°59'N, 117°53'W], 3 9 (CNC). Description. — Female (n = 5): Total length: 1.08 ± 0.1 1 mm; carapace length: 0.49 ± 0.04 mm; carapace width: 0.33 ± 0.04 mm; I, carapace smooth, shiny, light yellow to light brown, sometimes with diffused gray mark- j ings; 5-6 erect setae along midline; sternum I, light yellow slightly shaded with gray, Che- licerae yellow to light brown, promargin with 4-5 large teeth, retromargin with 5 denticles. Cheliceral stridulatory organ not visible with ' stereomicroscope. Abdomen unicolor, off- i white to light yellow, densely covered with | long semi-erect setae. Legs light yellow to [ light brown, tibia FII with two dorsal macro- j setae and tibia III-IV with one dorsal seta; j| metatarsus I with dorsal trichobothrium, Tml j 0.27-0.38, TmlV absent. Epigynum with atri- j um broad, deep, almost round; mediae septum slender; short, extending one-half length of J atrium; presence of a rim along side of atrium; I' (Fig. 22); posterior end of epigynal plate ris- j ing, recurving into a median triangular pro- j cess; spermathecae elongated reaching jj one-half length of atrium (Figs. 22, 23). j' Distribution. — Widespread species in Can- j ada, from British Columbia to Quebec and the j northern states of USA. 1 Habitat. — Sisicus penifusifer has been ■ mainly collected in forest litter (deciduous and [ coniferous), under logs, in duff and moss. Remarks. — ^The original description of the ^ species includes a description and illustrations of the male, but also a short description of the female which is not mentioned by Platnick 1 (2004). Bishop & Crosby (1938) did not, j however, include any illustrations of the fe- j male epigynum which is shown here for the first time. ’ Genus Walckenaeria Blackwall 1833 | Walckenaeria clavipalpis Millidge 1983 j Figs. 24-26 ' Walckenaeria clavipalpe Millidge 1983:135, figs. j 95, 117, 118. I Walckenaeria clavipalpis Millidge: Paquin et al. j 2000:272; Paquin & LeSage 2001:101; Buckle et | al. 2001:149; Paquin et al. 2001:20; Paquin & j Duperre 2003:125, figs. 1319-1322. Material examined. — CANADA: New- foundland: Gros Morne National Park Stan- ford River [49°4rN, 57°44'W], 1 9 (RPC); Gros Morne National Park east 1 9 Main Riv- er West [49°41 'N, 57°44'W], 3 d (RPC); Port- au-Choix [50°42'N, 57°22'W], 2 d, 1 9 (RPC); Quebec: Reserve faunique des Lau- rentides [47°4rN, 70°51'W], 1 d, 1 9 DUPERRE ET AL.— LINYPHIID AND THERIDIID DESCRIPTIONS 155 (MLC); La Haute-Gaspesie, Parc de la Gas- pesie; Mines Madeleine [48°57'N, 66°0rW], 2 d (CNC). Description.— Mflfe (n = 1): Total length: 2.55 mm; carapace length: 1.05 mm; carapace width: 0.85 mm; carapace dark brown, radi- ating lines and cephalic groove darker brown, midline shaded with black; sternum dark brown, margin darker; cephalic horn absent (Fig. 24). Chelicerae brown, promargin with 3 large teeth and 1 small tooth, retromargie with 2-3 denticles. Cheliceral stridulatory or- gan easily visible, well developed, ~ 1 1 ridges widely separated. Abdomen unicolor, dark gray, densely covered with medium length semi-erect setae. Legs orange, tibia LII with two dorsal macrosetae and tibia IILIV with one dorsal seta; metatarsus I with dorsal tri- chobothriurn, Tml 0.49, TmlV absent. Palpal tibia with four apophyses (3 dorsal, 1 ventral) (Fig. 25); tail piece long, apex billhook shaped; embolus long, coiled, covering 3/4 of the genital bulb length; embolic membrane long, narrow, covering 3/4 of the genital bulb length (Fig. 26). Distribution,— This rare species was known only from Mt Whiteface (New York) (Millidge 1983), the type locality, and the Gaspesie Park (Quebec). Based on these re- cords, the species was tentatively placed in the Alpine- Appalachian category (sensu LeSage & Paquie 2001) by Paquie et al. (2000). The records given here provide additional support for this placement as both localities confirm the alpine and Appalachian affinities of the species. Habitat,— Coniferous forest litter on the summits of North-East North America; —500 m and higher. Family Theridiidae Siindevall 1833 Genus Robertus O. Pickard- Cambrdidge 1879 Robertus crosbyi (Kastoe 1946) Figs. 27, 28 Ctenium crosbyi Kaston 1946:7, fig. 52. Robertus crosbyi (Kaston 1946): Brignoii 1983: 411; Aitchison-Benell & Dondale 1992:219, Be- langer & Hutchinson 1992:81; Paquin et al. 2001: 24; Paquin & Duperre 2003:218, figs. 2435- 2437. Material examined*— CANADA: Quebec: Riviere-du-Loup, lie Verte [47°50'N, 69032' W], 6 6,6$ (CNC); Pontiac, Les Col- lines-de-rOutaouais, Parc de la Gatineau, Lac Brown [45G6'N, 75°55'W], 1 d (CNC); Iles- de-la-Madeleine [47°24'N, 6r47'W], 1 ? (CNC); Manitoba: Riding Mountain National Park [46°4rN, 60°28'W], 1 ? (CNC); to.- Athabasca [54°45'N, 113°35'W], 1 9 (DJB). Description.- — Male (n =3): Total length: 2.31 ± 0.06 mm; carapace length: 1.13 ± 0.07 mm; carapace width: 0.92 ± 0.06 mm; cara- pace smooth, shiny, light brown to dark brown, cephalic groove slightly darker, radi- ating lines shaded with gray; sternum brown shaded with gray, margin darker. Chelicerae dark. Abdomen ueicolor, light gray to dark gray, sparsely covered with long erect setae. Legs light brown. Palpal tibia with one apoph- ysis, cymbium with one lateral apophysis lacking apical setae (Fig. 27). Therediid ter- minal apophysis elongated, narrowing into a hook bearing an additional process; median apophysis bearing two basal points, median apophysis projecting toward the apex of the palpus and curving behind the terminal apophysis; conductor sinuate, projecting api- cally (Fig. 28). Distribution.— In the original description, Kastoe (1946) lists only two records from New York. Based on the few records avail- able, this is a widespread but rarely collected species. Habitat.— crosbyi seems associ- ated with salt marshes, sea wrack, lakeshore litter and mosses in boggy areas. Genus Thymoites Keyserling 1884 Thymoites minnesota Levi 1964 Figs. 29, 30 Thymoites minnesota Levi 1964:467, figs. 74-76; Levi & Randolph 1975:47; Belanger & Hutch- inson 1992:85; Dondale et al. 1997:78; Paquin et al. 2001:24; Paquin & Duperre 2003:224 figs. 2509-2511. Material examined. — U.S.A.: North Da- kota: Bottineau County [county record only], 1 9 (DSU); CANADA: Nova Scotia: Locke- port [43°42'N, 65°07'W], 1 d (CNC); New Brunswick: Sackville [45°55'N, 63°23'W], 1 d (CNC); Quebec: Saint- Jean-sur-Richelieu, Le Haut-Richelieu, CAcadie [45°18'N, 73°20'W], 1 d, 1 9 (CPAD); Riviere-du- Loup, ile Verte [47°50'N, 69°32^W], 4 d (CNC); Iles-de-la-Madeleine [47°24'N, 6r47'W], 1 d (CNC); Ontario: Wawa 156 THE JOURNAL OF ARACHNOLOGY [47°59'N, 84°47'W], 1 $ (CNC); Saskatche^ wan: Lady Lake [52°02'N, 102°37'W], several d, several 9 (MCZ, DJB); SuDenis [52°09'N, 106°07'W], 1 d, 1 ? (DJB); Manitoba: 9 mi W Souris [49°37'N, 100°15'W], 1 9 (CNC); Alberta: Watertoe Lake National Park [49°0LN, 114°04'W], 1 d, 2 9 (CNC); Wee^ tzelLake [59°02'N, 114°28'W], 1 (7 (UASM); 5 mi S Armena [53°07^N, 112°57'W], 1 9 (AG); Baptiste Lake [54°45, 113°35], 1 d (DJB); 7 km W Bittern Lake [53"01'N, 113°03'W], 1 S {AG); Northwest Territories: Yellowknife [62°27'N, 114°2rW], 1 d (CNC); Yukon Territory: Old Crow [67°35'N, 137°53'W], 1 9 (CNC). Description.— (n =5): Total length: 2.18 ± 0.32 mm; carapace length: 0.77 ± 0.01 mm; carapace width: 0.76 ± 0.02 mm; cara- pace shiny, yellow with a tinge of orange, gray to black band along midiine and along the second half of carapace margin; chelicerae light yellow to yellow; sternum yellow with a tinge of orange, margin shaded with gray. Ab- domen unicolor, off-white or off-white with random pattern of white pigment, sparsely covered with decumbent setae; ventral side of abdomen often with a gray to black triangular mark above the spinnerets and two oval mark- ings on each side of the epigynum. Legs yel- low-orange. Epigyeal plate with atrium, deep, divided in two by a complete mediae septum; posterior margin of plate rising and forming a ledge connecting with the mediae septum (Fig. 29); copulatory ducts large, spermathe- cae oval, situated at anterior margin of the epi- gyeal plate (Figs. 29, 30). Distribution. — Widespread northern spe- cies. Habitat. — Thymoites minnesota has been recorded from limestone outcrops, moss, freshwater sedge marshes, salt marshes, and a male and female were collected in a pitfall trap in a cultivated field in LAcadie (Quebec). DISCUSSION Couples in copula are rarely used as a ref- erence for matching the sexes of a given spe- cies. Usually, we rely on the co-occurrence of two sexes in the same sample, or the occur- rence of an unmatched sex in a precise region or habitat in which only one species of a given genus is known. Although these practices may lead to incorrect matches of sexes, most as- sociations were done using these simple meth- ods, resulting in a very low number of mis- associations. In almost all the present cases, at least one couple was collected together in a given sample, thereby, facilitating the cor- ; rect associations between the sexes. Also, the fauna of the Quebec region is well known, therefore limiting the possibilities of a mis- match (see Belanger & Hutchinson 1992; Pa- quin et al. 2001). Knowledge of spider fauna is highly depen- dent on museum collections that gathered |i specimens from several types of surveys and || biodiversity studies. In the present case, ex- !| amination of material preserved in museums j and private collections allowed us to fill some |! gaps for species descriptions. The accessibil- !l ity of such material was essential to this study. ' Museum specimens provide not only precious i information about specimens, but may also be used to orient further collecting to discover an f unknown sex or to collect fresh specimens for [ DNA studies. | ACKNOWLEDGMENTS We wish to express our gratitude to C.D. | Dondale and J.H. Redner for allowing us to i use the material deposited in the CNC and | benefit from their years of work on Linyphi- S idae. We also would like to thank J. Miller for ' suggestions and comments on an earlier draft of this manuscript, L Agnarson for clarifying the terminology in use for theridiid palps, M. Larrivee and R. Pickavance for sharing data j on W. clavipalpis, C. Vink for grammatical j improvements and G. Hormiga and M. Har- | vey for their comments and review. ! 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The genus Walckenaeria Black- wall (Araneae, Linyphiidae). Journal of Arach- nology 11(3): 105-200. Paquin, P, & N. Duperre. 2003, Guide d’ identification des Araignees (Araneae) du Que- bec. Fabreries, Supplement 11. 251 pp. Paquin, R, N, Duperre & R. Hutchinson. 2001, Liste revisee des Araignees (Araneae) du Que- bec. Pp. 5 — 87. In Contributions a la connaiss- ance des Araignees (Araneae) d'Amerique du Nord (P. Paquin & D.J. Buckle eds.). Fabreries, Supplement 10. Paquin, P. & L. LeSage. 2001 [“2000”]. Diversite et biogeographie des Araignees (Araneae) du pare de conservation de la Gaspesie, Quebec. Proceedings of the Entomological Society of On- | tario 131:67-111. Paquin, P., L. LeSage & N. Duperre. 2000. First Canadian records of Tenuiphantes cracens (Zorsch) and Walckenaeria clavipalpis Millidge (Araneae: Linyphiidae), and thirteen new provin- cial records and a confirmation for Quebec. En- tomological News 1 12(4):271-277. Peck, S.B. 1988. A review of the cave fauna of Canada, and the composition and ecology of the invertabrate fauna of caves and mines in Ontario. Canadian Journal of Zoology 66:1197-1213. Platnick, N.L 2004. The world spider catalog, ver- sion 5.0. American Museum of Natural History. http://research.amnh.org/entornology/spiders/ catalogs 1-87/index.htmL ij Saaristo, M.I. & A.V. Tanasevitch. 1996. Redelim- j itation of the subfamily Micronetinae Hull, 1920 and the genus Lepthyphantes Menge, 1866 with I description of some new genera (Aranei, Liny- j phiidae). Bericht des Naturwissens-Medizinisch- i en Verein Innsbruck 83:163-186. I Simon, E. 1926. Les Arachnides de France. Syn- j opsis generale et catalogue des especes franqaises I de Fordre des Araneae. Tome 6, 2eme partie. pp 309-532. Paris. West, R., C.D. Dondale, & R.A. Ring. 1984. A re- vised checklist of the spiders (Araneae) of British Columbia. Journal of the Entomological Society of British Columbia 81:80-98. West, R., C.D. Dondale, & R.A. Ring. 1988. Ad- I ditions to the revised checklist of the spiders (Ar- aneae) of British Columbia. Journal of the En- tomological Society of British Columbia 85:77- 86. ! Manuscript received 12 January 2004, revised II November 2004. j 2006. The Journal of Arachnology 34:159-162 CAPTURE EFFICIENCY AND PRESERVATION ATTRIBUTES OF DIFFERENT FLUIDS IN PITFALL TRAPS Martin H. Schmidt,^ Yaen Clough, Wenke Schulz, Anne Westphalen and Teja Tscharntke: Agroecology, University of Gottingen, Waldweg 26, D-37073 Gottingen, Germany. E-mail: martin. schmidt@zos.unibe.ch ABSTRACT, Pitfall traps are widely used to capture arthropods. The type of fluid employed in the traps can affect size and condition of the catch. Direct comparisons of different fluids allow entomologists to avoid suboptimal solutions, and facilitate comparisons between studies using different fluids. We compared capture efficiency and preservation attributes between five fluids in a field experiment with special respect to spiders (Araneae) and ground beetles (Coleoptera, Carabidae). Catches in pure water, ethanol- water and ethanol-glycerin were less well preserved than in brine or ethylene glycol-water. Brine and ethanol-glycerin showed low capture efficiencies, presumably because their high specific density made arthropods float and thereby facilitated escape. Only the mixture of ethylene glycol and water combined good preservation attributes with high capture efficiency, and therefore represented the best solution. Keywords! Brine, ethanol, ethylene glycol, glycerin, pitfall traps Originally described by Barber (1931), pit- fall traps continue to be among the most wide- ly employed sampling methods for ground- dwelling arthropods, particularly spiders (Araneae) and ground beetles (Coleoptera, Carabidae). Consisting of cups sunk into the ground flush with the surface, pitfall traps are inexpensive, easy to use and operate round- the-clock, resulting in large, species-rich sam- ples (Clark & Blom 1992). A variety of liq- uids are employed to retain, kill and preserve the arthropods. Solutions of formalin and wa- ter were once common, but have been largely abandoned because of health hazards (van den Berghe 1992). Pure water is an alternative (Waage 1985), but mixtures with ethanol, glycerin, ethylene glycol or brine are often preferred because their conservation attributes are presumably better (Holopainee 1992; Teichmann 1994). The use of different pre- servatives also affects sampling efficiency and thereby complicates comparisons between studies. As only a few replicated field studies have been published that compare different preservatives, informed recommendations re- main difficult (Weeks & McIntyre 1997; Lem- ieux & Lindgren 1999). Here, we compared ' Current address: Martin H. Schmidt, Community Ecology, University of Bern, Baltzerstr. 6, CH-3012 Bern, Switzerland. sampling efficiencies and conservation attri- butes of five commonly used fluids in a field experiment. METHODS The preservatives compared in this study were (tap) water, brine (saturated solution of NaCl in water), 2:1 mixture of ethanol and water, 3:1 mixture of ethanol and glycerin, and 1:3 mixture of ethylene glycol (autom^o- bile antifreeze) and water. An unscented de- tergent was added to all liquids to break the surface tension and accelerate wetting and killing of arthropods (Topping & Luff 1995). The traps consisted of 0.2 liter plastic cups with an opening of 7.0cm diameter. They v/ere protected from rain with 25 X 25cm acrylic glass roofs. Two cm from the top of the cup, pieces of 2cm mesh hardware cloth were in- serted to hold off vertebrates (Hall 1991). For- ty of these traps were installed in a fallow on calcareous soil near Gottingen, Germany, in a grid with 5m distance between traps. Seventy ml of each of the five preservatives described above were added to the traps with eight rep- licates in a Latin square design. The traps were operated for seven days starting on 2 May 2003. Upon withdrawal, catches were transferred to polyethylene bottles and stored at 4°C for another week. Then, the volume of remaining liquid was recorded after pouring it 159 160 THE JOURNAL OF ARACHNOLOGY Table 1. — Differences between fluids in the percentage of damaged spiders (with detached legs, palps or opisthosomae), the amount of liquid retrieved, numbers of individuals (N), species (S) or genera (G) captured. Means ± SE. One-way ANOVA, or Kruskal-Wallis ANOVA when variance homogeneity was not met (Hymenoptera N and springtail N). Means preceded by different capitals are significantly different at F < 0.05. Tested variable Water Brine Ethanol-water Ethanol-glycerin Damaged spiders [%] ^28.9 -h 3.7 B9.O -+- 1.6 ^38.1 -h 6.0 ^33.3 ± 4.0 Liquid [ml] M6.8 -+- 1.0 ^52.6 1.4 ^^8.8 -+- 1.2 C29.6 -1- 0.9 Arthropod N 87.8 ■+- 9.7 63.9 -1- 8.6 79.0 7.2 68.5 + 10.8 Spider N M5.8 5.1 C23.5 -1- 3.2 AB40.0 -h 4.8 BC30.O -h 4.8 Spider S 7.5 0.6 5.9 -t- 0.5 6.5 -1- 0.7 6.0 4- 0.4 Ground beetle N 18.0 -+- 2.8 11.6 -1- 2.2 14.3 -1- 1.4 14.0 -+- 3.8 Ground beetle G AB4 4 0.3 ^33 -1- 0.3 A5.I -+- 0.3 B3.8 -h 0.4 Hymenoptera N 4.3 -h 1.0 17.3 -f- 9.8 8.1 -+- 4.4 3.5 1.2 Springtail N AB2.6 -+- 0.6 B1.5 -h 0.5 AB2.8 ± 0.8 A8.3 -H 2.0 Diptera N ^6.5 -+- 1.1 B1.6 -1- 0.5 B3.5 -U 0.7 B2.6 -h 0.7 through gauze, and the arthropods were trans- ferred to 80% ethanol. The condition of the catch was noted with particular attention to signs of decomposing processes, such as the percentage of spiders with detached body parts (legs, palps or opisthosomae). All ar- thropods were identified to order. Spiders were further identified to species, and ground beetles to genera. The number of genera was used as a surrogate of ground beetle species richness (Baldi 2003). The weather during the sampling period was dry and sunny, with an average temperature of 14.7 °C (2.9°-30.0°), mean wind velocity of 3.4 m/s (daily average 1. 4-6.4), and 8.5 hours of sunshine per day (0.5-14.2). Rain (1.5mm) occurred only on the last of the seven sampling days (data sup- plied by Deutscher Wetterdienst, Offenbach, Germany). RESULTS The condition of the samples differed mark- edly between preservatives. The percentage of spiders that had lost body parts was nearly three times as high in water, ethanol-water and ethanol-glycerin as in brine and ethylene-gly- col (Table 1). Additionally, all ethanol-water and two out of eight brine samples developed mold after one week in the refrigerator. Most liquid was retrieved from the traps filled with ethylene glycol and brine, representing 77% and 75% of the initial volume, respectively. Significantly less liquid was retrieved from traps filled with water (67%), ethanol-glycerin (42%) and ethanol-water (13%) at the end of the experiment (Table 1). The overall catch was 1522 spiders (com- prising 1232 Lycosidae, 248 Tetragnathidae, I and 42 individuals from six other families), | 607 ground beetles, 336 Hymenoptera (96% ants, Formicidae), 127 springtails (Collembo- ! la), 122 dipterans and 289 other arthropods. , While the total number of arthropods was not significantly different between liquids, the number of spiders, springtails and dipterans, I and the number of ground beetle genera showed significant treatment effects (Table 1). Thirty-five percent fewer spider individuals were captured in brine and ethanol-glycerin compared to the three remaining liquids. The ! number of ground beetle genera was 25% lower in brine and ethanol-glycerin than in j ethanol-water and ethylene glycol. The num- ^ ber of dipterans was 6.5 times as high in water ; as in ethylene glycol, with intermediate values , in the three remaining liquids, and 7.3 times as many springtails were captured in ethanol- glycerin than in brine and ethylene glycol. DISCUSSION Both preservation attributes and sampling efficiency differed between the fluids com- pared in this study. High losses of volume from ethanol-glycerin and ethanol-water sug- j gest that the ethanol had largely evaporated during one week of exposure. The develop- ment of mold in ethanol-water catches gives additional indication that most of the ethanol, and thereby the conservation attributes of the solution, had disappeared. The mold presum- i ably also kept back an additional part of the remaining liquid, explaining why markedly SCHMIDT ET AL.— SUITABILITY OF FLUIDS IN PITFALL TRAPS 161 Table 1. — Extended. Glycol ^4,35 P x" P M4.7 ± 3.7 9.5 <0.001 ^53.6 ± 3.2 118 <0.001 75.8 ± 7.8 1.1 0.38 ABC37.0 ± 6.1 3.2 0.026 7.3 ± 0.6 1.6 0.18 18.0 ± 2.4 1.1 0.38 ^5.0 ± 0.4 3.6 0.015 8.5 ± 5.8 2.0 0.73 ^0.8 ± 0.4 CEO ± 0.4 9.3 <0.001 10.8 0.028 less than the deployed amount of water could be retrieved, while losses from the pure water traps were minor. Water and brine catches smelled offensive, and water attracted high numbers of dipterans, which are further signs for the decay occurring in these catches. High percentages of spiders had lost body parts in the water, ethanol-water and ethanol-glycerin catches, indicating softening of the cuticle due to decomposition and/or chemical processes. Ground beetles appeared to be less vulnerable to decomposition than spiders (Holopainen 1992), and a certain degree of softening may even be desired because it facilitates mounting of specimens or preparation of genitalia. How- ever, in other ground beetle studies, ethylene glycol was found to be preferable to brine be- cause of its better conservation attributes (Lemieux & Lindgren 1999; Vennila & Ra- jagopal 2000). Therefore, a non-volatile pre- serving component like ethylene glycol is rec- ommended to reliably prevent decomposition in pitfall traps exposed for one week or longer. Numbers of spider individuals and beetle genera were lower in ethanol-glycerin and brine than in the three remaining liquids. Such differing sampling efficiencies can be ascribed to attraction or deterrence by the preservative (Teichmann 1994; Weeks & McIntyre 1997). However, the differences observed in our study suggest an additional mechanism. Ar- thropods usually float in liquids whose spe- cific gravity (SG) is distinctly higher than that of water (SG = 1.0). Reduced capture effi- ciencies in ethanol-glycerin and brine may hence be due to arthropods floating at the sur- face, which facilitated escape of newly trapped individuals falling on top of them. Brine (SG = 1.18-1.20) and glycerin (SG = 1 .26) were the liquids with the highest specific gravities employed in this study, and the spe- cific gravity of the ethanol-glycerin mixture presumably rose to similar values as brine, once most of the ethanol (SG — 0,80) had evaporated. Arthropods may also float in pure ethylene glycol (SG = 1.11), but sink in 1:3 mixtures with water, as has been confirmed with wolf spiders in the laboratory (MHS per- sonal observation). Hence, diluting ethylene glycol with water not only reduces expenses, but may also improve capture efficiency. In conclusion, ethylene glycol had better conservation attributes and/or higher sampling efficiencies for spiders and ground beetles than brine, pure water, or any combination containing ethanol. If there are no specific purposes like DNA preservation (Gurdebeke & Maelfait 2002) or attraction of slugs and snails (to ethanol), mixtures of ethylene glycol and water remain the first choice preservative for pitfall traps. As ethylene glycol is poten- tially hazardous to wildlife, a bitter agent should be added, or physical obstacles em- ployed to avoid access by vertebrates (Hall 1991; van den Berghe 1992). To date, only propylene glycol appears to be a comparably adequate, yet more expensive alternative (Weeks & McIntyre 1997). ACBCNOWLEDGMENTS Peter Gajdos, Matthias Schaefer, George Thomas and an anonymous reviewer gave 162 THE JOURNAL OF ARACHNOLOGY valuable comments on an earlier draft of this manuscript. MHS was supported by the Ger- man National Academic Foundation (Studien- stiftung des deutschen Volkes). LITERATURE CITED Baldi, A. 2003. Using higher taxa as surrogates of species richness: a study based on 3700 Coloep- tera, Diptera, and Acari species in Central-Hun- garian reserves. Basic and Applied Ecology 4: 589-593. Barber, H.S. 1931. Traps for cave-inhabiting in- sects. Journal of the Elisha Mitchell Scientific Society 46:259-266. Clark, W.H. & RE. Blom. 1992. An efficient and inexpensive pitfall trap system. Entomological News 103:55-59. Gurdebeke, S. & J.-P. Maelfait. 2002. Pitfall trap- ping in population genetic studies: finding the right “solution”. Journal of Arachnology 30: 255-261. Hall, D.W. 1991. The environmental hazard of eth- ylene glycol in insect pit-fall traps. Coleopterists Bulletin 45:193-194. Holopainen, J.K. 1992. Catch and sex ratio of Car- abidae (Coleoptera) in pitfall traps filled with ethylene glycol or water. Pedobiologia 36:257- 261. Lemieux, J.R & B.S. Lindgren. 1999. A pitfall trap for large-scale trapping of Carabidae: Compari- son against conventional design, using two dif- ferent preservatives. Pedobiologia 43:245-253. Teichmann, B. 1994. Eine wenig bekannte Konser- vierungsflussigkeit fiir Bodenfallen. Entomolo- gische Nachrichten und Berichte 38:25-30. Topping, C.J. & M.L. Luff. 1995. Three factors af- fecting the pitfall catch of linyphiid spiders (Ar- aneae: Linyphiidae). Bulletin of the British Ar- achnological Society 10:35-38. van den Berghe, E. 1992. On pitfall trapping in- vertebrates. Entomological News 103:149-156. Vennila, S. & D. Rajagopal. 2000. Pitfall trap sam- pling of tropical carabids (Carabidae: Coleop- tera)— evaluation of traps, preservatives and sampling frequency. Journal of the Bombay Nat- ural History Society 97:241-246. Waage, B.E. 1985. Trapping efficiency of carabid beetles in glass and plastic pitfall traps contain- ing different solutions. Fauna Norvegica Series B 32:33-36. Weeks, R.D. & N.E. McIntyre. 1997. A comparison of live versus kill pitfall trapping techniques us- ing various killing agents. Entomologia Experi- mentalis et Applicata 82:267-273. Manuscript received 18 November 2004, revised 29 May 2005. 2006. The Journal of Arachnology 34:163-169 DESCRIPTION AND ECOLOGY OF A NEW SOLIFUGE FROM BRAZILIAN AMAZONIA (ARACHNIDA, SOLIFUGAE, MUMMUCIIDAE) Lincoln S. Rocha: Instituto Butantan, Av. Vital Brazil 1500, CEP 05503-900, Sao Paulo-SP, Brazil. E-mail: linrocha@butantan, gov.br Martinho C. Carvalho: Departamento de Zoologia, ICC-Ala Sul, Universidade de Brasilia — UnB. 70910-900 Brasilia-DF, Brazil. Present address: Dep. de Biologia, Campus do Pici, Universidade Federal do Ceara-UFC, Fortaleza, Brazil ABSTRACT, Three regions of Brazilian Amazonia within the state of Rondonia were searched for the presence of solifuges by means of pitfall traps. A new solifuge, Mummucia taiete, was found at two sites inside Vilhena region, which are “Cerrado” enclaves surrounded by Amazonian forest. This new species is described here and is the seventh from Brazil. Populations of M taiete from these two sites were compared regarding some autoecological parameters. Results showed populations from the two sites are similarly diurnal and male biased, as observed in M. mauryi and M, coaraciandu. On the other hand, these populations differ in density and juvenile/adult ratio. RESUMO. Tres regioes da Amazonia brasileira pertencentes ao Estado de Rondonia foram investigadas quanto a, presenga de Solifugae por meio de armadilhas “pitfall”. Um novo solifugo, Mummucia taiete, foi encontrado dois locals no municfpio de Vilhena, que sao enclaves de Cerrado circundados por floresta Amazonica. Esta nova especie de Solifugae e descrita aqui e e a setima do territorio brasileiro. Popuiapoes de Mummucia taiete desses dois locals foram comparadas quanto a alguns parametros autoecologicos. Resultados indicam que as populaqoes se assemelham pelos habitos diurnos e pelo maior numero de machos, como observado para outras especies como Mummucia mauryi and Mummucia coaraciandu.'Poi outro lado, essas populagdes diferem em densidade e na razao jovens/adultos. Keywords: Solpugida, sun-spiders, camel-spiders, taxonomy The arachnid order Solifugae is distributed over the Americas, Africa and Eurasia, cur- rently comprising about 1,100 species (Har- vey 2002). However, South American distri- butional maps exhibit the presence of only six species in Brazil with large empty areas (Ro- cha & Cancello 2002b). That is mostly due to the lack of studies as shown by recent novel species descriptions (Xavier & Rocha 2001; Martins et al. 2004). We hereby describe a new species from “Cerrado” enclaves, sur- rounded by Amazonian forest, in Vilhena re- gion, situated in southwestern Brazilian Ama- zonia and provide notes on the habitat and abundance of this species. “Cerrado” is the Brazilian savannah-like vegetation, the largest domain after Amazon forest (Silva & Bates 2002). This vegetation occupies a large continuous area in central Brazil and many isolated remnants or “is- lands” in the Atlantic and Amazon forests bi- omes (Silva & Bates 2002; Prance 1996). The Cerrado comprises mainly open vegetation types such as grasslands, opee-scrablands and woodlands. Cerrado has been nominated as an important world biodiversity hotspot due to its diversity and high incidence of endemism (Myers et al. 2000). Solifuges are arthropod predators common in arid and semi-arid lands (Muma 1967). Besides the functional similar- ity between solifuges and spiders, the former resemble scorpions in relation to thermal tol- erance (Cloudsley-Thompson 1962), which enable them to be typical desert inhabitants (Cloudsley-Thompson 1977). They present an extreme wide foraging mode {sensu Piaeka 1966) differentiating them from other arach- nids, including wandering spiders. Solifuges are also prodigal excavators in order to meet various needs (Muma 1966a), and the role played by them in trophic webs of drier eco- systems should not be underestimated. 163 164 THE JOURNAL OF ARACHNOLOGY METHODS We surveyed nine Cerrado enclaves in the state of Rondonia, northwestern Brazil, with three study sites in each region: Vilhena, Pi= menta Bueno and Guajara-Mirim. We used 50 pitfall traps set 100 cm apart from each other at three sites. These traps were 500 ml plastic cups sunk in the soil so that the top of the cup remains at the soil level, with 9 cm diameter, filled with a 200 ml mixture of 9 parts of 80% G.L. alcohol and 1 part of 0.8% formalde^ hyde. The traps were provided with a protec- tive cover consisting of a 20 cm diameter white plastic plate fixed with sticks 5-8 cm height above the traps. The traps were opened for seven days. On all nine study sites we searched both during morning and night hours for arachnids. We also inspected 20 liter pitfall traps (used for herpetological surveys) at all three Vilhena sites plus two R Bueno sites. Night collecting was a one hour search per collector over each of 12 quadrats (30 X 5 meters) set up on each site. The surveys were conducted on the fol- lowing dates: Vilhena (30 August-14 Septem- ber 1999) and P. Bueno (11-29 July 2000) during the dry season and (southern hemi- sphere) winter, whereas the G. -Mirim surveys were conducted during summer (wet season, 12-21 January 2001). The material studied is deposited in the Institute Butanta, Sao Paulo, Brazil (IBSP) and the Universidade Nacional de Brasilia, Brasilia-DF, Brazil (UNB). Cheliceral teeth are named according to Muma (1951), where sizes of cheliceral teeth are ordered with Roman numerals, such that I is larger than II, etc. The telotarsal spination formulae are used as in Maury (1982). The nomenclature of podomeres is in accordance with Shultz (1989). TAXONOMY Family Mummuciidae Roewer 1934 Genus Mummucia Simon 1879 Mummucia taiete new species Figs. 1-8 Types. — Holotype male, Vilhena (12° 4U39"S, 60°05'53"W), State of Rondonia, Brazil, 30 August-14 September 1999, M.C. Carvalho (IBSP). Paratypes: 4 males and 5 fe- males, same data as holotype (IBSP); 1 male and 1 female, same data as holotype (UNB). Etymology. — The specific name “taiete” is from Tupi, ancient South American Indian language, and means “very good teeth”, and is to be treated as a noun in apposition. Diagnosis. — Mummucia taiete is the only species of Mummuciidae with the last three distal pleurites almost totally dark-brown (Figs. 1, 2). Description.- — Male: Coloration in 80% ethanol: Prosoma. Propeltidium white, with a central brown blotch, with the dorsal grooves between propeltidium and lateral lobes dark- brown. Ocular tubercle dark-brown. Peltidium white. Parapeltidium, mesopeltidium and me- tapeltidium similar to opisthosomal tergites. Chelicerae pale-brown, three longitudinal white stripes on ectal face joined dorsally above the fondal teeth. Pedipalps and legs brown, with slightly darker ends. Malleoli as in Fig. 3. Opisthosoma: Lateral borders of ter- gites white, with wide dark-brown stripe on the central half, which is darker near the pos- terior border of the tergites. Brown bifid setae with brown sockets when they are in white area of the tergites, and white sockets when in the dark-brown area. Pleurites (Figs. 1, 2) with dark-brown and pale-brown areas, being the three last distal almost entirely dark- brown. Sternites pale-brown, lateral borders slightly darker. First and second post-spirac- ular sternites with about 15 brown spots which include the sockets of some bifid bristles. All vestitural bristles and bifid bristles are trans- lucent pale-brown. Morphology and chaetotaxy: Prosoma. Propeltidium with some scattered bifid setae, slightly wider than long (Table 1) and sepa- rated from lateral lobes by dorsal grooves. Oc- ular tubercle prominent with bifid setae ante- riorly oriented. Distance between the two eyes slightly more than one eye diameter. Peltidium narrow, with a transverse row of bifid setae. Parapeltidium smooth. Mesopeltidium wider than long, semicircle-shaped, with several bi- fid setae on posterior border. Metapeltidium wider than long, with several bifid setae. Che- licerae (Figs. 4, 5): stridulatory apparatus on mesal face with six parallel narrow grooves; ectal face with several short bristles and sev- eral setae, bifid or acuminate; movable finger with one anterior, one intermediate and one principal tooth, graded in size from distal to proximal II, III, 1. Fixed finger with three an- terior teeth, one intermediate and one princi- pal tooth, graded in size from distal to prox- ROCHA & CARVALHO— DESCRIPTION AND ECOLOGY OF NEW SOLIFUGE 165 Figures 1-8. — Mummucia taiete new species, holotype male unless stated otherwise: 1. Left pleurites; 2. Left pleurites of male paratype; 3. Left malleolus V; 4. Left chelicera, mesal view; 5. Left chelicera, ectal view; 6. Left leg IV; 7. Right chelicera, ectal view, of female paratype; 8. Genital operculum of female paratype. imal III, V, II, IV, 1. The three anterior teeth placed in a slightly prominent projection so that they are not completely aligned with other teeth. Six ectal fondal teeth graded in size from distal to proximal I, II, V, III, IV, VI, being third and sixth ectal teeth from distal to proximal occasionally vestigial or absent. Three mesal fondal teeth graded I, II, III, the first distal separated from the others by a di- astema; flagellum (Figs. 4, 5) thin, translucent drop-shaped vesicle, laterally flattened and with a longitudinal ectal opening. Pedipalp: tarsi immovable, without spines, densely cov- ered by differentially sized bifid bristles, with some very long setae on basitarsi and tibiae (about twice the length of pedipalpal tibia). Legs: with several differentially sized bifid bristles and some bifid setae. Some very long 166 THE JOURNAL OF ARACHNOLOGY Table 1. — Morphometric characters of Mummucia taiete, in millimeters (except propeltidium length/ width ratio). Morphometric character Holotype Males {n = 5) Female paratype Females {n = 5) Total length 9.20 8.00-9.20 12.00 10.00-12.00 Cheliceral length 2.50 1.80-2.50 2.80 2.50-2.80 Cheliceral width 0.90 0.70-0.90 1.00 0.84-1.00 Propeltidium length 1.72 1.30-1.72 1.63 1.56-1.63 Propeltidium width 2.03 1.60-2.03 2.25 2.19-2.25 Propeltidium length/width ratio 0.85 0.81-0.85 0.72 0.71-0.72 Pedipalpus length 6.88 6.50-6.88 6.40 5.84-6.40 Leg I 5.75 5.75-6.00 5.50 5.20-5.50 Leg IV 11.88 9.20-11.88 8.96 8.00-8.96 dorsal setae (about twice the length of basi~ tarsus IV). Leg I thin, without claws and spines. Legs II and III: tibiae with a distal pair of ventral spines; basitarsus with three retro- lateral spines and 1.1.2 ventral spines; telotar- si two-segmented with 2.2.272.2 ventral spines. Leg IV (Fig. 6): tibia with an anterior row of 1.1.1 ventral bifid spines and a distal pair of ventral spines; basitarsus with 1.1. 1.2 ventral spines; telotarsi three-segmented, with 2.2.2/272.2 ventral spines. Opisthosoma. Ter- gites with rounded borders, covered by bifid setae and bifid bristles. Sternites densely cov- ered by bifid bristles. Posterior border of sec- ond post-spiracular sternite with a row of about 50 ctenidia. Morphometric dimensions in Table 1, Female: Similar to male, but with the fol- lowing particular features. Morphology and chaetotaxy: Prosoma. Eyes separated by twice their diameter. Chelicerae (Fig. 7): stridulatory apparatus with 8 parallel narrow grooves. Movable finger with 1 anterior, 1 intermediate and 1 principal teeth graded in size from distal to proximal I, III, 11. Fixed finger with three anterior teeth, one intermediate and one prin- cipal tooth, graded in size from distal to prox- imal III, V, I, IV, IL Five ectal fondal teeth graded in size from distal to proximal I, II, IV, III, V. Genital operculum prominent, fan- shaped, round-bordered, with central longitu- dinal opening (Fig, 8). Morphometric charac- ters in Table 1. Remarks. — According to Maury (1998), it is impossible to reliably distinguish the genera of Mummuciidae, so the most conservative decision is to consider the new species de- scribed here as belonging to the typical genus Mummucia until more precise information about the taxonomy and phylogeny of the group become available. The same decision was adopted in the description of Mummucia mauryi Rocha 2001 (Xavier & Rocha 2001) and M. coaraciandu Pinto-da-Rocha & Rocha ' 2004 (Martins et al. 2004). j The cheliceral dentition of M. taiete and M. ^ coaraciandu are quite similar; remarkably this j feature is generally species-specific in Soli- ' fugae (Maury 1984; Rocha 2002). These two | species can only be distinguished by the col- ; oration of the posterior three pleurites, which are almost totally dark-brown in M. taiete whereas they are predominantly pale-brown in M. coaraciandu. \ The color pattern of the pleurites are known ' for four species of Mummuciidae and may be , considered as species-specific. The mummu- ' ciids Metacleobis fulvipes Rower 1934 (Rocha & Cancello 2002a), Mummucia mauryi (Xa- vier & Rocha 2001), Mummucia coaraciandu \ (Martins et al. 2004), Gaucha fasciata Mello- ; Leitao 1924 (Maury 1970; Rocha, unpub. data) present distinct patterns of pleurite col- , oration, with slight intraspecific variation. Un- ' fortunately, pleurite coloration has not been : studied in the remaining 15 species of Mum- i muciidae. j It is not known if the coloration difference between M. coaraciandu and M. taiete sug- gests that they are two actual and reproduc- tively isolated species. Alternatively, this dis- tinction could be merely a geographical polymorphism due to low gene flow between the populations (putative species), since they are isolated by about 1300 km and by several rivers, ecosystems and other barriers. Any- I ROCHA & CARVALHO— DESCRIPTION AND ECOLOGY OF NEW SOLIFUGE 167 Table 2.- — -Characteristics of two Cerrado en- claves near Vilhena, Rondonia state, Brazil v/here M. taiete was collected (data from Colli, G.R. in prep=). Cerrado “stricto sensu” = a woodland; Cer- rado field = open scrubland. Cerrado Area “stricto Antropic Cerrado (Km^) sensu” field field “forest” Site 1 73.15 83.9% 8.2% 7.6% 0.3% Site 2 10.06 85.2% 9.4% 5.4% 0% how, pleurite coloration is indeed diagnostic for M. coaraciandu and M. taiete and fits at least the definition of the typological concept of a species. ECOLOGICAL RESULTS AND DISCUSSION We recorded the occurrence of M taiete at only two study sites in the Vilhena region with densities of 29.65 individuals/m Vday at site 1 and 17.52 iedividuals/niVday at site 2. Both sites are Cerrado enclaves with similar vege- tation cover (Table 2). Total number of indi- viduals captured: n = 66; n 2 == 39. These figures show both the high density and mobility of this species. The observed fre- quency distributions of the number of M. taie- te caught in 50 pitfall traps fit the Poisson dis- tribution in both areas (Kolmogorov-Smimov one-sample test, K = 0.368, P < 0.01, n = 50) showing a random distribution pattern of M. taiete movements. We found no solifuges in the other seven study sites. Data from other arachnids will appear elsewhere (Carvalho et al. in prep.). We found a decreasing activity in M. taiete in the early morning hours when we arrived at the sites between 7:30-8:30 a.m., when many live individuals were found in the 20 1 pitfall traps. This suggests that activity may begin at dawn (no individuals were seen dur- ing the eight surveys) (Maury 1984; Xavier & Rocha 2001; Marties et al. 2004). The sex- ratio (operational) was biased toward males (X" = 4.694, d.f. = 1, P < 0.05; with Yates correction; data from both areas were pooled after perform heterogeneity chi-square analy- sis: x^ = 0.29, d.f. — 1, N.S.; Chi-square anal- ysis with Yates correction Ho = 1:1; site #1: X^ ^ 3,368, d.f. - 1, N.S.; site #2: ^ 0.941, d.f. — 1, N.S.), probably due to higher male 100% n 80% - 60% - 40% - 20% - 0% H L J ^ »- ■ - - , site 1 site 2 Figure 9.— Operational sex-ratio of M. taiete is male biased with Yates correction = 4.694, d.f. = 1, P < 0.05, n = 36; data from both areas were pooled after perform heterogeneity chi-square anal- ysis), Mature individuals were captured in two Cer- rado enclaves near Vilhena, Rondonia state, Brazil. sitei = 19, n site 2 = 17). Capture effort: 50 pitfall traps (9 cm diameter, 1 meter apart) were exposed for one week. activity (Miima 1975, 1980; Xavier & Rocha 2001; Martins et al. 2004) or to early male maturity (Muma 1966b). The occurrence of one or both of these factors could explain the male biased sex-ratio we found in the pitfall trap data (Fig. 9). We tested the null hypoth- esis of an 1:1 juvenile/adult ratio and Site 1 seems to have a young, rapidly expanding population (x^ with Yates correction test: x^ = 11.045, d.f. - 1, P < 0.001, n = 66) and site 2 a relatively slow growth population (x^ ” 0.41, d.f. - 1, N.S., N = 39) (Fig. 10). This difference in the dynamic of these populations may reflect constraints due to the smaller size of Site 2. This survey was conducted at the end of the dry season, thus juveniles would 100% . 80% - 60% - 40% - 20% - 0% -- sitei site 2 Figure 10. — Proportion of juveniles in M taiete populations from two Cerrado enclaves near Vil- hena, Rondonia state, Brazil. Individuals were caught by 50 pitfall traps (9 cm diameter, 1 meter apart) exposed for one week in each site, (y^ with Yates correction test Ho = 1:1, site 1: = 11.045, d.f. = i, F < 0.001, n = 66; site 2: y^ = 0.41, df. = 1, N.S., n 3i,2 = 39). 168 THE JOURNAL OF ARACHNOLOGY reach maturity after the onset of the rainy sea- son. Both sites have white sandy soil and this may facilitate colonization by M. taiete due to the excavation behavior of Solifugae. Similar soil features appear to be favorable for M. coaraciandu, another solifuge from Brazilian Cerrado, which is more abundant in sites with sandy soils (Martins et al. 2004). Accordingly to these observations, Dean & Griffin (1993) suggest that sandy soils are a key factor de- termining the occurrence of many solifuge species. However, two other R Bueno sites (Sites 12 & 12A) also have white sandy soils and may potentially be colonized by M. taiete. Prance (1996) observes that savannahs on white sandy soil have a high incidence of en- demism among plant species. All three G. Mi- rim sites are rock outcrops and seem unlikely to be colonized by M. taiete (Colli unpub. data, for details of vegetation structure and physical characteristics for all nine sites; we use here the same site labels as Colli). Mum- mucia taiete may be a distribution-restricted species with high density populations in Cer- rados of white sandy soils with a particular combination of Cerrado types allowing direct incidence of solar radiation in patches of bare soil surface. It is worth noting that these two sites are the most similar regarding vegetation cover compared to the other seven sites sur- veyed (Colli et al, unpub. data). The data presented here coupled with the above-mentioned characteristics derived from the literature suggest that Solifugae deserve further investigation as a key predator of ar- thropod fauna in the Brazilian Cerrados, a poorly known biodiversity hotspot. ACKNOWLEDGMENTS We thank Dr. Guarino R. Colli, coordinator of the project “Structure and dynamics of the biota in natural and antropic Cerrado frag- ments, lessons to Conservation Biology” which provided logistic support, sponsored by “PROBIO”, a biodiversity program of the Brazilian Ministry of Environment (MMA) and Brazilian Council of Scientific Research (CNPq). M.C. Carvalho received a post-doc- toral fellowship from CNPq. L.S.R. would like to thank Dr. Antonio Brescovit for pro- viding optical equipment used in the descrip- tion of the new species and Dr. Eduardo Na- varro for assiatance with Tupi grammar. LITERATURE CITED Cloudsley-Thompson, J.L. 1962. Lethal tempera- tures of some desert arthropods and the mecha- nism of heat death. Entomologia Experimentalis et Applicata 5:270-280. Cloudsley-Thompson, J.L. 1977. Adaptational bi- ology of Solifugae (Solpugida). Bulletin of the British Arachnological Society 4:61-71. Dean, W.R.J. & E. Griffin. 1993. Seasonal activity patterns and habitats in Solifugae (Arachnida) in the southern Karoo, South Africa. South African Journal of Zoology 28:91-94. Harvey, M.S. 2002. The neglected cousins: what do we know about the smaller arachnid orders? Journal of Arachnology 30:357-372. Martins, E.G., V. Bonato, G. Machado, R. Pinto- Da-Rocha & L. S. Rocha. 2004. Description and ecology of a new species of sun spider (Arach- nida: Solifugae) from the Brazilian cerrado. Jour- nal of Natural History 38:2361-2375. Maury, E.A. 1970. Sobre la presencia de Gaucha fasciata Mello-Leitao 1924 en la Argentina. Phy- sis 79(29):357-362. Maury, E.A. 1982. Solifugos de Colombia y Ven- ezuela (Solifugae, Ammotrechidae). Journal of Arachnology 10:123-143. Maury, E.A. 1984. Las familias de Solifugos Ame- ricanos y su distribucion geografica (Arachnida, Solifugae). Physis, Buenos Aires 42:73-80. j Maury, E.A. 1998. Solifugae. Pp. 560-568. In Bio- diversdidad de Artropodos Argentinos. (J.J. Mor- rone & S. Coscaron, eds). Ediciones SUR. La Plata. Muma, M.H. 1951. The arachnid order Solpugida | in the United States. Bulletin of the American ! Museum of Natural History 97:35-141. ! Muma, M.H. 1966a. Burrowing habits of North i American Solpugida (Arachnida). Psyche, Cam- , bridge 73:251-260. ! Muma, M.H. 1966b. The life cycle of Eremobates | durangonus (Arachnida: Solpugida). Florida En- ! tomologist 49:233-242. ! Muma, M.H. 1967. Basic behavior of North Amer- ican Solpugida. Florida Entomologist 50:115- 123. Muma, M.H. 1975. Long term can trapping for pop- ulation analyses of ground-surface, arid-land arachnids. Florida Entomologist 58:257-270. Muma, M.H. 1980. Comparison of three methods for estimating solpugid (Arachnida) populations. j Journal of Arachnology 8:267-270. [ Myers, N., R.A. Mittermeier, G.C. Mittermeier, G.A.B. Fonseca & J. Kent. 2000. Biodiversity ; hotspots for conservation priorities. Nature 403: ! 853-859. Pianka, E.R. 1966. Convexity, desert lizards, and | spatial heterogeneity. Ecology 47:1055-1059. Prance, G.T 1 996. Islands in Amazonia. Philosoph- ' ROCHA & CARVALHO^-DESCRIPTION AND ECOLOGY OF NEW SOLIFUGE 169 ical Transactions of the Royal Society of London B 351:823-833. Rocha, L.S. 2002. Solifugae. Pp. 439-448. In Am- azonian Arachnida and Myriapoda (Adis, J. ed). Sofia. Pensoft Publishers. Rocha, L.S. & E.M. Cancello. 2002. Redescription of Metacleobis fulvipes Roewer from Brazil (So- lifugae, Mummuciidae). Journal of Arachnology 30:104-109. Rocha, L.S. & E.M. Cancello. 2002b. South Amer- ican Solifugae: New records, occurrence in hu- mid forests and concurrence with termites. Newsletter of the British Arachnological Society 93:4-5. Shultz, J.W. 1989. Morphology of locomotor ap- pendages in Arachnida: evolutionary trends and phylogenetic implications. Zoological Journal of the Linnean Society 97:1-56. Silva, J.M.C. & J.M. Bates. 2002. The Cerrado: a tropical savannah hotspot. Bioscience 52:225- 233. Xavier, E. & L.S. Rocha. 2001. Autoecology and description of Mummucia mauryi, a new species from Brazilian semi-arid caatinga. Journal of Arachnology 29:127-134. Manuscript received 11 May 2004, revised 1 Feb- ruary 2005. 2006. The Journal of Arachnology 34:170-175 TWO NEW PURSE-WEB SPIDERS OF THE GENUS ATYPUS (ARANEAE, ATYPIDAE) FROM KOREA Seung-Tae Kim, Hun-Sung Kim, Myung-Pyo Jung, Joon-Ho Lee: Entomology Program, School of Agricultural Biotechnology, Seoul National University, Seoul 151-921, Korea. E-mail: stkim2000@hanmail.net Joon Namkung: 933 Wabu-eup, Namyangju 472-902, Korea ABSTRACT. Two new species of the genus Atypus, Atypus sternosulcus new species from Andong and A. suwonensis new species from Suwon, are newly described from Korea. Keywords: Purse-web, Atypus sternosulcus, Atypus suwonensis, Asia, taxonomy Spiders of the genus Atypus are known as purse-web spiders as they construct silk-lined tunnels in the ground that extend above the soil surface, usually against the vertical side of a tree or rock. The tube is covered with sand and debris and is difficult to detect. Males are active mostly from June to August, and females then guard their egg sacs during August and September (Schwendinger 1990). Mating takes place inside the tube and the spi- ders stay together for several months, after which the male dies or is eaten by the female (Im & Kim 2000). Females of these primitive spiders may five for several years. Worldwide, 19 species of the genus Atypus have been recorded from the United States, Europe and south-east Asia (Platnick 2004). Three species have been described from Ko- rea: A. coreanus Kim 1985, A. magnus Na- mkung 1986 and A. quelpartensis Namkung 2001 (Kim 1985; Namkung 1986, 2001). Atypid spiders are characterized by a male sternum with marginal ridges, a short, straight and spike-like embolus, a straight conductor and a distally widened vulva with bulbous or pyriform receptacula and with two lateral patches of pores on the genital atrium (Gertsch & Platnick 1980). Kraus & Baur (1974) utilized various taxonomic characters to distinguish between the European species, such as the segmentation of the posterior spin- nerets, features of the patellar membrane, morphology of sigilla I and IV, and the male palpal conductor, palpal furrow and male metatarsal spines. Schwendinger (1990) noted and discussed the granular texture on the male chelicerae and front legs, and the cymbial pit for distinguishing species. METHODS This paper describes two new atypid spi- ders, Atypus sternosulcus from Girae stream, Andong, Gyeongsangbuk-do and Atypus su- wonensis from Suwon, Gyeonggi-do in Korea. Male specimens of all Korean atypid spiders were examined, and we reviewed published descriptions to compare taxonomic characters (Kim 1985; Yaginuma 1986; Schwendinger 1990; Chen & Zhang 1991; Song et al 1999; Namkung 1986, 2001). The specimens ex- amined were: A. karschi: 1 3, 15 August 1995, Kumamoto, Kyusu, Japan, Kim and J. Namkung; A. coreanus: 1 A, 17 May 1982, Mt. Ungil, Gyeonggi-do, Korea, J. Namkung; A. magnus: 1 6,1 June 1986, Jigdong, Gyeonggi-do, Korea, J. Namkung; A. quel- partensis: 1 3, 4 July 1989, Jungmun, Jeju island, Korea, J. Namkung (Table 1). The external morphology of the specimens was observed and illustrated utilizing a ste- reoscopic microscope, and metric characters were measured with an ocular micrometer. All measurements are given in mm. Leg measure- ments are given in the order of femur, patella -i- tibia, metatarsus and tarsus, in parentheses. Abbreviations used in this paper are: AER = anterior eye row; PER = posterior eye row; AME = anterior median eye; ALE = anterior lateral eye; PME = posterior median eye; PLE = posterior lateral eye; MOQ = median oc- 170 KIM ET AL.— PURSE- WEB SPIDERS FROM KOREA 171 Table 1. — Comparisons of taxonomic characters of male of some Asian atypid spiders. Species PLS, number of seg- ments Leg patella, retrolateral face Sigilla I Sigilla IV Conductor, bend in upper comer Furrow on palpal femur Spines on metatar- sus IV Granules on femur I(n) Distri- bution A. karschi 4 pigmented marginal oval strong shallow present domed (domed) Japan China Taiwan A. heteropthecus 4 white remote from margin oval medium shallow absent domed (domed) China A. coreanus 4 pigmented marginal oval strong deep present domed (domed) Korea A. magnus 4 pigmented marginal oval strong deep present smooth (smooth) Korea Russia A. quelpartensis 4 white remote from margin oval strong deep present smooth (smooth) Korea A. sternosulcus 4 white marginal suboval strong shallow present domed (smooth) Korea A. suwonensis 4 white remote from margin oval strong shallow present smooth (smooth) Korea ular quadrangle; ALS = anterior lateral spin- nerets; PMS = posterior median spinnerets; PLS == posterior lateral spinnerets. The spec- imens studied are lodged in the deposited in the Laboratory of Insect Ecology, Seoul Na- tional University, Seoul, Korea (SNU). TAXONOMY Family Atypidae Thorell 1870 Genus Atypus Latreille 1804 Atypus sternosulcus new species Figs. 1-7 Type. — Holotype male, Giran stream, An- dong, Korea (36°3ror' N, 128°50'3rU), 3 June 2003, S.T. Kim, H.S. Kim, M.P. Jung and J.H. Lee (SNU). Etymology — The specific name is a com- bination of ‘stern’ from Greek meaning breast and Latin ‘sulcus’, a furrow or pit, referring to the pit on the sternum of the male holotype. Diagnosis. — This species is similar to A. karschi Donitz 1887 in general appearance, but differs as follows: the coloration of the abdomen; the chelicera with 12 teeth and 2 denticles on promargin (Fig. 6), instead of 13 in A. karschi (Yaginuma 1986, fig. le) and their alignment; the shape of vestigial and pit- ted anterior sternal sigilla (Fig. 5) (A. karschi: Yaginuma 1986, fig. lb); and the shape of up- per lateral edge of conductor (Figs. 2-4) (A. karschi: Yaginuma 1986: fig. Ip). Description. — Male: Total length 16.5 (in- cluding chelicerae and excluding spinnerets). Body length 11.7. Carapace 5.1 long, 4.7 wide. Abdomen 6.6 long, 4.2 wide. Chelicerae 5.0 long, 1.7 wide. Endite 2.7 long, 1.9 wide. Labium 0.3 long, 1.1 wide. Sternum 3.4 long, 3.1 wide. AER 1.2, PER 1.3. Leg measure- ments; I: 14.9 (4.7, 4.5, 3.5, 2.2); II: 12.7 (3.7, 3.9, 3.0, 2.1); III: 12.0 (3.4, 3.5, 2.8, 2.3), IV: 15.2 (4.5, 4.4, 3.5, 2.8); pedipalp 5.8 (2.4, 2.3, -, 1.1). Carapace lustrous and quadranglar, reddish brown with narrow black border, gently nar- rowed posteriorly; cephalic region dark red- dish brown and elevated, lateral margins with black stripes; thoracic region reddish brown, fiat, gently rounded on both sides, emargin- ated posterior margins bearing conspicuous pleurites. Median groove broad weakly U- shaped and deeply imprinted, behind midpoint of carapace, occupying about 1/6 of carapace width at that point; cervical and radial furrows distinct and deeply imprinted (Fig. 1). Eye tu- bercle black, 0.7 long and 0.8 wide. AER and PER slightly recurved when viewed from above. Eye sizes and interdistances: AME 0.26, ALE 0.23, PME 0.17, PLE 0.20; AME- AME 0.20, AME-ALE 0.07, AME-PLE 0.23, PME-PME 0.63, PME-PLE almost touching, PLE-PLE 0.83, ALE-PLE-PME contiguous. MOQ 0.5 long, front width 0.7 and back width 0.9 (Fig. 7). Chelicerae dark reddish brown, well developed with deep longitudinal furrow at base of retrolateral surface; 12 teeth and 2 172 THE JOURNAL OF ARACHNOLOGY Figures 1-7. — Atypus sternosulcus new species: 1. Body, dorsal view; 2. Left palp, prolateral view; 3. Left palp, retrolateral view; 4. Left palpal bulb, ventral view; 5. Sternum, left half; 6. Left chelicera, retrolateral view; 7. Eyes, dorsal view, Upper corner of conductor indicated by arrow. denticles on promargin of cheliceral furrow (Fig. 6); prolateral and retrolateral surface with distinct granular texture. Sternum dark reddish brown with ridges at margins. Four pairs of sigilla deeply imprinted. Anterior sig- ilia vestigial and deeply pitted between labium and endite. Sigilla sizes and interdistances: I 0.3 long and 0.2 wide, almost touching at mar- gin, II 0.2 long and 0.3 wide, 0.6 from margin, III 0.6 long and 0.3 wide, 0.5 from margin, IV 0.7 long and 0.5 wide, 0.4 from margin; I- I 1.1, II-II 1.5, III-III 1.3, IV-IV 0.5, LII 0.6, II-III 0.3, III-IV 0.2 (Fig. 5). Endite and la- bium dark reddish brown. Legs 4123, dark reddish brown and armed with short spines; prolateral side of femur I with granular tex- KIM ET AL.— PURSE=WEB SPIDERS FROM KOREA 173 Figures 8-14. — Atypus suwonensis new species. 8. Body, dorsal view; 9. Left palp, prolateral view; 10. Left palp, retrolateral view; 11. Left palpal bulb, ventral view; 12. Sternum, left half; 13. Left chelicera, retrolateral view; 14. Eyes, dorsal view, Upper corner of conductor indicated by arrow. ture; tarsi light reddish brown and pseudoseg- mented; retrolatral membranous area on pa™ tella I without pigments; trichobothria in two distally convergent rows on basal 2/3 tibiae L IV (left): 4 + 6, 5 + 6, 5 + 5, 6 + 6 and in single row on distal half of metatarsi LIV: 3, 3, 3, 4. Abdomen suboval and dull grayish brown; dorsal scutum (3.8 long and 2.0 wide) blackish gray enclosing dull blackish gray ter™ gite (Fig. 1); lung patches light grayish brown. Spinnerets light blackish gray. ALS 0.5; PMS 0.9; PLS 4 segmented: basal joint 0.6, median 0.7, subapical 0.7, apical 0.5. Pedipalp dark reddish brown; palpal cymbium without basal pit; palpal femur without furrow; bulb small and globe=hke; embolus short and stout, 174 THE JOURNAL OF ARACHNOLOGY spine-shaped; upper distal corner of conductor slightly bent upwards (Figs. 2-4). Female: Unknown. Distribution. — Korea (Giran stream, An- dong, Gyeongsangbuk-do). Ecological remarks. — The sole specimen was collected in a pitfall trap near a stream beside a hillock. Atypus suwonensis new species Figs. 8-14 Type. — Holotype male, Seodung-dong, Su- won, Korea (37°15'41"N, 126°59'16"E), 24 June 2000, T.W. Kim (SWU). Etymology. — The specific name is an ad- jective referring to the type locality. Diagnosis. — This species is similar to Aty- pus coreanus Kim 1985 in general appear- ance, but differs as follows: the alignment of cheliceral promarginal teeth (Fig. 13) (A. co- reanus: Kim 1985: 6, figs. 1-3); the shape of upper lateral edge of conductor (Figs. 9-11) (A. coreanus: Kim 1985: p. 6, fig. 7; Na- mkung 2001; p. 25, figs. b-c). Description. — Male: Total length 14.7 (in- cluding chelicerae and excluding spinnerets). Body length 10.6. Carapace 5.0 long, 4.9 wide. Abdomen 5.6 long, 3.6 wide. Chelicerae 4.8 long, 1.8 wide. Endite 2.8 long, 1.7 wide. Labium 0.3 long, 1.1 wide. Sternum 3.7 long, 3.4 wide. AER 1.3, PER 1.4. Leg measure- ments; I: 13.7 (4.6, 4.2, 3.0, 1.9); II: 12.0 (3.7, 3.6, 2.5, 2.2); III: 10.8 (3.2, 3.1, 2.6, 1.9), IV: 14.2 (4.1, 3.9, 3.6, 2.6); pedipalp 5.3 (2.1, 2.1, 1.1). Carapace lustrous and quadranglar, reddish brown with narrow black border, gently nar- rowed backward; cephalic region dark reddish brown and elevated, margined by black stripes; thoracic region reddish brown and flat, gently rounded on both sides, emarginated posterior margins bearing conspicuous pleu- rites. Median groove weakly W-shaped and deeply imprinted, positioned at about 2/3 of carapace length, occupying about 1/5 of car- apace width at that point; cervical and radial furrows distinct and deeply imprinted (Fig. 8). Eye tubercle black, 0.8 long and 0.7 wide. AER and PER slightly recurved from the above. Eye sizes and interdistances: AME 0.26, ALE 0.23, PME 0.17, PLE 0.18; AME- AME 0.20, AME- ALE 0.10, AME-PLE 0.26, PME-PME 0.73, PME-PLE almost touching, PLE-PLE 1.02, ALE-PLE-PME contiguous. MOQ 0.6 long, front width 0.7 and back width i 1.0 (Fig. 14). Chelicerae dark reddish brown, ! well developed with deep longitudinal furrow at base of retrolateral surface; 12 teeth on pro- margin of cheliceral furrow (Fig. 13); prola- j teral and retrolateral surface with distinct granular texture. Sternum dark reddish brown with ridges at margins. Four pairs of sigilla j deeply imprinted. Sigilla sizes and interdist- ! ances: I 0.6 long and 0.4 wide, 0.7 from the margin, II 0.3 long and 0.4 wide, 0.6 from the j margin. III 0.6 long and 0.4 wide, 0.5 from | the margin, IV 0.7 long and 0.6 wide; I-I 0.9, II-II 1.4, III-III 1.4, IV- VI 0.4, I-II 0.06, ILIII 0.03, IILIV 0.09 (Fig. 12). Endite and labium dark reddish brown. Legs 4123, dark reddish brown and armed with short spines; prolateral side of femur I with granular texture; tarsi light reddish brown and pseudosegmented; re- trolatral membranous area on patella I without pigments; trichobothria in two distally con- vergent rows on basal 2/3 tibiae I-IV (left): 6 + 7, 6 + 6, 5 + 5, 6 + 6 and in single row on distal half of metatarsi I-IV: 4, 3, 3, 7. Ab- domen suboval and dull black; dorsal scutum (4.0 long and 2.1 wide) blackish brown en- i closing yellowish brown tergite (Fig. 8); lung j patches light grayish brown. Spinnerets light blackish gray. ALS 0.5; PMS 0.9; PLS 4 seg- mented: basal joint 0.6, median 0.8, subapical 0.7, apical 0.6. Pedipalp dark reddish brown; palpal cymbium without basal pit; palpal fe- i mur without furrow; bulb small and globe i like; embolus short and stout spine-shaped; upper distal corner of conductor conspicuous- j ly bent upwards (Figs. 9-11). j Female. — Unknown. Distribution. — Korea (Suwon, Gyeonggi- do). Ecological remarks. — The sole specimen was collected in a pitfall trap. ACKNOWLEDGMENTS We thank Mr. T.W. Kim, Sungshin Woman’s University, who collected and provided the specimen of A. suwonensis. We thank Dr. R.J Raven and an anonymous referee for their j critical review of the manuscript. This work ! was funded by Ministry of Environment (Pro- ject 2004-09001-0012-0) and supported by the Brain Korea 21 Project, Korea. LITERATURE CITED Chen, Z.E & Z.H. Zhang. 1991. Fauna of Zhejiang: i Araneida. Zhejiang Science and Technology , Publishing House, Zhejiang, 356 pp. I KIM ET AL.-~~PURSE-WEB SPIDERS FROM KOREA 175 Gertsch, WJ. & Ni. Platnick. 1980. A revision of the American spiders of the family Atypidae (Ar- aneae^ Mygalomorphae). American Mnseum Novitates 2704:1-39. Im, M.S. & S.T Kim. 2000. Field Guide of Korean Spiders. Konkuk University Press, Seoul, 285 pp. Kim, J.P. 1985. A new species of genus Atypus (Ar- aneae: Atypidae) from Korea. Korean Arachnol- ogy 1(2): 1-6. Kraus, O. & H. Baur. 1974. Die Atypidae der West- Palaarktis: Systematik, Verbreitung und Biologic (Arach.: Araneae). Abhandlungen aus der Natur- wissenschaften Verein, Hamburg (N.F.) 17:85- 116. NamkuEg, J. 1986. A new species of the genus Aty- pus Latreille, 1804 (Araneae: Atypidae) from Korea. Acta Arachnologica 35:29-33. Namkung, J. 2001. The Spiders of Korea. Kyo-Hak Publ. Co. Seoul, Korea. 647 pp. Platnick, N.L 2004. The World Spider Catalog, Ver- sion 5.0. American Museum of Natural History, online at http://research.amrih.org/entomology/ spiders/catalog/index. html [cited 15 December 2004] Schwendinger, P.J. 1990. A synopsis of the genus Atypus (Araneae, Atypidae). Zoologica Scripta 19:353-366. Song, D.X., M.S. Zhu & J. Chen. 1999. The Spiders of China. Hebei Science and Technology Pub- lishing House, Shijiazhuang, 640 pp. Yaginuma, T. 1986. Spiders of Japan in Color (new ed.j. Hoikusha Publishing Company, Osaka. 305 pp. Manuscript received 12 November 2003, revised 3 February 2005. 2006. The Journal of Arachnology 34:176-185 COPULATORY BEHAVIOR AND WEB OF INDICOBLEMMA LANNAIANUM FROM THAILAND (ARACHNIDA, ARANEAE, TETRABLEMMIDAE) Matthias Burger: Natural History Museum, Department of Invertebrates, Bernastrasse 15, CH“3005 Bern, Switzerland. E-mail: burgermatthias@hotmaiLcom Alain Jacob: University of Bern, Department of Conservation Biology, Baltzerstrasse 6, CH-3012 Bern, Switzerland and Natural History Museum, Department of Invertebrates, Bernastrasse 15, CH-3005 Bern, Switzerland. Christian Kropf: Natural History Museum, Department of Invertebrates, Bernastrasse 15, CH-3005 Bern, Switzerland. ABSTRACT. The present study reports for the first time on the behavior prior to, during and after the copulation of a member of the haplogyne spider family, Tetrablemmidae and describes the web of this species. Prior to copulation, male and female of Indie oblemma lannaianum from Thailand sometimes avoided each other or the female scared the male away, apparently by vigorous vibrations of her body. When first copulations were initiated, they lasted from 1.21 to 3.8 h with an average of 2.25 ± 0.71 h {n — 17). Some females accepted a second male for mating 3-9 days after first mating. There was no significant difference between the duration of first and second copulations but significantly more trials were needed to induce the second copulations. In the copulatory position, the male was inverted and faced in the same direction as the female. He seized the female’s opisthosoma with apophyses on his chelicerae which fit into grooves on a female’s ventral plate in this way building a locking mechanism during copulation. The pedipalps were inserted alternately. The web of I. lannaianum consisted of a longish narrow sheet, which was made of many short threads forming a zigzag pattern and additional long oblique threads overdrawing the sheet and functioning as signal threads. Keywords: Haplogynae, copulation, locking mechanism, courtship Spiders show an impressive array of vari- ous copulatory patterns and positions (e. g., Gerhardt 1933; von Helversen 1976; Foelix 1996; Huber & Eberhard 1997). Although there have been numerous studies on the mat- ing of a variety of spiders, the copulatory be- havior of many species still remains unknown. Bristowe and Gerhardt described the mating behavior of several entelegyne species, and of haplogynes including members of the families Scytodidae, Pholcidae, Segestriidae, Dysderi- dae and Oonopidae (Bristowe 1929-1931; Gerhardt 1926-1930, 1933). In addition to Gerhardt’s comprehensive work (reviewed by Huber 1998a), the most detailed descriptions of reproductive biology and copulatory mech- anisms in haplogynes are given for the family Pholcidae (Uhl 1993a, 1993b, 1998; Uhl et al. 1995; Huber 1994, 1995, 1997, 1998b, 2002; Huber & Eberhard 1997; Yoward 1998; Seng- let 2001; Schafer & Uhl 2002). However, studies on the copulatory behavior of many haplogynes are still missing. This is especially true for members of the family Tetrablemmi- dae as their behavior has not yet been ob- served in detail. Tetrablemmids are armored spiders which mainly live as soil-dwellers in the litter habitat of tropical rain forests (Shear 1978; Lehtinen 1981; Burger 2005). They show a character- istic pattern of abdominal sclerotization, and the carapace or the chelicerae of males are often strongly modified (Shear 1978; Lehtinen 1981; Schwendinger 1989, 1994), The family Tetrablemmidae is systematically placed as sister group of the Dysderoidea (Coddington & Levi 1991; Platnick et al. 1991). The diminutive size (body length less than 2 mm) of most tetrablemmids and the fact that many species are hard to find make behavioral observations difficult. In the present study we investigate the mating behavior of a tetra- 176 BURGER ET AL.— COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 177 blemmid spider and compare the copulation duration of first and second copulations. We also describe avoidance behaviors of males and females, female aggressive behavior prior to copulation, and we provide details of the web of this species. METHODS Specimens of the tetrablemmid Indicoblem- ma lannaianum Burger 2005 were collected in the primary evergreen hill forest of Doi Suth- ep, 1600 m elevation, near Chiang Mai (18°48' N, 98°59' E) in northern Thailand, from 11-21 July 2003 by sieving. Types are deposited in the Geneva Natural History Mu- seum, Switzerland and the Natural History Museum of Bern, Switzerland. Indicoblemma lannaianum inhabits the middle humid leaf lit- ter layer alongside little streams. Sixty-four females and forty-four males were caught and kept singly in little snap cap glass jars (3 cm diameter and 5 cm height) with ground gyp- sum, which was moistened every day by one or two drops of water so that the air humidity was almost saturated. The spiders were fed Collembola (Folsomia Candida Willem). All spiders were mature when collected and thus their mating history is unknown. For the mating behavior studies, the spiders were used one or two days after they were collected. Copulations were observed with a binocular microscope (Wild M3; 6.4 X, 16X, 40 X) and partially photographed using a dig- ital camera (Canon Power Shot G2). All sixty- four females and forty-four males were used for the mating observations. For each pairing, the male was carefully removed from his glass jar with a paint-brush and placed into the fe- male’s jar. The spiders were left together for 30-60 min. If no copulation was initiated dur- ing that time, the male was put back in his own glass jar and another male was offered to the female after a recovery period of 30 min. The first palpal insertion by the male was tak- en as the beginning of copulation. The end of copulation was defined as the moment when the spiders physically separated. Post-copula- tory behavior was observed for 15 minutes in each case. Eleven females which had copulated once were given the opportunity to mate a second time. One day after a female had copulated in the lab for the first time, different males were offered to her one after another (1-3 males per day). When a female copulated for the second time no more males were offered. If a female did not accept a second male within 14 days, the procedure was stopped and the female was considered as single mated. Only the six females that had mated twice were included in the statistical comparison of first and second copulations. To compare first and second copulations, two-tailed Wilcoxon Sign-Rank-tests for paired data were applied. We tested for a difference in copulation du- ration and for a difference in the number of trials needed until successful first and second copulations took place. We used nonparamet- ric statistics because most of our groups to test differed significantly from a normal distribu- tion [duration of second copulations (Shapiro Wilk W-test; W = 0.775621, P < 0.0351); number of trials for the first (W — 0.639893, P < 0.0014) and second copulations (W = 0.763674, P < 0.0270)]. Averages are given ± standard deviation. RESULTS Pre-copulatory behavior. — When a male (Figs. 1, 2) was placed in the female’s jar (Figs. 3, 4), both spiders usually walked around and appeared to meet each other ac- cidentally. A male who came into contact with the threads of a female’s web appeared to commence searching for the female. The fe- male then reacted by turning towards him. No male was ever seen filling his pedipalps with sperm prior to copulation. Sometimes males and females stood immobile at a distance of about 3-5 mm facing each other for 8-10 minutes before contacting (Fig. 5). When the spiders met frontally, the male either grasped the female directly and took the copulatory position (Figs. 6-10) or he pushed her back and upwards with his front legs. In the latter case both of them palpated each other with their front legs for a few seconds before the male took the copulatory position by going backwards along the ventrum of the female (Fig. 11). When the male approached the fe- male from the side or from behind he jumped at her and gripped her back with his chelic- erae. If the female kept running around, the male grasped her opisthosoma with his legs (Fig. 12). Before mating, the male crept under her from the side and took the copulatory po- sition (Figs. 6-10). Locking mechanism. — When the male was 178 THE JOURNAL OF ARACHNOLOGY Figures 1-8. — Indicoblemma lannaianum, male and female. 1. Male, dorsal view; 2. Male in web, anterior dorsal view, arrow pointing to signal thread; 3, 4. Female, dorsal view; 5. Male (left side) facing the female prior to copulation; 6. Male (left side) and female in copula, arrow pointing to embolus of male pedipalp; 7. Male (left side) and female in copula; 8. Male (right side) and female in copula. BURGER ET AL.— COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 179 Figures 9—12. — Indicoblemma lannaianum prior and during copulation, 9. Male (left side) and female in copula; 10. Male (prosoma on the left side, ventral view) grasping the female (ventral plates in optical cut, dorsal view) with apophyses on his chelicerae during copulation (asterisks); 11. Male (left side) and female palpating each other with the front legs prior to copulation before the male moves into the copu- latory position (arrow); 12, Male (left side) grasping the female’s opisthosoma prior to copulation before creeping under her (arrow) and taking the copulatory position. Scale bars = 0.5 mm (9, 11, 12), 0.2 mm (10). 180 THE JOURNAL OF ARACHNOLOGY 4.0 3.5 I 3.0' CQ 2.5' S 2.0H 3 Q. 8 1.5' 1.0 r • • • • t first copulations 13 c 18- i 16- Q. 8 12- 10 ■ C0 8 8- 6 4H 2 0 • first copulation o second copulation o o o . 15 '1 ' ^ I — I I 3 4 5 6 female-no. Figures 13-17. — Copulation durations, trials for successful copulations and web of Indicohlemma Ian- naianum. 13. Copulation duration of first copulations (n = 17); 14. Copulation duration of first and second copulations (n = 6; numbers beneath box-plots indicate female-no. corresponding to Fig. 15); 15. Number of trials needed for each female to induce successful first and second copulation; 16, 17. Web. Abbrevi- ations: sh = horizontal sheet, th = oblique signal threads. in the copulatory position, he only used apophyses on his chelicerae to grasp grooves on a ventral plate of the female’s opisthosoma (asterisks in Fig. 10). The tips of his legs were not used to hold the female’s opisthosoma (Figs. 6-9). In this position the male often bent and stretched his legs in quick succes- sion. Avoidance behavior. — We observed three types of behavior that appeared to be avoid- ance behavior: (i) Sometimes, when put to- gether, the two spiders ran quickly in different directions, (ii) After facing each other and staying motionless for several minutes, the fe- male or the male (or both) sometimes turned around and walked away without copulating, (iii) After having palpated each other with the front legs (once or several times in quick suc- cession), the spiders sometimes walked away from each other. Female aggressive behavior, — We some- times observed that when a male approached a female (or had faced her for some time or had already come into contact with her) she seemed to scare him away by vigorous vibra- tions of her body, especially the front legs. Such apparent female aggressive behavior was observed 18 times. BURGER ET AL.— COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 181 Copulatory behavior. — Seventeen out of 64 females accepted a male for copulation (first copulations). Forty-one trials were nec- essary to induce successful first copulations. Those first copulations lasted from 1.21 to 3,8 h (median = 2.06; lower quartile =1,7; upper quartile = 2.74; Fig. 13). Eleven of the 17 females that had copulated once were tested to see if they would accept a second mate. Six of them accepted a second male for copulation 3-9 days after first copulation (average 4.3 ± 2.3). The second copulations lasted from 0.36 to 5.75 h (median = 1.53; lower quartile = 0.48; upper quartile = 2.89; Fig. 14). There was no significant difference between the du- ration of first and second copulations (T = -4.5; df = 5; P < 0.44; Fig. 14). For the females that mated twice, eight tri- als were necessary to induce successful first copulations whereas 37 trials totally were nec- essary to induce successful second copula- tions (Fig. 15). The numbers of trials needed to induce successful first and second copula- tions significantly differed (T = 10.5; df = 5; P < 0.031; Fig. 15). In the copulatory position (Figs. 6-10) the male was inverted and faced in the same di- rection as the female. His legs did not touch the female’s opisthosoma. The male only used his chelicerae to grasp the female (see “lock- ing mechanism” above). After copulation had begun, all his legs stayed motionless and slightly bent. The male rested on the patellae of his legs III and IV and on the anterior dor- sal part of his opisthosoma. During copulation both spiders remained calm and almost motionless. From time to time the female could take a few steps and slightly change her position. The palps were inserted alternately (arrow in Fig. 6, Fig. 10). Towards the end of copulation, some females started to run around or tried to knock off the male using their legs. In all cases, the male sprang away from the female when copulation was finished by turning a somersault. Post-copulatory behavior. — After their separation, the spiders mostly walked away from each other. In some cases, they remained close together and showed intense self-groom- ing. The male often ran his pedipalps through his chelicerae. When the spiders met a second time by walking around after copulation, they either both ran quickly in different directions and kept out of each other’s way, or they pal- pated each other with the front legs for a few seconds before they separated again (see “avoidance behavior” above). Sometimes the female scared away the male (see “female ag- gressive behavior” above). Uncertain cases. — In some pairings, after the male had seized the female’s opisthosoma with his chelicerae (or had already taken the copulatory position), the female ran until the male loosened his hold and let her go. Some- times the female pushed her legs against the male’s legs and she seemed trying to knock the male off. Such male-female interactions were observed five times and lasted from 1.5- 10.62 minutes (average 5.33 ± 3.42). These uncertain cases were not counted as copula- tions and consequently not included for the analyses shown above. Web characterization and function. — The web inhabited by /. lannaianum consisted of a longish narrow horizontal sheet which was made of many short threads forming a zigzag pattern (sh in Figs. 16, 17). The sheet should be attached along dry leaves and small branches on the ground. Long oblique addi- tional threads functioning as signal threads overlaid the sheet and were partly connected with it (th in Figs. 16, 17). The spiders often stayed in contact with these threads (arrow in Fig. 2, Fig. 16). They immediately reacted to a prey touching the threads and successfully captured it. No particular retreat for the spider was observed. DISCUSSION Pre-copulatory behavior. — The main rea- sons for a male to court before copulation are probably to identify himself as a mate of the same species, to avoid being mistaken for prey or to stimulate the female and convince her of his quality (Eberhard 1985, 1996; Foe- lix 1996; Huber 1997; Huber & Eberhard 1997; Bartos 1998). In many haplogyne spi- ders the male courtship behavior prior to cop- ulation is restricted to abdominal vibrations or simple leg and palp movements (Bristowe 1929; Gerhardt 1929; Dabelow 1958; Uhl et al. 1995; Huber 1994, 1995, 1998b, 2002; Huber & Eberhard 1997; Bartos 1998; Senglet 2001). In pholcids, males often keep on mov- ing their pedipalps during copulation (copu- latory courtship) (Gerhard 1927; Uhl et al. 1995; Huber 1994, 1995, 1998b, 2002; Huber & Eberhard 1997; Schafer & Uhl 2002). Other 182 THE JOURNAL OF ARACHNOLOGY forms of male courtship behavior are tapping or jerking the female’s web (Bartos 1998), cutting threads of the female’s web (Uhl et al. 1995; Bartos 1998; Senglet 2001), or spread- ing the chelicerae (Jackson & Pollard 1982). In exceptional cases male pholcids even per- form gustatorial courtship (Huber 1997). Males of /. lannaianum showed several be- haviors that could have some sort of pre-cop- ulatory courtship function. Some males pal- pated the female and pushed her backwards or the male bent and stretched his legs in quick succession before copulation started. The grasping of the female’s opisthosoma by che- liceral apophyses could also have a courtship function (Huber 1999). In 5% of the cases, distinctive female aggressive behavior was observed: the male may have been ready to copulate, but the female prevented any further interaction by scaring him away. This behav- ior is a striking indication for pre-copulatory female choice as those females often mated with other males afterwards. Similar female aggressive behavior in haplogynes was ob- served in certain pholcids prior to copulation (Huber 1994, 1995; Huber & Eberhard 1997; Bartos 1998) or after copulation (Bartos 1998; Senglet 2001). The fact, that significantly more trials were needed for successful copu- lation if females had already mated (Fig. 15) could indicate that females become choosier with increasing copulation number as sug- gested by Schafer & Uhl (2002) for Pholcus phalangioides Fuesslin, 1775. Nevertheless copulations also seemed to take place without pronounced pre-copulatory courtship as some- times males just grasped the females force- fully with their chelicerae and took the cop- ulatory position directly. Locking mechanism. — A cheliceral lock- ing mechanism (by apophyses or modified cheliceral hairs) during copulation in haplo- gyes was reported for certain scytodids (Da- below 1958) and pholcids (Huber 1994, 1995, 1998b, 2002; Uhl et al. 1995; Senglet 2001). Lehtinen (1981) proposed that some tetra- blemmids use apophyses on their chelicerae to grasp corresponding grooves on a ventral plate of the female during copulation. The present study confirms this suggestion for the first time by live observations. Copulation duration. — Copulation dura- tion is quite variable among different spiders. Elgar (1995) suggested that haplogynes have shorter copulations because of the simplicity of their copulatory apparatus. However, stud- ies have shown that the copulatory organs of some haplogynes are in fact quite complex (e. g., Uhl 2000; Burger et al. 2003), and little is known about the copulation duration of many haplogynes. A few of the haplogyne spiders investigated so far copulate longer than one hour (Gerhardt 1927-1929, 1933; Uhl 1993a; Bartos 1998; Senglet 2001; Schaf- er & Uhl 2002). The longest copulation known for a haplogyne spider was longer than 5 hours in the oonopid Silhouettella loricatula (Roewer 1942) (sub Dysderina loricata) (Bristowe 1930). It seems obvious that the function of a pro- longed copulation in general cannot be ex- plained by prolonged sperm transfer only (Eberhard 1985; Suter & Parkhill 1990), Dur- ing long copulations, apparent risks are ac- cepted, which should favor brief mating (e, g. increased danger from predators or interrup- : tion by another male or by the female before j sperm transfer is completed; Eberhard 1996). Copulation duration may correlate with the ! amount of transferred sperm (Engqvist & Sauer 2003) and/or with fertilization success j (Andres & Rivera 2000) but Schafer & Uhl (2002) showed that longer copulations do not indicate higher fertilization rates in their study. The prolonged copulation duration in /. lannaianum could be explained by sperm i competition hypotheses such as sperm prece- dence (Suter & Parkhill 1990; Elgar 1998) , and/or by hypotheses of sexual selection by cryptic female choice (the male could initiate ‘ processes in the female during copulation which increase his chances of siring her off- spring; Eberhard 1985, 1996). The prolonged ' copulation could also serve as mate guarding. By guarding the female, a male can restrict : access to females from other males (Sillen- Tullberg 1981 ; Wynn & Vahed 2004) and con- sequently guard and protect his own trans- ferred ejaculate (Schofi &. Taborsky 2002). These are all hypotheses yet to be tested but it appears that /. lannaianum could be a good species for testing these ideas. The second copulation of female no. 4 (Fig. 14) seemed to last disproportionately long when compared to the other second copula- tions, Excluding it from the analysis would result in a trend towards a decreased duration i of the second copulations. Schafer & Uhl I BURGER ET AL.— COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 183 (2002) suggested that shorter second copula- tions resulted from a stronger conflict of in- terest between the sexes over paternity in sec- ond matings. Copulatory position.-^-The evolution of the mating positions in spiders was discussed by von Helversen (1976). The most plesio- morphic copulatory position is the one taken by most mygalomorphs and haplogyees like Oonops (Gerhardt 1930), Segestria or Filis- tata (Gerhardt 1928, 1929): the male ap- proaches the female frontally, pushes her body back and upwards and inserts his pedipalps simultaneously or alternately (von Helversen 1976). Among different spider groups, a change from the plesiomorphic to a derived copulatory position took place coevergeetly: the male rests on the ventral side of the female and both partners face in the same direction. This position is seen in the oonopids SilhoueF tella (Bristowe 1930) and Xestaspis (Gerhardt 1933). Indie oblemma lannaianum takes the de- rived copulatory position. The behavior be- fore some copulations could be a remnant of the one shown by spiders that take the pie- siomorphic position: the male pushes the fe- male back and they palpate each other with the front legs for a few seconds (Fig. 11) be- fore he goes backwards along the veetrum of the female and takes the derived copulatory position (Figs. 6-10). Web characterization and function,— The web of L lannaianum resembles the only web known so far of a tetrablemmine spider. In BrignoUella vulgaris Lehtieen 1981, it is a small dense sheet attached along the surface of large dry leaves (Lehtieen 1981). Schwee- dinger (1989) described the web constructed by Perania nasuta Schwendinger 1989, a member of the subfamily Pacullinae. It is a loose sheetweb in which the spider hangs up- side down at eight. ACKNOWLEDGMENTS We are most grateful to Dr. Peter Schwee- dinger from the Geneva Natural History Mu- seum, who discovered the species and gave us important information about the type locality in Thailand. The tap to Thailand was financed by a travel grant of the Swiss Academy of Sciences (SAS) which is greatly acknowl- edged. We sincerely thank Prof. Dr. Claus We- dekind for helpful comments on the manu- script. AJ. thanks the Natural History Museum of Bern and the Swiss National Sci- ence Foundation for financial support. LITERATURE CITED Andres, J.A. & A.C. Rivera. 2000. Copulation du- ration and fertilization success in a damselfly: an example of cryptic female choice? Animal Be- haviour 59:695-=703. Bartos, M. 1998. Quantitative analyses of male courtship behaviour in Pholcus phalangioides (Fuesslin, 1775) (Araneae, Pholcidae). In: Seiden P.A., editor. Proceedings of the 17* European Colloquium of Arachnology, Edinburgh 1997. 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Male Gryllus bima- culatus guard females to delay them from mating with rival males and to obtain repeated copula- tions. Journal of Insect Behavior 17:53-66. Yoward, P.J. 1998. Sperm competition in Pholcus phalangioides (Fuesslin, 1775) (Araneae, Pholci- dae)— shorter copulations gain higher paternity re- ward than first copulations. In: Selden P.A., editor. Proceedings of the 17* European Colloquium of Arachnology, Edinburgh 1997. British Arachnolog- ical Society, Burnham Beeches, p 167-170. Manuscript received 27 August 2004, revised 23 February 2005. 2006. The Journal of Arachnology 34:186-193 PREY CHOICE BY NESTICODES RUFIPES (ARANEAE, THERIDIIDAE) ON MUSCA DOMESTICA (DIPTERA, MUSCIDAE) AND DERMESTES ATER (COLEOPTERA, DERMESTIDAE) Marcelo N. Rossi and Wesley A.C. Godoy: Departamento de Parasitologia, IB, Universidade Estadual Paulista (Unesp), Botucatu, Sao Paulo, Brazil. E-mail: rossilife@ibb.unesp.br ABSTRACT. Nesticodes rufipes is widely distributed in tropical and subtropical regions, being strongly associated with humans. However, few behavioral and ecological studies have investigated interspecific interactions between these spiders and insects of medical and veterinary importance. Here, we have in- vestigated prey choice by N. rufipes when two different prey species, Musca domestica and Dermestes ater, were offered simultaneously. We also quantified the capture of these prey types by this predator in a poultry house and analyzed the association between prey-choice with physical characteristics of the prey. Finally, we discuss whether there is an antagonistic intraguild interaction in such a system composed of N. rufipes (top predator), D. ater (predator of larvae of M. domestica and prey of N. rufipes) and M. domestica {N. rufipes'' prey). We found that Musca domestica were more abundant than D. ater in N. rufipes webs in the poultry house. Spiders given a choice of adults of M. domestica plus adults of D. ater, and also on adults plus larvae of M. domestica, preyed more on adult flies than on the other prey types. This preference was probably associated with the lesser mass and shorter lengths of adult flies. Our experiments demonstrated that the predation impact of N. rufipes on D. ater is low when compared to M. domestica. This result provides evidence that an antagonistic interaction between these predators does not occur, suggesting that they are in fact acting either synergistically or additively on M. domestica prey. Keywords: Prey selection, housefly, spider predation, poultry house Many spiders eat a wide variety of prey species (usually insects), and they usually pre- sent a sedentary foraging behavior (Wise 1993), suggesting that selection for habitat, not for prey, should be the rule. However, sev- eral prey capture specializations can be seen (Greenstone 1979; Riechert & Luczak 1982; Uetz 1992; Alderweireldt 1994; Onkonbury & Formanowicz 1997; Nyffeler 1999; Toft 1999), and some may have been an important influence on the evolution of insect defense behavior (Uetz 1990). It has been recognized that the choice of habitat (patch) in spiders is of primary importance through its effect on feeding rates, growth and reproduction (Riechert 1981; Morse & Stephens 1996). Nevertheless, once in a feeding patch, spiders typically are confronted with an array of po- tential prey species. Indiscriminate feeding is not advantageous because prey vary enor- mously in quality due to toxicity or nutrient content. Thus, active prey selection by spiders serves to find the optimal compromise be- tween three “nutritional goals”: to maximize energy intake, to balance nutrient composition of the body, and to minimize toxin consump- tion (Toft 1999). Prey selection has been defined by Hassell (1978) as follows: “Preference for a particular prey is normally measured in terms of the de- viation of the proportion of that prey attacked from the proportion available in the environ- ment.” A common form of prey specialization shown by spiders is on prey size (Uetz 1992), evidenced by some studies comparing the prey of spiders to that available in the envi- ronment (Uetz & Hartsock 1987; Uetz 1990). Spiders are major components of the gen- eralist predator guild that characterizes inter- mediate trophic levels in many terrestrial sys- tems (Moulder & Reichle 1972; Manley et al. 1976; Spiller & Schoener 1996). Theory sug- gests that prey suppression by multiple pred- ator species can lead to a variety of outcomes depending on the nature of the predator-pred- ator interaction. Predator effects can be en- 186 ROSSI & GODOY— PREY CHOICE BY NESTICODES RUFIPES 187 hanced when predators interact either addi- tively or syeergistically (Finke & Denno 2002). Antagonistic interactions, on the other hand, result in diminished prey suppression, either because one predator disrupts the for- aging behavior of another predator (Moran et al. 1996), or consumes the other predator (Po- lis & Holt 1992; Rosenheim 1998; Wise & Chen 1999). Nesticodes rufipes (Lucas 1846) (Araeeae, Theridiidae) (referred to as Theridion rufipes in references) is widely distributed in tropical and subtropical regions, extending to temper- ate zones, and these spiders construct irregular webs with a disordered aspect (Gonzalez 1989). Its exact distribution is not easy to de- termine, since it is strongly associated with humans (Downes 1988; Gonzalez & Estevez 1988; Gonzalez 1989). Behavioral and eco- logical studies considering predation by M ru- fipes are scarce. Fox (1998) highlighted the strategic importance of these spiders to the natural control of Aedes aegypti (Diptera, Cu- licidae), since the spiders incorporate a para- lyzing substance in the webs, which paralyzed the mosquitoes through contact, increasing their capture efficiency. Barreto et al. (1987) also mentioned the importance of N. rufipes as predators of Rhodnius prolixus (Hemiptera, Reduviidae), Musca domestica (Linnaeus 1758) (Diptera, Muscidae) has a cosmopolitan distribution and high syeanthropic indices (Smith 1986; Fer- reira & Lacerda 1993), being also of consid- erable medical and veterinary importance (Harwood & James 1979; Smith 1986; Levine & Levine 1991), This species lives in human dwellings, poultry houses, supermarkets and garbage, growing on a wide variety of sub- strates such as food and vertebrate excrement (Axtell & Arends 1990; Bowman 1995). Al- though there are some chemical techniques aimed to control M. domestica in poultry houses, the negligent human behavior related to the correct application of chemicals has in- tensified the search for potential natural ene- mies of houseflies in order to dimmish chem- ical applications (Cunha & Lomonaco 1996). Therefore, the understanding of the strength of interspecific interactions between M. do- mestica and its predators is of major impor- tance. Dermestes beetles grow in organic matter, such as carrion and dung that accumulate in poultry houses (Cloud & Collison 1986). Der- mestes ater (DeGeer 1774) (Coleoptera, Der- mestidae) feeds and scavenges on animal products. However, sometimes it feeds on oth- er insects, thus acting as a predator (Veer et al. 1996). For example, D. ater causes serious economic damage to sericiculture, because the beetles feed on high numbers of Bombyx mori (Lepidoptera, Bombycidae) (Kumar et al. 1988; Bai & Mahadevappa 1996). According to Lomonaco & Prado (1994), M. domestica (91. 82%) and Chrysomya pu- toria (Diptera: Calliphoridae) (6.47%) were the most abundant fly species sampled in a poultry house located in the city of Uberlandia (MG), Brazil. These authors also observed that D. ater was one of the most frequent nat- ural enemies of larvae and pupae of M. do- mestica in that system. As M. domestica (adults) and D. ater (adults and larvae) are usually seen in N. rufipes webs in poultry houses, and D. ater attacks and feeds on M. domestica, it is of major importance to un- derstand the strength of interspecific interac- tions among these animals in such a system. Here, we investigated prey choice by N. ru- fipes when two different prey species, D. ater and M. domestica, were provided at the same time as primary food sources. We also quan- tified the capture of these prey species by this predator in a poultry house, comparing the re- sults with the prey choice experiment. Corre- lations of prey choice with physical charac- teristics of prey types are also presented. Finally, we discuss whether there is evidence of antagonistic ietraguild interactions in such a system composed of N. rufipes (top preda- tor), D. ater (intermediate predator and prey of N. rufipes), and M. domestica {N. rufipes'' prey). METHODS Field observations. — An experimental poultry house located in the city of Botucatu- SP (Brazil) (22°52'20"S; 48°26'37"W) was chosen to collect insects captured by N. rufi- pes webs. From September 2001 to August 2002, all poultry house parts (walls, door crevices, wood supports, chicken cages, etc.) were examined monthly. When a web site containing N. rufipes was found, all arthropod carcasses caught in the web were removed and put into small glass tubes for identification. Spiders were never removed from their web 188 I! j sites in order not to diminish their abundance, and we did not distinguish males from fe^ males, or even spiderlings from adults, that were inhabiting the webs. The carcasses were then taken to the laboratory where prey were identified. We recorded from each web the species of prey and also the respective month of collection. Voucher specimens (spiders and insects) from this study are deposited in the Invertebrate Collection of the Department of Parasitology, Unesp (Botucatu-SP), Brazil. We compared which prey species were cap- tured throughout the year by plotting the total number of flies and beetles (adults + larvae) captured monthly. In the same plot, we in- cluded the number of web sites observed by month. Rearing of prey species*- — -While visiting the poultry house, we collected larvae of houseflies and adults of D. ater from small samples of chicken feces deposited below the cages and put them into small glass tubes. All insects were then taken to the laboratory where larvae of M. domestica were reared in vials containing wet ground animal ration (25 °C under 12 h light). After pupation, vials were kept in cages of nylon mesh on a wood frame (30 cm X 30 cm X 30 cm) where water and sugar were provided for adults. Adults of D. ater were kept in plastic boxes (15 cm X 45 cm X 30 cm) (25 °C under 12 h light) with large pieces of cotton which allows females to lay their eggs. Wet cotton and fish (sar- dines) were added weekly as water and food sources, respectively. Prey choice.-— Forty-five adult females of N. rufipes were captured in several buildings located on the campus of the University of the State of Sao Paulo (Botucatu, Brazil) from January-March 2003, and kept individually in clear plastic containers [10.5 cm X 11.5 cm (900 ml)] in the laboratory (25 °C under 12 h light). All spiders were of similar size range (15 mm). Before the prey choice experiments were carried out, a nylon mesh (10 cm X 3 cm) was internally fixed in each container in order to allow spiders to build their webs. All spiders were fed with both houseflies and der- mestid beetles for a month (insects were ran- domly offered twice a week) in order to attain similar nutritional status. After twenty-four hours of food deprivation (sufficient time for spiders to build their webs), fifteen containers with spiders received THE JOURNAL OF ARACHNOLOGY j i five larvae (third instar) and five adults of M. !; domestica each. Another fifteen containers re- |l ceived five larvae (fifth instar) of D. ater and ^ five adults of M. domestica each, and the re- I' maining containers received five adults of D. f ater and five adults of M. domestica each. Before adding the different prey types into the spider containers, flies were immobilized by chilling in a freezer for three minutes. Af~ j ter that, flies were removed and put in a Petri |i dish together with the other insects [D. ater \ (larvae or adult) or larvae of M. domestica)] [ previously removed from their laboratory I rearing cages. When flies began to move, ail | ten insects were carefully dropped in the bot- 1 tom of a spider container, without touching the ! spider web. This procedure prevented flies from being captured quickly due to their su- perior flying ability and it insured that flies i could be easily separated prior to the experi- j ments. In the first two minutes (approximate- i ly) inside the spider containers, flies just walked and then started flying. Prey consump- tion evaluation started twenty-four hours fol- lowing the introduction of the insects. The number of prey eaten by spiders was recorded according to the combination of prey types, and an analysis of variance (ANOVA) (Zar 1999) was computed comparing the j mean proportions [arcsin (V^oportion)] of adults of M. domestica consumed, since it was common for all combinations of prey. A Least Significant Difference test (LSD) was com- puted comparing the pairs of transformed mean proportions of adults of M. domestica consumed between the different prey combi- nation treatments (adults + larvae of M. do- mestica, adults of M. domestica + larvae of D. ater, and adults of M. domestica + adults of D. ater). To test the hypothesis that prey choice was random, we compared the mea- sured proportion of prey captured to the prob- ability that prey capture was random (i.e. 50% chance of capturing house flies). We did this by constructing utests on the arcsin (Vproportioe) transformed data, which com- pared the mean transformed value to arcsin (Vas). Size of prey. — After the prey-choice ex- periments, many larvae (fifth instar) and adults of D. ater as well as larvae (third instar) and adults of M domestica were randomly re- moved from their respective rearing cages ROSSI & GODOY— PREY CHOICE BY NESTICODES RUFIPES 189 Months Figure !.■ — ^Number of M. domestica (adults) and D. ater (adults plus larvae) carcasses collected from October 2001 to September 2002 in a poultry house located in Botucatu (SP), Brazil. The number of web sites observed is also included. and, from, there, twenty insects from each prey type were again randomly removed. These in- sects were first killed with ether solution (90%) and then measured (body length mea- sured from anus to head without measuring wings for adult flies) by using a graduated mi- crometric ocular coupled to a stereoscopic mi- croscopy and weighed with a semi-analytical scale. Student’s f-tests were computed com- paring pairs of mean weights and lengths for each combination of prey types. Thus, we test- ed whether the lighter and shorter prey were also the more preferable ones. RESULTS Musca domestica carcasses were much more abundant than D, ater (adults + larvae) on N. rufipes webs for most of the 12 months of collection (Fig. 1). The spiders in the poul- try house ate more 5.5 times as many flies {n = 44) than dermestid beetles (« = 8) over the course of the year-long study (Fig. 1). Spiders captured a total of sixteen species of prey. In- sects from orders Coleoptera (4836%) and Diptera (34.02%) represented 8238% of all prey captured, and for all months sampled M. domestica was predominant as prey, since it represented 24.59% of the insects captured, followed by the coleopterans Alphitobius dia- perinus (Tenebrionidae) (20.90%), Aphodius (Scarabaeidae) (10 25%), Gnathocerus (Te- nebrionidae) (6.15%), and D. ater (3.28%). All dipterans except M domestica represented only 9.4% of prey. Even though several prey were captured, in figure 1 we present data only related with the arthropod species studied here. In the prey choice experiments, the number of adult flies consumed by spiders was signif- icantly different when different combinations of prey types were offered {df = 2; MS = 0.808; F = 4.185; F = 0.023), and the com- bination of adults of M. domestica plus adults of D. ater showed the highest average pro- portion of adult flies consumed (Table 1). The Student’s t-tests showed that when spi- ders were placed in cages with adults of M. domestica plus adults of D. ater, or with adults of M domestica plus larvae of M. do- mestica, spiders were selective and took more adult flies than the other prey (Table 1). Al- though the combination of adults of M. do- 190 THE JOURNAL OF ARACHNOLOGY » Table 1 . — Mean proportion of spiders that fed on adults of M. domestica given different combinations of potential prey. Student’s r-tests were used to test for significance of difference using the transformed mean proportions [arcsin (Vproportion)] of adults of M. domestica consumed and the probability of 50% |' [arcsin (Vo.5)] of consumption. *Significant at F < 0.0 L In addition, proportions followed by different [ letters differed statistically from each other [Least Significant Difference {LSD) test] at F < 0.05. n = Number of observations for each group, « = 14 and « = 13 means that one and two spiders did not feed on any prey during experimentation, respectively. Combination of prey types Mean proportion (± SD) n t F Adults X Larvae of M. domestica 0.73 ± 0.44 a 13 3.45 0M02* Adults of M. domestica X Larvae of D. ater 0.71 ± 0.80 a 15 1.86 0.074 Adults of M. domestica X Adults of D. ater 0.96 ± 0.26 b 14 13.0 0.000* smaller mass and shorter lengths, probably be- | cause it would facilitate their being killed and i handled by spiders. Prey activity would also | explain why spiders captured disproportion- ately more adults of M. domestica than the j other prey types (Table 1). According to Prov- ! eecher & Coderre (1987), prey activity is be- i lieved to influence functional responses and I switching of spiders for some prey. Although I all prey species were highly active in the ex- | perimental containers, only adults of M. do- j mestica could do a three-dimensional explo- i ration in the container, since it flew over all mestica plus larvae of D. ater presented a nonsignificant result for adult flies consumed (F = 0.074), a strong tendency of spiders to consume more flies was evidenced (Table 1). Adults of M. domestica weighed less when compared to the other prey offered to spiders, and it also had smaller body size since Stu- dent’s Atests were significant for all compar- isons in all combinations of prey (Figs. 2, 3). DISCUSSION The preference of N. rufipes for adults of M. domestica might be associated with their Figure 2. — Comparisons of mean weights (grams) of prey according to different combinations of prey types. A Student’s Atest was computed for each combination and all analyses were statistically significant (All analyses had n = 20 and 38 degrees of freedom). Combination 1: adults + larvae of M. domestica (Avalue = —13.27; F < 0.01); Combination 2: adults of M. domestica + larvae of D. ater (Avalue = — 20.60; F < 0.01); Combination 3: adults of M. domestica + adults of D. ater (Avalue = —12.69; F < 0.01). ROSSI & GODOY-^PREY CHOICE BY NESTICODES RUFIPES 191 Figure 3. “—“Comparisons of mean lengths (mm) of prey according to different combinations of prey types. A Student’s t-test was computed for each combination and all analyses were statistically significant (All analyses had n = 20 and 38 degrees of freedom). Combination 1: adults + larvae of M. domestica (r-value = ""2.65; P < 0.05); Combination 2: adults of M. domestica + larvae of D. ater (lvalue = “20.71; P < 0.01); Combination 3: adults of M. domestica + adults of D. ater (lvalue = “7.15; P < 0.01). areas of the container, probably increasing its frequencies of encountering the predator. The other prey types only walked intensively in the bottom of the container with the exception of adults of D. ater, which occasionally flew. The higher rate of consumption of adults of M. domestica when this prey was offered con- comitantly with adults of D. ater (Table 1) is possibly associated with the rigidity of beetle exoskeletoes, which may increase their rate of escape from spider attacks. We observed that all spiders actively cap- tured their prey rather than passively waiting for prey to fall randomly in their webs (sit- and- wait strategy). This behavior was possible because the available time given to spiders to build their webs (24 hours) was insufficient to enable them to weave large and dense webs. Large webs would allow spiders to catch prey only by a prey-web contact. Hence, the way that we set up the experiments ensured that webs were just used by spiders to increase their area of attack, forcing them to actively choose a prey type. It is important to state that all spiders wove webs in all parts of the con- tainers, including the bottom, enabling them to capture all prey available. Thus, we con- clude that preference of spiders for housefly adults was determined by prey behavior and physical characteristics of prey (length and weight) in addition to active spider prey choice. Finke & Deneo (2002) studied the com- bined impact of two salt-marsh-inhabiting in- vertebrate predators, the mirid Tytthus vagus (Heteroptera, Miridae) and the wolf spider Pardosa littoralis Banks 1896 (Araeeae, Ly- cosidae), on suppression of their shared prey, the planthopper Prokelisia dolus (Hemiptera, Delphacidae), in simple and complex habitats. They observed that in simple habitats, the predators interacted antagonistically, due to intraguild predation of rnirids by spiders, and predation pressure on the planthopper popu- lation was relaxed. However, for structurally complex habitats this antagonistic interaction was dampened by providing a refuge for mir- ids from spider predation. Our experiments demonstrated that the predation impact of N. rufipes on D. ater is low when compared to that on M. domestica (Fig. 1), and it provides some evidence that an antagonistic interaction between these predators (and scavenger) may not occur, suggesting that they are in fact act- 192 ing either synergistically or additively on M. domestica prey. Considering that more than a hundred of pathogens are associated with M. domestica, such as those causing typhoid fever, cholera, tuberculosis, parasitic helminthiasis and pro- tozoosis (Harwood & James 1979; Smith 1986; Levine & Levine 1991; Chavasse et al. 1999; Fischer 1999), synergistic and additive interactions between D. ater and N. rufipes have important practical implications since it may increase the likelihood of a natural sup- pression of housefly populations established in poultry houses. However, functional re- sponse studies of D. ater and N. rufipes on larvae and adults of M. domestica, respective- ly, are encouraged in order to understand the actual contribution of these predators in di- minishing natural or experimental housefly populations. ACKNOWLEDGMENTS Four anonymous reviewers provided valu- able insights and offered several helpful sug- gestions for improving early versions of this manuscript. M.N.R. is particularly grateful to Fapesp (Fundagao de Amparo a Pesquisa do Estado de Sao Paulo - Contract N° 01/06368- 2) for financial support, W.A.C.G. has been supported by a research fellowship from CNPq (Conselho Nacional de Desenvolvi- mento Cientifico e Tecnologico), We also thank Professor Luzia Aparecida Trinca for statistical advice. LITERATURE CITED Alderweireldt, M. 1994. Prey selection and prey capture strategies of linyphiid spiders in high- input agricultural fields. Bulletin of the British Arachnological Society 9:300-308. Axtell, R.C. & J.J. 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Journal of Stored Products Re- search 32:69-89. Wise, D.H. 1993. Spiders in Ecological Webs. Cam- bridge University Press, Cambridge. Wise, D.H. & B. Chen. 1999. Impact of intraguild predators on survival of a forest-floor wolf spi- der. Oecologia 121:129-137. Zar, J.H. 1999. Biostatistical Analysis. Prentice Hall, Upper Saddle River, NJ. Manuscript received 1 Eebruary 2004, revised 1 March 2005. 2006. The Journal of Arachnology 34:194-205 A REVIEW OF PHOLCID SPIDERS FROM TIBET, CHINA (ARANEAE, PHOLCID AE) Feng Zhang, Ming-Sheng Zhu and Da-Xiang Song: College of Life Sciences, Hebei University, Baoding, Hebei 071002, China. E-mail: zhangfeng@mail.hbu.edu.cn. ABSTRACT. The pholcid spiders from Tibet, China, are reviewed. Seven species belonging to three j genera are recorded. A new genus, Tibetia, is established, and four new species, Pholcus medog, P. zham, \ Belisana gyirong and B. mainling are described. And two new combinations are formed: Tibetia everesti | (Hu & Li 1987) is transferred from Pholcus, and Belisana yadongensis (Hu 1985) is transferred from Spermophora. Keywords: Taxonomy, new species, new combination, Asia I The spider family Pholcidae currently con- tains 75 genera and 868 species (Platnick 2004) throughout the world. Members of the family vary in habitus, size and life style. Also, the pholcids are among the most com- mon spiders occurring in houses. These spi- ders have elongate or globose abdomens and frequently very long and thin legs (about 4- 20 times as their body length) with false seg- ments in tarsi, and are thus called daddy-long- legs, although in a few pholcid species, the legs are quite short (only about 1 mm). The overall coloration of pholcids is quite variable, but the legs are often characteristically annu- lated. The eye region is more or less elevated, bearing eight or six eyes. If the smallest AMEs are present, the others are in two triads. The presence of the cheliceral stridulatory or- gans is variable. The male chelicerae are fre- quently equipped with pairs of special apoph- yses which are often species-specific in morphology and the pedipalps are conspicu- ously large and strong (Huber 1995, 1999) and their complex morphology has been well demonstrated by Uhl et al. (1995). Externally the female genitalia are usually relatively sim- ple, but the internal morphology is very com- plicated (Huber 1998). Pholcids spin messy, loose and irregular webs, and the males and females often hang inverted in the same webs. The females carry the spherical egg cluster under their chelicer- ae. When disturbed or under threat of attack, most pholcid species violently vibrate within their webs to scare off antagonists. Also sol- itary species on firm ground can shake their bodies in such a rhythm that they virtually dis- appear from the human eye (Saaristo 2001). i Many pholcid species are pantropical and syn- ! anthropic. They live in dark recesses, such as j within houses and other buildings, in caves, ; under rocks and loose bark, as well as in leaf I litter. ! Previously, only three pholcid species be- ! longing to two genera have been recorded from the Tibetan region of China (Hu 1985; Hu & Li 1987; Hu 2001): Spermophora ya~ | dongensis Hu 1985, Pholcus everesti Hu & Li ' 1987 and Pholcus affinis Schenkel 1953. j After examination of pholcid specimens i collected from Tibet in 2002 and 2003, we | here deal with seven species of this family : belonging to three genera, including a new ge- ; nus, Tibetia, and four new species: Pholcus medog, P. zham, Belisana gyirong and B. mainling. Also, two new combinations are proposed: Belisana yadongensis (Hu 1985) (transferred from Spermophora), Tibetia ev- \ eresti (Hu & Li 1987) (transferred from Phol- cus). Additionally, the common species, Phol- cus manueli Gertsch 1937 (senior synonym of Pholcus affinis Schenkel 1953, by Senglet 2001) is listed. METHODS This paper is mostly based on material col- lected from Tibet by staff members from He- bei University, with the exception of Pholcus manueli Gertsch 1937. Terminology is stan- dard for Araneae. Carapace length was mea- i 194 ZHANG ET AL.— PHOLCIDS FROM TIBET 195 sured from the anterior face of the ocular area to the rear margin of the carapace medially, excluding the clypeus. Total length is the sum of carapace and abdom.en length, regardless of the petiole. The measurements of legs are pre- sented as follows; total length (femur + pa- tella plus tibia + metatarsus T tarsus). The left male pedipalp is used for illustrations. Epigyea were cleared in a warm solution of potassium hydroxide (KOH), transferred to water and temporarily mounted for drawing. All measurements are given in millimeters. Type specimens are deposited in the College of Life Sciences, Hebei University (HU). The following abbreviations are used: ALE “ anterior lateral eyes; AME = anterior me- dian eyes; PLE = posterior lateral eyes; PME = posterior median eyes; MO A ~ median oc- ular area; AME -ALE — distance between AME and ALE; ALE-PLE — distance be- tween ALE and PLE; AME-AME = distance between AMEs; PME-PLE — distance be- tween PME and PLE; PME-PME — distance between PMEs. TAXONOMY Family Pholcidae C.L. Koch 1851 Genus Pholcus Walckenaer 1805 Pholcus Walckenaer 1805: 80; Simon 1893: 470- 471; Huber 2000: 77; Huber 2001: 108-111; Hu 2001: 8L Type species. — Aranea phalangioides Fuesslin 1775, by subsequent designation. Diagnosis. — Medium to large-sized phol- cids with cylindrical opisthosoma, mostly with AMEs present. The most useful charac- ters that distinguish Pholcus from other gen- era are the projections of the bulb, tradition- ally called the uncus, the appendix and the embolus. Other characters are the conserva- tive male chelicerae (a pair of dark frontal apophyses and a pair of light lateral apophy- ses), the shape of the procursus (usually with ventral boss), and the knob or worm-shaped apophysis on the often roughly triangular or oval epigyeum (Huber 2001). Remarks. — The genus Pholcus, with more than 110 species mainly from the pan-Pacific region and Africa, is the largest pholcid genus. Most nominal species seem to be correctly placed, although they are poorly revised at the specific level. The real taxonomic problem is its relationship to other genera of the Pholcus group(see Huber 2001). Pholcus manueli Gertsch 1937 Pholcus manueli Gertsch 1937: 1, figs. 6-7; Senglet 2001: 62, figs. 60-66. Pholcus affinis Schenkel 1953: 23, figs. 12a-b; Song et al. 1999: 52, figs. IIH, 22D-G; Hu 2001: 81, figs. 7.1-4 (first synonymized with P. man- ueli by Senglet in 2001). Type matermh— Pholcus manueli: the type specimens from New Jersey are deposited in the American Museum of Natural History, New York (not examined). Pholcus affinis: 1 male paratype from Tcheuly of China is deposited in the Natural History Museum of Basel, Switzerland, ex- amined by Senglet (2001); other type material unknown. Material examined.- — None from Tibet. Description. — See Gertsch (1937) and Hu (2001). Distribution. — China: Tibet (Gyiroeg), Hebei, Zhejiang, Jiangsu, Sichuan, Shannxi, Shanxi, Inner Mongolia, Liaoning, Jilin; Rus- sia, Japan, U.S.A. Remarks. — ^Judging from the figures drawn by Hu (2001), we are confident that this spe- cies is correctly identified and it is confirmed from Tibet. Pholcus medog new species Figs. 1-8 Material examined. — CHINA: Tibet: Ho- lotype male, Medog County (29°12'N, 95°18'E), 17 August 2003, Feng Zhang (HU). Paratypes: 5 females, 2 males, same data as holotype; 19,1^, Medog County, 10 Au- gust 2003, Feng Zhang (HU); 1 9,3 juve- niles, Baibung Town (29°12'N, 95°06'E), Me- dog County, 13 August 2003, Feng Zhang (HU). Etymology. — The species name is a noun in apposition derived from the type locality. Diagnosis. — This species resembles P. po- dophthalmus Simon 1893 (Song et aL 1999: 58, figs. 240-V), but can be readily distin- guished from the latter by the shape of uncus, the bifurcate appendix of the male pedipalp and the color pattern of the carapace. Description, — Male (holotype): Total body length 6.25: cephalothorax 1.17 long, 1.71 wide; abdomen 5.08 long, 1.40 wide. Prosoma shape as in Fig 1. Carapace short, broad and 196 THE JOURNAL OF ARACHNOLOGY Figures 1-8. — Pholcus medog, new species: 1. male prosoma, dorsal view; 2. male opisthosoma, dorsal view; 3. male sternum, ventral view; 4. male chelicerae, frontal view; 5. left pedipalp, prolateral view. 6. same, retrolateral view; 7. epigynum, ventral view; 8. same, dorsal view. Scale lines: 1 mm (1-3), 0.5 mm (4-8). almost circular, ochre, with brown mark broadly connecting to ocular area. Cephalic region raised, with two brown slender central marks, ocular area dark yellow. Clypeus 0.43 high, ochre, without marks. Except AMEs, other six eyes in two triads, each triad on the top of a relatively longer eye stalk. Distance AME-AME 0.05. Diameter AME 0.09, ALE 0.26, PME 0.18, PLE 0.23. Chelicerae shaped as in Fig. 4, with pair of black apophyses dis- tally and pair of unsclerotized rounded apoph- yses proximolaterally. Labium light yellow. Endites gray. Sternum (Fig. 3) dark gray, with irregular yellow patches centrally. Legs ex- ceedingly long and slender, femora, patellae and tibiae ochre, with dark rings, metatarsi and tarsi brown. Measurements of legs: I 52.51 (13.05 + 13.50 + 22.95 + 3.01); II 34.23 (9.00 + 9.21 + 14.4 + 1.62); III 23.47 (6.49 + 6.30 + 9.23 + 1.45); IV 32.11 (8.89 + 8,68 T 12.83 + 1.71). Leg formula: 1243. Abdomen cylindrical, pale ochre, dorsum with large brown pattern as in Fig. 2, venter with central long brown stripe. Spinnerets yellow- ish brown. Uncus of male pedipalp large and slightly triangular, heavily sclerotized and pro- vided with many teeth on the edge; appendix spilt into two parts; embolus lying between the uncus and appendix, soft and transparent (Fig. 5); procursus with ventral boss (Fig. 6). Female: In general very similar to male. Total body length 4.86-6.23. One specimen measured: total length 5.95: cephalothorax 1.45 long, 1.53 wide; abdomen 4.50 long, 1.31 wide. Clypeus 0.55 high. Both eye rows recurved. Except AMEs, other six eyes in two triads, each triad on a slightly elevated tuber- cle. Distance AME-AME 0.05, AME- ALE 0.13, PME-PME 0.30, PME-PLE 0.04. Di- ameter AME 0.08, ALE 0.18, PME 0.15, PLE 0.17; MO A 0.26 long, front width 0.19, back width 0.56. Measurements of legs: I 40.81 (10.07 + 10.21 + 18.00 + 2.53); II 24.53 (6.75 + 6.03 + 10.13 + 1.62); III 17.58 (4.97 + 4.82 T 6.66 T 1.13); IV 24.08 (6.93 + 6.53 T 9,00 + 1.62). Leg formula: 1243. Epigyn- um roughly triangular, with a knob-shaped apophysis on the top of it (Fig. 7). ZHANG ET AL.-— PHOLCIDS FROM TIBET 197 Figures 9-16. — Pholcus zham, new species: 9. male prosoma, dorsal view; 10. male opisthosoma, dorsal view; 11. male sternum, ventral view; 12. male chelicerae, frontal view; 13. left pedipalp, prolateral view; 14. same, retrolateral view; 15. epigynum, ventral view; 16. same, dorsal view. Scale lines: 1 mm (9, 10), 0.5 mm (11-16). Habitat*— Untidy webs are made under cliff or rock crevices. Generally, a male and a female hang upside down in the same web. Distribution.— Known from Medog Coun- ty, Tibet. Pholcus zham new species Figs. 9-16 Material examined.— CHINA: Tibet: Ho- lotype male, Zham Town (27°54'N, 85°54'E), Nyalam County, 30 August 2002, Feng Zhang and Zhi-Sheng Zhang (HU). Paratypes: 6 fe- males, same data as holotype (HU). Etymology.— The species name is a noun in apposition derived from the type locality. Diagnosis.— This new species resembles P. medog (Figs. 1-8), but can be readily distin- guished from the latter by: the shape of the uncus, the noe-bifurcate appendix of male pedipalp and the chelicerae with two pairs of unsclerotized apophyses centrally. Description. — Male (holotype): Total body length 6.33: cephalothorax 1.56 long, 1.73 wide; abdomen 4.77 long, 1.26 wide. Prosoma shape as in Fig. 9. Carapace short, broad and almost circular, ochre, with brown mark broadly connecting to ocular area. Cephalic region raised, with a brown longitudinal mark centrally, ocular area yellow. Clypeus 0.80 high, ochre, without marks. Except AMEs, other six eyes in two triads, each triad on the top of a relatively longer eye stalk. Distance AME-AME 0.07. Diameter AME 0.11, ALE 0.22, PME 0.16, PLE 0.17. Chelicerae shaped as in Fig. 12, with pair of black apophyses distally, and two pairs of unsclerotized round- ed apophyses proximolaterally and proximo- centrally respectively. Labium light yellow. Endites gray. Sternum (Fig. 11) dark gray, with roughly 5 pairs of irregular yellow patch- es on it. Legs exceedingly long and slender, femora, patellae and tibiae ochre, with dark rings, metatarsi and tarsi brown. Measure- ments of legs: I 56.61 (13.77 + 13.95 + 25.65 198 THE JOURNAL OF ARACHNOLOGY -f 3„24); II 36.90 (9.76 + 10.04 + 14.85 + 2.25) ; III 24.57 (7.02 + 6.75 + 9.45 + 1.35); IV 32.91 (9.59 A 8.83 + 12.83 + 1.66). Leg formula: 1243. Abdomen cylindrical, pale ochre, dorsum with many brown spots as in Fig. 10, venter with central long brown stripe. Spinnerets yellowish brown. Uncus of male pedipalp large and slightly rectangular, heavi- ly sclerotized and provided with many scales on the edge; sclerotized appendix hook- shaped and rod-like; embolus lying between the uncus and appendix, soft and transparent (Fig. 13); and procursus with ventral boss (Fig. 14). Female: In general very similar to male. Total body length 7.36-8.82. One specimen measured: total length 8.82: cephalothorax 1.70 long, 1.72 wide; abdomen 4.32 long, 1.48 wide. Clypeus 0.49 high. Both eye rows recurved. Except AMEs, other six eyes in two triads, each triad on a slightly elevated tuber- cle. Distance AME-AME 0.06, AME-ALE 0.12, PME- PME 0.33, PME-PLE 0.08. Di- ameter AME 0.10, ALE 0.22, PME 0.14, PLE 0.21; MOA 0.31 long, front width 0.27, back width 0.58. Measurements of legs: I (12.96 + 13.95 + 25.61 + 3.19); II (9.45 + 9.54 + 14.37 + 2.21); III (6.75 + 6.44 + 9.00 + 1.26) ; IV (9.72 + 8.82 + 12.96 + 1.74). Leg formula: 1243. Epigynum roughly triangular, with a knob-shaped apophysis on the top of it (Fig. 15). Habitat. — Untidy webs are made under rocks. Distribution. — Known only from type lo- cality in Nyalam County, Tibet. Tibetia new genus Type species. — Pholcus everesti Hu & Li 1987. Etymology, — The generic name refers to Tibet, the Xizang Autonomous Region. The gender is feminine. Diagnosis. — The genus Tibetia is similar to several other genera (Wugigarra Huber 2001, Trichocyclus Simon 1908, Physocyclus Simon 1893 and Artema Walckenaer 1837) through the possession of the peculiar set of structures on the procursus (dorsal apophysis and ventral pocket). It can be distinguished from Wugi- garra by the absence of a characteristic worm- shaped process on the male bulb (Figs, 21- 22) and the absence of stridulatory files in females, and the epigynum with a median de- pression (Figs. 23, 51); from Trichocyclus by the absence of a weak zone dorsally on male cymbium (Fig. 56), and the mediae depression of the female epigynum; from Physocyclus by the bulbal apophysis and the shape of the pro- cursus (Fig. 56), the absence of embolus on the bulb (only the sperm duct opening be seen) (Fig. 52) and the shape of the epigynum (Figs. 23, 51); and from Artema by the epi- gynal depression centrally (Fig. 23, 51), the absence of embolus and the shape of bulbal apophysis. Additionally, the new genus also differs from the Holocneminus Berland 1942 (Ber- land 1942, figs. 5a-f), which is apparently widely distributed in eastern Asia, by the ab- sence of stridulatory files in the female, and the normal tarsus of female pedipalp (not strongly dilated) (Fig. 52). Description. — See description of single species below. Remarks. — Judging from the known dis- tribution of Physocyclus, it appears that Phy- socyclus is a New World genus, with the ex- ception of the pantropical Physocyclus globosus, while the new genus is found only in Tibet. Thus the two genera appear to be allopatric in distribution. Tibetia everesti (Hu & Li 1987) new combination Figs. 17-24, 51-56 Pholcus everesti Hu & Li 1987: 260, fig. 8; Song, Zhu & Chen 1999: 57; Hu 2001: 82, fig. 8. 1-7. Type material.- — Hu & Li (1987) described both sexes from Nyingchi and Namling Coun- ties, Tibet. The type specimens are deposited in Shandong University, China, not examined. Material examined. — CHINA: Tibet: 5 9 , 2 6, Zetang Town (29°12^N, 9r42'E), Ne- dong County, 25 August 2002, Mieg-Sheng Zhu and Feng Zhang (HU); 1 9,1 6, Rigaze City (29°12'N, 88°48'E), 6 September 2002, Jun-Xia Zhang (HU); 1 9, Nyingchi County (29°30'N, 94°!8'E), 21 August 2003, Feng Zhang (HU); 19,16, Bayi Town (29°36^N, 94°12'E), Nyingchi County, 2 August 2003, Ming-Sheng Zhu and Zhi-Sheng Zhang (HU); 1 9, Lhasa City (29°36'N, OUOb'E), 30 July 2002, Ming-Sheng Zhu and Jun-Xia Zhang (HU). Diagnosis. — See generic diagnosis above. Description. — Male: Total body length ZHANG ET AL.— PHOLCIDS FROM TIBET 199 Figures 17-24. — Tibetia everesti, new species: 17. male, dorsal view; 18. male left chelicerae, retrela- teral view; 19. male sternum, ventral view; 20. male chelicerae, frontal viev/; 21. left pedipalp, prolateral viev/; 22. same, retrolateral view; 23. epigynum, ventral view; 24. same, dorsal view. Scale lines: 0.5 mm (17, 19), 0.3 mm (21-24), 0.2 mm (18, 20). 1.63-2.24. One specimen measured: total length 2.24: cephalothorax 0.81 long, 1.15 wide; abdomen 1.43 long, 1.00 wide. Prosoma shape as in Fig 17. Carapace oval, wider than long, with distinct thoracic groove and brown mark centrally. Cephalic region slightly raised, ocular area light yellowish. Clypeus 0.30 high, unmodified, yellowish, with light brown marks. Except AMEs, other six eyes in two traids, on a moderately elevated ocular area. Distance AME-AME 0.04, AME=-ALE 0.04, PME-^PME 0.13, PME^PLE 0.03. Di^ ameter AME 0.06, ALE 0.10, PME 0.13, PLE 0.10. MOA 0.22 long, front width 0.16, back width 0.29. Chelicerae (Figs. 20-21) with stridulatory ridges and pair of large black apophyses proximolaterally. Labium gray. En- dites pale. Sternum (Fig. 19) yellowish, with- out patches on it. Legs long, yellow, with dark rings subdistally on femur, patella plus tibia proximally, and tibia subdistally. Measure- ments of legs: I 11.15 (3.33 + 3.42 + 3.50 + 0.90); II 9.50 (2.75 + 2.97 + 3.06 + 0.72); III 8.70 (2.39 -f 3.06 + 2.57 + 0.68); IV 10.31 (3.42 + 3.06 + 3.15 + 0.68). Leg for- mula: 1243. Abdomen globular, gray, with black spots dorsally. Spinnerets whitish yel- low. Procursus (Fig. 22) relatively simple and with dorsal apophysis, ventral pocket indis- tinct; bulb (Fig. 21) consisting of the proximal globular part and the distal sclerotized apoph- ysis, without embolus, but the sperm duct opening can be seen (Fig, 52). Female: In general very similar to male. Total body length 1.88-2.34. One specimen measured: total length 2.34: cephalothorax 0.86 long, 1.04 wide; abdomen 1.50 long, 1.04 wide. Clypeus 0.33 high. Both eye rows recurved. Distance AME- AME 0.03, AME- ALE 0.03, PME-PME 0.14, PME-PLE 0.02. Diameter AME 0.05, ALE 0.10, PME 0.10, PLE 0.10. MOA 0.20 long, front width 0.13, back width 0.29. Measurements of legs: I lost; II 8.02 (2.52 + 2.45 + 2.46 + 0.59); III 6.85 (2.03 + 2.07 + 2.16 + 0.59); IV 8.97 (2,88 + 2.70 + 2.70 + 0.69). Epigynum roughly rect- angular, with a depression centrally (Fig. 23). 200 THE JOURNAL OF ARACHNOLOGY I Figures 25-31. — Belisana gyirong, new species: 25. male prosoma, dorsal view. 26. male opisthosoma, dorsal view. 27. male sternum, ventral view. 28. male chelicerae, frontal view. 29. male left chelicera, retrolateral view. 30. left pedipalp, prolateral view. 31, same, retrolateral view. Scale lines: 0.5 mm (25- 27), 0.3 mm (30-31), 0.2 mm (28-29). Distribution. — Known only from several localities in Tibet. Belisana Thorell 1898 Belisana Thorell 1898: 278; Simon 1903: 988; Si- mon 1909: 81; Deeleman-Reiehold 1986: 46-47; Huber 2001: 124-126. Type species. — Belisana tauricornis Tho- rell 1898, by original designation Diagnosis. — Small-sized, pholcids with roughly globular or higher-than-long opistho- soma. Six eyes in two triads, AMEs absent, and eyes not elevated. Distance between PMEs less than two times diameter of PME. The genitalic structure is somewhat similar to that of Spermophora (Deeleman-Reinhold 1986), but can be distinguished from Sper- mophora by the distance between the PMEs which is less than two times the diameter of PME, but is more than three times the diam- eter of PME in Spermophora. Remarks. — The distinction between Beli- \ Sana and Spermophorides requires further clarification (see Huber 2001). Belisana gyirong new species Figs. 25-31 Material examined. — CHINA: Tibet: Ho- lotype male, Gyirong Town (28°24'N, 85°12'E), Gyirong County (28°54'N, 85°12'E), 2 September 2002, Ming-Sheng Zhu and Jun-Xia Zhang (HU). Paratype: 1 male, same data as holotype (HU). Etymology. — The species name is a noun in apposition derived from the type locality. Diagnosis. — This new species resembles B. yadongensis (Hu 1985), but can be readily distinguished from the latter by the long apophysis of the bulb and the subdistal apoph- yses of chelicerae bending internally. Description. — Male (holotype): Total length of body 2.23: cephalothorax 0.86 long, ZHANG ET AL.— PHOLCIDS FROM TIBET 201 Figures 32--40. — Belisana mainling, new species: 32. male prosoma, dorsal view; 33. male opisthosoma, dorsal view; 34. male sternum, ventral view; 35. male left chelicera, retrolateral view; 36. male chelicerae, frontal view; 37. left pedipalp, prolateral view; 38, same, retrolateral view; 39. epigynum, ventral view; 40. same, dorsal view. Scale lines: 0.5 mm (32-33), 0.3 mm (34), 0.2 mm (35-40). 0.91 wide; abdomen L37 long, 0.87 wide. Prosoma shape as in Fig. 25, Carapace oval, slightly longer than wide, without thoracic groove, with brown mark on each side of car- apace. Cephalic region not raised. Ocular area light yellowish, with an ochre mark centrally. Clypeus 0.22 high, unmodified, without marks. Six eyes in two triads. Distance PME- Figures 41-50. — Belisana yadongensis: 41. male prosoma, dorsal view; 42, male opisthosoma, dorsal view; 43. male chelicerae, frontal view; 44, male sternum, ventral view; 45, 46. left pedipalp, prolateral view; 47, 48. same, retrolateral view; 49. epigynum, ventral view; 50. same, dorsal view. Scale lines: 0.5 mm (41-42, 44), 0.2 mm (43, 45-50). Figures 51-56. — Tibetia everesti, new species; 51. epigynum, ventral view, showing copulatory open- ing; 52. tip of female pedipalp; 53. male right palpal organ, showing sperm duct opening; 54. male right chelicera, showing the end of black apophysis proximolaterally; 55. male right chelicera, showing strid- ulatory ridges; 56. male right pedipalp, showing shape of the procursus. THE JOURNAL OF ARACHNOLOGY 202 ZHANG ET AL.— PHOLCIDS FROM TIBET PME 0.09. Diameter ALE 0.09, PME 0.09, PLE 0.09. Chelicerae (Figs. 28-29) with pair of simple, black and long apophyses subdis-= tally and pair of rounded light apophyses proximolaterally. Labium and eedites whitish. Sternum yellowish, without patches on it. Measurements of legs: I 19.06 (5.22 + 5.72 + 5.89 + 2.23); II 13.66 (3.38 + 4.05 + 4.95 + 1.28); III 9.18 (2.70 + 2.61 + 3.15 + 0.72); IV11.21 (3.33 + 3.15 + 3.83 + 0.90). Leg formula: 1243. Abdomen (Fig. 26) almost globular, whitish with black spots dorsally. Bulb consisting of proximal globular part and distal sclerotized apophysis (Fig. 30). Female: Unknown. Distribution. — Known only from the type locality in Tibet. Belisama mainling new species Figs. 32-40 Material examined.^CHINA: Tibet: Ho- lotype male, Mainling County (29°12'N, 94°06'E), 19 August 2002, Ming-Sheng Zhu and Jun-Xia Zhang (HU). Paratypes: 2 fe- males, same data as holotype (HU). Etymology.— The species name is a noun in apposition derived from the type locality. Diagnosis.— This new species resembles B. yadongensis (Hu 1985), but can be readily distinguished from the latter by the tip of the bulbal apophysis with two pointed apiculi, the internal structure of epigynum., and the apoph- yses subdistally of chelicerae bending inter- nally. Description.— Mflie (holotype): Total length of body 2.18: cephalothorax 0.73 long, 0.78 wide; abdomen 1.45 long, 1.05 wide. Prosoma shape as in Fig. 32. Carapace oval, wider than long, without thoracic groove, with brown mark on each side of carapace. Ce- phalic region not raised. Ocular area light yel- lowish, with a ochre bar centrally. Clypeus 0.23 high, unmodified, without marks. Six eyes in two triads. Distance PME-PME 0.10. Diameter ALE 0.09, PME 0.08, PLE 0.08. Chelicerae (Figs. 35-36) with pair of simple black long apophyses subdistally and pair of rounded light apophyses proximolaterally. La- bium and endites whitish. Sternum (Fig. 34) yellowish, without patches on it. Measure- ments of legs: I 10.13 (2.56 T 2.83 + 3.51 + 1.23); II 7.45 (2.03 + 2.18 + 2.49 + 0.75); III 5.89 (1.57 + 1.93 + 1.71 + 0.68); IV 6.63 (1.87 + 1.92 + 2.16 + 0.68). Leg formula: 203 1243. Abdomen (Fig. 33) almost globular, white with black spots dorsally. Bulb consist- ing of the proximal globular part and the distal sclerotized apophysis (Fig. 37). Female: In general very similar to male. Total body length 1.58-1.66. One specimen measured: total length 1.66: cephalothorax 0.67 long, 0.70 wide; abdomen 0.99 long, 0.58 wide. Clypeus 0.19 high. Distance PME- PME 0.08. Diameter ALE 0.08, PME 0.08, PLE 0.08. Measurements of legs: I 8.07 (2.23 + 2.38 + 2.45 + 1.01); II 5.90 (1.79 + 1.76 + 1.69 + 0.66); III 4.04 (1.16 + 1.14 + 1.19 + 0.55); IV 5.35 (1.73 + 1.53 + 1.48 + 0.61). Leg formula: 1243. Epigynum roughly oval (Fig. 39). Distribution. — Known only from the type locality in Tibet. Belisana yadongensis (Hu 1985) new combination Figs. 41-50 Spermophora yadongensis Hu 1985: 148, figs. 1- 10; Song et al. 1999: 65; Hu 2001: 85, figs. 10.1- 10. Type material.- — Hu (1985) described both sexes from Yadong County, Tibet. The type specimens are deposited in Shandong Univer- sity, China, not examined. Material examined. — -CHINA: Tibet: 19 9, 11 c^, Yadong County (27°24'N, 88°54'E), 3 September 2002, under stone heap, Feng Zhang and ZM-Sheng Zhang (HU); 2 9,16, Xiayadong Town, Yadong County, 4 Septem- ber 2002, Feng Zhang and Zhi-Sheng Zhang (HU). Description. — Male: Total body length 1.39-1.71. One specimen measured: total body length 1.71: cephalothorax 0.70 long, 0.67 wide; abdomen 1.01 long, 0.78 wide. Prosoma shape as in Fig. 41. Carapace oval, slightly longer than wide, without thoracic groove, with brown mark on each side of car- apace. Cephalic region not raised. Ocular area light yellowish, with an ochre bar centrally. Clypeus 0.18 high, unmodified, without marks. Six eyes in two triads. Distance PME- PME 0.12. Diameter ALE 0.08, PME 0.06, PLE 0.07. Chelicerae shaped as in Fig. 43, with pair of simple large black apophyses sub- distally and pair of rounded light apophyses proximolaterally. Labium and eedites whitish. Sternum yellowish, without patches on it. Legs light ochre, without dark ring and spine. 204 THE JOURNAL OF ARACHNOLOGY Measurements of legs: I 9.91 (2.48 + 2.80 + 3.42 + 1.21); II 7.16 (1.94 + 2.16 + 2.34 + 0.72); III 5.66 (1.52 + 1.89 + 1.67 + 0.58); IV 6.36 (1.80 + 1.87 + 2.07 + 0.62). Leg formula: 1243. Abdomen almost globular, whitish with black spots dorsally. Bulb con- sisting of the proximal globular part and the distal sclerotized apophysis. Female: In general very similar to male. Total body length 1.63“1.88. One specimen measured: total body length 1.84: cephalotho- rax 0.67 long, 0.67 wide; abdomen 1,17 long, 0.92 wide. Clypeus 0.17 high. Distance PME- PME 0.13. Diameter ALE 0.08, PME 0.08, PLE 0.06; Measurements of legs: I 7.84 (2.12 + 2.25 + 2.48 + 0.99); II 5.66 (1.63 + 1.65 + 1.70 + 0.68); III 4.37 (1.30 + 1.28 + 1.31 + 0.48); IV 5.59 (1.70 + 1.62 + 1.62 + 0.65). Leg formula: 1243. Epigynum roughly rect- angular. Distribetioe. — -Only found in Yadong County, Tibet. ACKNOWLEDGMENTS Thanks due to Mark Harvey and Paula Cushing who gave invaluable assistance in the preparation of this manuscript, also to two anonymous referees for their valuable com- ments. We are grateful to J.X. Zhang and Z.S. Zhang for collecting some specimens and for their valuable advice. Many thanks are due to B.A. Huber and Saaristo for the provision of some valuable references. This work was sup- ported by Foundation of Hebei University (No. 2005409), China, and in part by the Nat- ural Science Foundation of Ikbei to F. Zhang (C2006000975). LITERATURE CITED Deeleman-Reinhold, C.L. 1986. Studies on tropical Pholcidae 11. Redescription of Micromerys grac- ilis Bradley and Calapnita vermiformis Simon (Araneae, Pholcidae) and description of some re- lated new species. Memoirs of the Queensland Museum 22:205-224. Gertsch, W.J. 1937. New American spiders. Amer- ican Museum Novitates 936:1-7. Hu, J.L. 1985. A new species of spider of the genus Spermophora from Xizang Autonomous Region, China (Araneae: Pholcidae). Acta Zootaxonom- ica Sinica 10:148-151. Hu, J.L. 2001. Spiders in Qinghai-Tibet Plateau of China. Henan Science and Technology Publish- ing House, 658 pp. Hu, J.L. & A.H. Li. 1987. The spiders collected from the fields and the forests of Xizang Auton- omous Region, China, IL Agricultural Insects, Spiders, Plant Diseases and Weeds of Xizang 2: 247-353. Huber, B.A. 1995. Copulatory mechanism in Hoi- ocnemus pluchei and Pholcus opilionoides, with notes on male cheliceral apophyses and stridu- iatory organs in Pholcidae (Araneae). Acta Zool- ogica, Stockholm 76:291-300. Huber, B.A. 1998. On the “valve” in the genitalia of female pholcids (Pholcidae, Araneae). Bulletin of the British Arachnological Society 11:41-48. Huber, B.A, 1999, Sexual selection in pholcid spi- ders (Araneae, Pholcidae): artful chelicerae and forceful genitalia. Journal of Arachnoiogy 27: 135-141. Huber, B.A, 2000. New World pholcid spiders (Ar- aneae: Pholcidae): a revision at generic level. Bulletin of the American Museum of Natural History 254:1-348. Huber, B.A. 2001. The pholcids of Australia (Ara- neae; Pholcidae): taxonomy, biogeography, and relationships. Bulletin of the American. Museum of Natural History 260:1-144. Koch, C.L. 1851. Ubersicht des Arachriidensys- terns. Vol. 5. J.L. Lotzbeck, Niimberg. Platnick, N.I. 2004. The world spider catalog, ver- sion 5.0. American Museum of Natural History, online at http: //research. amnh.org/entomology/ spiders/catalog/index. html Saaristo, M.I. 2001. Pholcid spiders of the granitic Seychelles (Araneae, Pholcidae). Phelsuma 9:9- 28. Schenkel, E. 1953. Chinesische Arachnoidea aus dem Museum Hoangho-Peiho in Tientsin. Bole- tim do Museu Nacional, Rio de Janeiro 119:1- 108. Senglet, A. 2001. Copulatory mechanisms in Ho- plopholcus, Stygopholcus (revalidated), Pholcus, Spermophora and Spermophorides (Araneae, Pholcidae), with additional faunistic and taxo- nomic data. Mitteilungen Der Schweizerischen Entomologischen Gesellschaft 74:43-67. Simon, E. 1893. Histoire Naturelle des Araignees. 2® edition, l(2):256-488. Librairie Encyclope- dique de Roret, Paris. Simon, E. 1903. Histoire Naturelle des Araignees. 2® edition, 2(4):669-1080. Librairie Encyclope- dique de Roret, Paris. Simon, E. 1909. Etude ser les Aracheides du Ton- kin (lere partie). Bulletin Scieetifique de la France et de la Belgique 42:69-147. Song, D.X., M.S. Zhu & J. Chen. 1999. The Spiders of China. Hebei Science and Technology Pub- lishing House, Shijiazhuang. Thorell, T. 1898. Secoedo saggio sui ragni Birmani. IL Retitelariae et Orbitelariae. Aenali del Museo Civico di Storia Naturale di Genova 39:271-378, Uhl, G., Huber, B.A. & Rose, W. 1995. Male ped- ipalp morphology and copulatory mechanism in ZHANG ET AL.— PHOLCIDS FROM TIBET 205 Pholcus phalangioides (Fuesslin, 1775) (Ara- neae, Pholcidae). Bulletin of the British Arach- nological Society 10:1-9. Walckenaer, C.A. 1805. Tableau des araneides ou caracteres essentiels des tribus, genres, families et races que renferme le genre Aranea de Linne, avec la designation des especes comprises dans chacune de ces divisions. Paris. Manuscript received 19 March 2004, revised 2 March 2005. 2006. The Journal of Arachnology 34:206-213 MAINOSA, A NEW GENUS FOR THE AUSTRALIAN ^SHUTTLECOCK WOLF SPIDER’ (ARANEAE, LYCOSIDAE) Volker W. Frameeau: Department of Terrestrial Invertebrates, Western Australian Museum, Welshpool DC, Western Australia 6986, Australia. E-mail: volker.framenau@museum.wa.gov.au ABSTRACT. A new monotypic genus, Mainosa, is described to accommodate the Australian ‘shuttle- cock wolf spider’, Mainosa longipes (L. Koch 1878) (= Lycosa mainae McKay 1979, new synonymy) as the type species. The male of this species is described for the first time. Mainosa longipes differs from other wolf spiders in having a the distinct color pattern of the abdomen, with white transverse bars and lines on a dark surface, and unusually long legs in males. Its genital morphology confirms M. longipes as a member of the subfamily Lycosinae. Mainosa longipes inhabits areas in South Australia and Western Australia with dry sandy soils in Acacia litter, where it constructs palisades around the entrance of its burrow. It appears to reproduce in winter. Keywords: Lycosinae, Australia, turret-building, palisade, taxonomy Australia has long been recognized for its unique fauna and flora. Two main reasons for Australia’s large number of endemic species are the long period of time it has been in geo- graphic isolation and its comparatively stable geological history (Hopper et ah 1996). Aus- tralia is thought to have split from the south- ern supercontinent Goedwana in the Late Pa- leocene, about 65 million years ago (e.g. Heatwole 1987), and some regions in Western Australia have not been subject to major geo- logical changes in the form of glaciation or continental uplifts (mountain building) since the Triassic more than 250 million years ago (e.g., Heatwole 1987; Hopper et al. 1996). Consequently, the spider fauna of Australia is very diverse and recent estimates suggest that some 20,000 species exist (Yeates et al. 2003). Wolf spiders belong to one of the predom- inant spider families in Australia. The pres- ence of large areas of open woodland, inland diffuse waterways (including salt lakes) and expansive arid and semi-arid regions that are all favored habitat areas of this family seems to account for this dominance (Main 1976, 1981). Recent studies have shown that the Australian and New Zealand wolf spider fau- nas contain some unique elements, such as the genera Artoria Thorell 1870, Anoteropsis L. Koch, 1878, Tetralycosa Roewer 1960, No- tocosa Vink 2002 and Venatrix Roewer 1960 (Framenau 2002, 2005; Framenau et al. 2006; Framenau & Vink 2001; Vink 2002). As part of his monumental monograph on Australian spiders Die Arachniden Austral- iens, L. Koch (1878) described the female of a wolf spider species with a very unusual col- or pattern of white transverse bars on an oth- erwise blackish-brown abdomen, Anoteropsis longipes L. Koch 1878. The type material of this species, part of the ‘Bradley Collection’, is considered lost (Framenau 2005). More than 100 years later, McKay (1979) described Lycosa mainae McKay 1979 from Western Australia with a very similar color pattern, also solely based on a single female and some immature spiders (Fig. 1). Earlier, Main (1976) had described the burrow of this spider and called this species the ‘shuttlecock wolf spider’ as these lycosids construct a palisade of litter around their burrow, reminiscent of a badminton shuttlecock (Fig. 2). A comparison of L. Koch’s (1878) and McKay’s (1979) de- scriptions of A. longipes and L. mainae strongly suggests that both species are actu- ally the same. After the examination of more than 15,000 Australian records of wolf spi- ders, I have found no other species that even remotely resembles that described by L. Koch (1878) and McKay (1979). An exhaustive investigation of the spider collection of the Western Australian Museum revealed some additional material of this spe- cies, including two males that match the fe- males of A. longipes and L. mainae. The ped- 206 FRAMENAU— NEW AUSTRALIAN WOLF SPIDER GENUS MAINOSA 207 Figure 1. — Penultimate female of Mainosa longipes from Lorna Glen Station, Western Australia (WAM T58395). The body length of this specimen is 10.5 mm. Figure 2. — Female of Mainosa longipes at the entrance of its burrow near Murchison Station, Western Australia (photograph courtesy of Fred and Jean Hort, Swan View). ipalp structure clearly identifies it as a member of the subfamily Lycosinae (sensu Dondale 1986). Consequently, the species cannot be a member of the genus Anoteropsis, as recently postulated in a revision of this ge^ nus (Vink 2002), since Anoteropsis is consid- ered member of an unnamed subfamily very different to the Lycosinae (Framenau et al. 2006). This species has also no somatic or genitalic similarities with Lycosa Latreille 1804, a putatively Mediterranean genus in which L. mainae was originally described (see e.g, Zyuzin & Logunov 2000). Here, L. mainae is considered a junior syn- onym of A. longipes. A new endemic Austra- lian genus, Mainosa, is erected to accommo- date the unusual 'shuttlecock wolf spider’ from Western Australia and South Australia, as it is not possible to place it in any other currently described genus within the Lycosi- dae. METHODS Descriptions are based on specimens pre- served in 70% ethyl alcohol. A female epi- gynum was cleared in lactic acid overnight for examination of the internal genitalia. The il- lustrations of epigyna and male pedipalps omit the setae for clarity. The morphological nomenclature follows Framenau & Vink (2001) and Framenau (2002). All measure- ments are in millimetres (mm). Since juvenile spiders can be clearly identified by the distinct color pattern, the species distribution is doc- umented based on both mature and immature specimens. Abbreviatioes.-^Eye^.- anterior (AE), an- terior median (AME), anterior lateral (ALE), posterior (PE), posterior mediae (PME), pos- terior lateral (PLE). Measurements (adult spi- ders, if not otherwise stated): total length (TL), carapace length (CL) and width (CW), abdomen length (AL) and width (AW). Col- lections: QM = Queensland Museum, Bris- bane; WAM = Western Australian Museum, Perth. SYSTEMATICS Subfamily Lycosinae Sumderall 1833 Mainosa new genus Type species.“-”A«otero/?5iA longipes L. Koch 1878. Etymology. — The genus is named in honor of Barbara York Main. Barbara’s contribution to Australian aracheology is legendary, and after two years at the Western Australian Mu- seum, I still have not finished sorting through her part of the immense wolf spider collec- tion. It also preserves McKay’s (1979) ac- knowledgment of Barbara’s contribution to the naming of L. mainae. The gender is fem- inine. Diagnosis. — Mainosa can be distinguished from ail other known wolf spiders by the unique coloration of the abdomen, consisting II 208 of white transverse bars and lines on a brown to black surface (Fig. 1). In addition, males have unusually long legs, with a large ratio of leg length to carapace width (WAM T62713; leg 1 - 7.0; leg 2 = 6.4; leg 3 = 6.13; leg = 8.6). For example this leg ratio is twice as large as the average for the species within the Australian lycosine genus Venatrix (leg 1 = 3.6; leg 2 = 3.2; leg 3 = 3.0, leg 4 = 4.2; data for 17 species derived from Framenau & Vink 2001). Description.- — ^Medium sized wolf spiders (XL 6“ 15 mm). Males smaller than females. Carapace elevated in head region, more prO“ noueced in males than females (Fig. 3). Row of AE narrower than row of PME. Row of AE slightly procurved (Fig. 4). Capet flanks steep in males (Fig. 4), but a gentle slope in fe- males. Spiders overall very dark, reddish- brown to black, however, the carapace has white setae medially resulting in a distinct me- dian band in live specimens (Fig. 1). Abdo- men with distinct light transverse bars and lines on a dark surface (Fig. 1). Chelicerae with three promargieal and three retromargin- al teeth. Leg formula IV>I>II>III. Males with very long and thin legs, each at least 6 times as long as the carapace width. Females with dense scopulae on the tarsi of all legs, metatarsi Fill and the apical two thirds of tib- iae I +11. Triangular mediae apophysis of male pedipalp directed retrolaterally and without ventral process (Fig. 5). Bulb rotated slightly clockwise, so that the large subtegulum is sit- uated prolaterally (rather than basally) and the base of the embolus apically (rather than ap- icoprolateral) (Fig. 5). Terminal apophysis sickle-shaped (Fig. 7). Female epigyum with inverted T-shaped median septum, of which the longitudinal part is indistinct and the lat- eral edges of the posterior transverse part are slightly bent anteriorly (Figs. 8, 10). Included species. — Mainosa longipes (L. Koch 1878). Distribution. — -As for species (Fig. 11), Mainosa longipes (L. Koch 1878), NEW COMBINATION (Figs. 1-11) Anoteropsis longipes L. Koch 1878: 973-974, plate 85, figs. 2, 2a. Lycosa maini McKay 1979: 260-263, figs. 7a-e; McKay, 1985b: 80. NEW SYNONYMY. Lycosa mainae McKay.- Platnick 1989: 372. THE JOURNAL OF ARACHNOLOGY j Tjpes.-—Anoteropsis longipes: holotype fe- ji male, Australia, locality not given in L. Koch (1878), Bradley Collection (presumed lost, see Framenau in press; not examined), Lycosa 1 mainae: holotype female. Western Australia, j 88 km N. -of Murchison River, 27°42'S, | 114°09'E, 30 January 1969, on red soil with mulga, Acacia aneura, in turret burrow, R.J. McKay (WAM 69/115), examined. Paratypes: AUSTRALIA: Western Australia: 1 immature j female, Billabong Roadhouse, near Shark Bay |j turn-off, 26°49'S, 114°36^E, 5 December |! 1972, R.J. McKay (QM W4668); 1 juvenile, | 5 miles N. of Menzies, 29°4rS, 121°02'E, 1 | September 1954, FN24, B.Y. Main (WAM 68/ | 821); 2 penultimate females, Mount Magnet | area, 28°05'S, 117°52'E, station 7, 323 mile ! peg Mt Magnet, 7-8 December 1968, R.J. j McKay, J. Gilbert, J. Ayres (WAM 69/1031, ; 69/1036); 1 juvenile, Murchison, 19 km N., | 29°38'S, 115°57'E, 20 February 1962, A.R. j Main (WAM 68/820); 2 juveniles, Norseman, | 76 km N., 3r33'S, 12r47'E, 26 December 1968, W.H. Butler (WAM 69/105-6); 1 Ju- venile, 220 mile peg, Paynes Find, 29°15'S, 117°4rE, 8 December 1968, R.J. McKay, J. ' Gilbert, P. Snowball (WAM 68/819); 1 juve- I nile, Tarin Rock Reserve, 33°06'S, 118°1LE, , 22 May 1971, palisade burrows on loam and ! litter, A. Baynes (W^AM 71/1859); 1 penulti- mate female, 1 penultimate male, Wubin, 32 km N.E., 30°06'S, 116°37'E, 14 July 1968, in large turret burrow, R.J. McKay, J. Gilbert, J. Ayres (WAM 68/817-8). All paratypes ex- amined. Other material examined.— AUSTRA- LIA: South Australia: 1 penultimate d, Dub- lin, 34°27'S, 138°2rE, 16 May 1986, B.Y. Main, FNl 1 (WAM T627 1 8); 1 juvenile, Mal- labie Shed Tank, 18 miles W., 3r28'S, 130°20'E, 23 December 1952, B.Y. Main, FN40 (WAM T62719). Western Australia: 1 juvenile, Arnolds Tank (PWD tank 488), N= of Wialki, 28°39'S, 122°36'E, 24 April 1957, A.R. Main, FNl, BYM 1957/AlO, palisade, thick litter under Acacia (WAM T46842); 1 d, Francois Peron National Park, 25°52'31"S, 113°32'59"E, 24 August-10 November 1994, wet pitfall traps, A. Sampey et al., WAM/ CALM Carnarvon Survey (WAM T48038); 1 juvenile, Francois Peron National Park, 1.6km NW of Monkey Mia Road, 25°50^20"S, 113°36'23"E, 18 January-23 May 1994, wet pitfall traps, M.S. Harvey et ah, WAM/CALM FRAMENAU— NEW AUSTRALIAN WOLF SPIDER GENUS MAINOSA 209 tegulum A / I cym^m sub" \ /: tegulum / .- \ terminal terminal palea apophysis embolus median \ ^ apophysis / 8 median septum spermatheca Figures 3-10. — Mainosa longipes (L. Koch, 1878): Male from ‘Sieda’, near Grass Patch, Western Australia (WAM T62713): 3. Carapace, lateral view; 4, Eyes, frontal view; 5. Left pedipalp, ventral; 6. Left pedipalp, retrolateral; 7. Apical part of bulb. Female holotype of Lycosa mainae, 88 km N. of Murchison River, Western Australia (WAM 69/115): 8. Epigynum, ventral view; 9. Epigynum, dorsal view. Female from Nerren Nerren, Western Australia (WAM 94/1940): 10. Epigynum, ventral view. Scale bar: (3) 2.70 mm, (4) 1.44 mm, (5-6) 0.66 mm, (7) 0.42 mm, (8-10) 0.80 mm. 210 THE JOURNAL OF ARACHNOLOGY Carnarvon Survey, site PE4 (WAMT62720); Id, ‘Sieda’, near Grass Patch, Fitzgerald Lo- cation 41, 33°13^56"S, 121°46'00"E, 20 Sep- tember 1988, A.E Longbottom, in house on desk near telephone, S233 (WAM T62713); 1 juvenile. Grass Patch, E. of, Fitzgerald Loca- tion 71, 33°13'S, 12r43'E, 17 December 1980, A.E Longbottom, web-lined burrow amongst leaf litter, S62 (WAM T53625); 1 ju- venile, Gutha, 15 miles N., 28°44'S, llb^OUE, 18 August 1953, B.Y. Main, palisade in Aca- cia litter, no silk, FN5 (WAM T53473); 1 pen- ultimate 9 , Kellerberrin, 3r33'S, 117°43'E, 1 April 1993, G.T. Smith, leaves on entrance, M17 (WAM T62714); 1 penultimate 9, Lorna Glen Station, 26°19'S, 12r02'E, 28 April 2004, K.E.C, Brennan, G. Owen, M. Moir, PR. Langlands (WAM T58395); 1 juvenile, Morawa, S.E., nature reserve on Lochada Road, 29°15'29"S, 116°2r43"E, 12 October 1999, B.Y. Main, dug from burrow with pal- isade (WAM T62716); 1 9, Nerren Nerren Station, 4.0km E of Nerren Nerren boundary fence, E of North West Coastal Highway, 27°00'21"S, 114°32'29"E, 15 October 1994, J.M. Waldock, J. Riley, dug from burrow with palisade, WAM/CALM Carnarvon Survey, site NE 4 (WAM 94/1940); 1 penultimate 9, Nerren Nerren Station, 5.9km E of Nerren Nerren boundary fence, E of North West Coastal Highway, 27°03'28"S, 114°36'25"E, 18 October 1994, J.M. Waldock, WAM/CALM Carnarvon Survey, site NE5 (WAM 94/1941); 1 juvenile, Norseman, 29.2 miles E., on Eyre Highway, 32°12'S, 122°17'E, 8 December 1953, B.Y Main, FN8 (WAM T46843); 1 9, North West Coastal Highway, 20.3km E, on Woodleigh— Byro Road, 22°12'31"S, 114°34'35"E, 12 October 1994, M.S. Harvey et ak, WAM/CALM Carnarvon Survey, site W02, from burrow with Acacia leaves pali- sade (WAM T62730); 1 juvenile, Oudabunna Station, 15.9km S. of Wydgee Homestead, 29°04'S, 117°45'E, 8 August 1982, B.Y. Main, FN13 (WAM T62715); 1 penultimate 9, Pay- nes Find, 29°15^S, 117°41'E, 1 August 1982, B.Y Main, FN14 (WAM T62717). Diagnosis. — Mainosa longipes displays a unique color pattern among known wolf spi- ders. The abdomen is dark brown to black, with white transverse bars and lines in the posterior half of the abdomen (Fig. 1), less distinct in males. Description. — Male (based on WAM FRAMENAU=— NEW AUSTRALIAN WOLF SPIDER GENUS MAINOSA 211 T62713): Carapace: head region strongly ele- vated (Fig. 3); overall dark brown, medially in front of fovea slightly lighter; indistinct dark radial pattern; covered with mainly sil= ver-white setae, that are particularly dense be- tween eyes and towards the carapace margins; some black setae between median and lateral bands on carapace flanks; one long brown bristle between AME, six long brown bristles below AE; clypeus high, more than one di- ameter of AME (Fig. 4). Eyes: row of AE shorter then row of PME; row of AE procur- ved (Fig. 4). Sternum: light brown with dense, black pigmentation; covered with brown bris- tles, which are longer towards the margin. La- bium: brown; front end truncate and white. Chelicerae: dark brown with a dark longitu- dinal band; a few white setae and, medially, a few long brown bristles; three retromarginal teeth, with the apical slightly smaller; three promarginal teeth, with the median largest. Pedipalp (Figs. 5~7): embolus long and slen- der with its tip pointing slightly apically, ter- minal apophysis sickle-shaped (Fig. 7). Ab- domen: very dark grey with indistinct light lanceolate heart mark in anterior half, its front end a more distinct orange patch; orange- whitish transverse bar medially and some thin transverse orange-whitish lines in posterior half; white setae medially in a band that wid- ens posteriorly, otherwise brown setae; venter uniformly dark brown, laterally with irregular light spots; spinnerets yellow brown. Legs: leg formula IV > I > II > III; uniformly dark brown, coxae ventrally yellow-brown; spina- tion of leg I: femur: 2 dorsal, 1 apicoprolater- ai; tibia: 3 ventral pairs, 2 prolateral; metatar- sus: 3 ventral pairs, 1 retrolateral, 1 apicoventral, 1 apicoprolateral, 1 apicoretro- lateral. Female (based on holotype of L. mainae WAM 69/115): Carapace: very dark reddish- brown with indistinct radial pattern; most se- tae rubbed off, some silver-white setae on car- apace flanks and between eyes. Eyes: row of AE shorter than row of PME, row of AE slightly procurved. Sternum: brown, long brown setae of increasing length and density towards margins. Labium: as in male. Chelic- erae: black, few brown setae medially, denti- tion as in male, Epigynum (Figs. 8~10): ven- tral view: longitudinal part of median septum indistinct, but with distinct posterior trans- verse part of which the lateral ends are bent forward; no anterior hoods (Figs. 8, 9); dorsal view: small spermathecae with dorsal appen- dix (Fig. 10). Abdomen: very dark brown with a wide light-brown patch anteriorly; white transverse bars in posterior half; covered in brown setae, whitish setae in transverse bars, some longer light brown setae in area of lan- ceolate heart mark. Venter as in male; spin- nerets brown. Legs: leg formula IV > I > II > III; uniformly dark brown, apical segments somewhat darker; dense scopulae on tarsi, metatarsi and apical two thirds of tibiae of leg I and II, on tarsi and metatarsi of leg III and tarsi of leg IV; spination of leg I: Femur: 3 dorsal, 2 apicoprolateral, 1 apicoretrolateral; patella: 1 prolateral; tibia: 3 ventral pairs, 1 prolateral; metatarsus: 3 ventral pairs, 1 api- co ventral. Measurements: Male, WAM T62713 (fe- male holotype of L. mainae, WAM 69/115): TL 5.92 (14.10), CL 3.24 (6.58), CW 2.12 (4.51). Eyes: AME 0.19 (0.22), ALE 0.11 (0.22), PME 0.32 (0.70), PLE 0.25 (0.48). Row of eyes: AE 0.68 (1.41), PME 0.79 (1.60), PLE 0.95 (1.83). Sternum (length/ width) 1.49/1.27 (2.54/1.97). Labium (length/ width) 0.36/0.44 (0.71/0.89). AL 2.26 (6.96), AW 1,97 (5.17). Legs: lengths of segments (femur + patella/tibia + metatarsus A tarsus — total length): Pedipalp 1.41 + 1.41 + — + 0.94 - 3.76, I 3.81 -f 4.65 + 4.09 + 2.26 = 14.81, II 3.38 + 4.37 + 3.81 + 1.97 = 13.53, III 3.24 + 4.09 A 3.81 + 1.83 = 12.97, IV 4.65 + 5.50 + 5.78 + 2.26 = 18.15 (Pedipalp 2.54 + 2.63 +— + 2.07 = 7.24, I 4.70 + 5.64 + 3.48 + 1.88 = 15.70, II 4.51 + 5.36 + 3.29 -f 1.79 - 14.95, III 3.85 + 4.51 + 3.29 + 1.69 - 13.34, IV 5.08 + 6.67 + 5.92 + 3.10 = 20.77). Variation: The dimensions of a second male, WAM T48038 (2 females, WAM 94/ 1940 and WAM T62730) are: TL 9.31, CL 5.08, CW 3.29 (TL 12.41, CL 6.77, CW 4.98 and TL 13.24, CL 6.49, CW 4.04). Remarks. — The female holotype of Ano- teropsis longipes, described from Bradley's Collection, is considered lost (Framenau 2005). However, L. Koch's (1878) description of the distinct color pattern of this species and his illustration of the female genitalia allow an accurate identification of this species. Lud- wig Koch (1878) did not give any locality for his specimen, however, it is not unlikely that the spider was from Western Australia as the 212 THE JOURNAL OF ARACHNOLOGY Bradley Collection included other spiders col- lected in this state, e.g., the type material of Tetralycosa oraria (L. Koch 1876) from King George Sound near Albany (L. Koch 1876; see also Framenau et al. 2006). Habitat preferences and life cycle. — Mai- nosa longipes appears to prefer open Acacia woodland and mallee with red clay to sandy soils (McKay 1979). Here, it constructs pali- sades of elongate leaves or phyllodes around the mouth of its burrow (Fig. 2). Palisades are generally constructed in heavy leaf litter be- low shrubs and trees, usually on the side of the tree where the afternoon sun falls (McKay 1979). A penultimate and a mature male were caught in May, a second mature male in Sep- tember which suggests that this species is re- productively active in winter. Distribution. — South Australia and West- ern Australia south of 25°S latitude (Fig. 11). DISCUSSION Mainosa longipes belongs to the subfamily Lycosinae as the male pedipalp has a trans- verse median apophysis with a sinuous chan- nel on its dorsal surface (Dondale 1986). The closest relatives may be found in the genus Dingosa Roewer 1955, represented by Din- gosa simsoni (Simon 1898) and the currently misplaced Australian lycosines 'Pardosa' ser~ rata (L. Koch 1877) and ‘P.’ Humphrey si McKay 1985a. Similar to Mainosa, Dingosa species construct turrets around their burrows and males have extremely elongated legs. However, genital morphology and coloration of Dingosa differ considerably from Mainosa. The male pedipalp in this genus has a large palea region with a broad, truncated (not sick- le-shaped) terminal apophysis. The median apophysis of these species is not triangular, but slim and elongated apically. In addition, the coloration of the Australian Dingosa is very different as the abdomen displays a char- acteristic serrated pattern with dark chevrons but no transverse bars. A revision of this ge- nus, that contains a further two undescribed Australian representatives, is forthcoming. Turret-building is not only restricted to Australian wolf spiders. Some species of the Holarctic genus Geolycosa Montgomery 1904, such as G. missouriensis (Banks 1895), also construct palisades around the opening of their burrow entrance (Wallace 1942; G. Strat- ton pers. comm.). The New Zealand Notocosa bellicosa (Goyen 1888) extends the opening i of its burrow with a rim of silk into which it | incorporates pieces of debris (Vink 2002). The benefits of these palisades are currently un- known. In mygalomorph spiders of the genus ' Aname L. Koch 1873, burrow turrets appear i to have some significance in relation to reg- ular sheet-flood events (Main 1993). Alterna- tive functions may include a barrier against debris that could otherwise fall into the bur- row. The palisades could also play an impor- tant role in foraging. Prey may be attracted to the palisade as an elevated resting place and the turret also provides the spider with a van- tage point since they can be seen sitting on the top of the turret during the day (pers. obs.). Finally, palisades may have an impor- tant thermoregulatory function such as to avoid hot surface air to penetrate the burrow. ACKNOWLEDGMENTS I am indebted to Barbara Main for sharing | her immense arachnological knowledge with j me every Thursday morning, and Julianne Waldock and Mark Harvey for their never- ! ending support at the Western Australian Mu- ‘ seum. I thank Barbara Baehr, Robert Raven j and Owen Seeman for their hospitality whilst i sorting through the collection of the Queens- ’ land Museum in Brisbane, where one of the paratypes of Lycosa mainae is housed. I am grateful to Fred and Jean Hort from Swan | View in Western Australia for permission to use their photograph of M. longipes (Fig. 2) j in this publication. Peter Langlands provided ' me with the live specimen of M. longipes il- | lustrated in Fig. 1. Melissa Thomas, Julianne Waldock, Mark Harvey, Torbjprn Kronestedt | and Cor Vink provided helpful comments on : earlier drafts of this manuscript. This study forms part of a revision of the Australian wolf | spiders that is funded by the Australian Bio- logical Resource Studies (ABRS) to Mark Harvey (Western Australian Museum) and Andy Austin (University of Adelaide). LITERATURE CITED j Dondale, C.D. 1986. The subfamilies of wolf spi- ders (Araneae: Lycosidae). Actas X Congreso In- ternacional de Aracnologia, Jaca, Espana 1:327- ^ 332. : Framenau, V.W. 2002. Review of the genus Artoria \ Thorell (Araneae: Lycosidae). Invertebrate Sys- ! tematics 16:209-235. ' Framenau, V.W. 2005. The genus Artoria in Aus- FRAMENAU— NEW AUSTRALIAN WOLF SPIDER GENUS MAINOSA 213 tralia: new synonymies and generic transfers (Ar- aneae, Lycosidae). Records of the Western Aus- tralian Museum 22:265-292. Framenau, VWI & CJ. Vink. 2001. Revision of the genus Venatrix Roewer (Araneae: Lycosidae). Invertebrate Taxonomy 15:927-970. Framenau, V.W, TB. Gotch & A.D. Austin, 2006. The wolf spiders of artesian springs in arid South Australia, with a revalidation of Tetralycosa (Ar- aneae, Lycosidae). Journal of Arachnology 34: 59-94. Heatwole, H, 1987. Major components and distri- butions of the terrestrial fauna. Pp. 101-135. In Fauna of Australia Volume lA. General Articles. (D.W. Walton, ed.). Australian Government Pub- lishing Service, Canberra. Hopper, S.D., M.S. Harvey, J.A. Chappill, A.R. Main & B.Y. Main. 1996. The Western Austra- lian biota as Gondwanan heritage— a review. Pp. 1-46. In Gondwanan heritage: past, present and future of the Western Australian biota. (S.D. Hopper, J.A Chappill, M.S. Harvey & A.S. George, eds). Surrey Beatty & Sons, Chipping Norton. Koch, L. 1876. Die Arachniden Australiens, nach der Natur beschrieben und abgebildet. Bauer and Raspe, Niimberg. Vol. 1, pp. 741-888. Koch, L. 1877. Die Arachniden Australiens, nach der Natur beschrieben und abgebildet. Bauer and Raspe, Niimberg. Vol. 1, pp. 889-968. Koch, L. 1878. Die Arachniden Australiens, nach der Natur beschrieben und abgebildet. Bauer and Raspe, Niimberg. Vol. 1, pp. 969-1044. Latreille, RA. 1804. Tableau methodique des in- sects. Noveau Dictionnaire d’Histoire Naturelle Paris 24:129-295. Main, B.Y. 1976. Spiders. Collins, Sydney. Main, B.Y. 1981. Australian spiders: diversity, dis- tribution and ecology. Pp. 809-852. In Ecologi- cal Biogeography of Australia. (A. Keast, ed.). Dr. W. Junk Publishers, The Hague, Boston, Lon- don. Main, B.Y. 1993. From flood avoidance to foraging: adaptive shifts in trapdoor spider behaviour. Memoirs of the Queensland Museum 33:599- 606. McKay, R.J. 1979. The wolf spiders of Australia (Araneae: Lycosidae): 12. Descriptions of some Western Australian species. Memoirs of the Queensland Museum 19:241-275. McKay, R.J. 1985a. The wolf spiders of Australia (Araneae: Lycosidae): 1. A new species of Par- dosa. Memoirs of the Queensland Museum 22: 101-104. McKay, R.J. 1985b. Lycosidae. Pp. 73-88. In Zoo- logical Catalogue of Australia, Vol. 3. Arachni- da, Mygalomorphae, Araneomorphae in Part, Pseudoscorpionida, Amblypygida, Palpigradi (D.W. Walton, ed.). Australian Government Pub- lishing Service, Canberra. Platnick, N.I. 1989. Advances in Spider Taxonomy, 1981-1987. Manchester University Press, Man- chester. Vink, C.J. 2002. Fauna of New Zealand. Number 44. Lycosidae (Arachnida: Araneae). Manaaki Whenua Press, Lincoln (New Zealand). Wallace, H.K. 1942. A revision of the burrowing spiders of the genus Geolycosa (Araneae, Lycos- idae). American Midland Naturalist 27:1-62. Yeates, D.K., M.S. Harvey & A.D. Austin. 2003. New estimates for terrestrial arthropod species- richness in Australia. Records of the South Aus- tralian Museum Monograph Series 7:231-242. Zyuzin, A. A. & D.V. Logunov. 2000. New and little-known species of the Lycosidae from Azer- baijan, the Caucasus (Araneae, Lycosidae). Bul- letin of the British Arachnological Society 11: 305-319. Manuscript received 10 November 2004, revised 3 March 2005. 2006. The Journal of Arachnology 34:214-220 ECOLOGY OF THESTYLUS AURANTIURUS OF THE PARQUE ESTADUAL DA SERRA DA CANTAREIRA, SAO PAULO, BRAZIL (SCORPIONES, BOTHRIURIDAE) Humberto Y. Yamaguti and Ricardo Pinto-da-Rocha: Departamento de Zoologia, Institute de Biociencias, Universidade de Sao Paulo, Caixa Postal 11461, CEP 05422-970, Sao Paulo, SP, Brazil. E-mail: humbertotete@yahoo.com.br ABSTRACT. Individuals of a Thestylus aurantiurus Yamaguti & Pinto-da-Rocha 2003 population in the Parque Estadual da Serra da Cantareira (Sao Paulo, SP, Brazil) show an increase of activity throughout the year. This increase is related to the reproductive season of these scorpions, from September to No- vember. The abundance of scorpions was related to environmental factors in four different areas of the park. More scorpions were collected in the higher areas, far away from water sources of the park and not exposed to flooding. A short description of the Thestylus aurantiurus burrows is also presented. Keywords: Scorpiones, Thestylus aurantiurus, Atlantic Rainforest, seasonality, relative abundance Scorpions are extremely sedentary animals. According to Polis (1990a), they share the re- cord with many spiders for the lowest arthro- pod metabolic rates ever recorded. Some spe- cies spend 97% of their lives inside their burrows, and they can exist one year without feeding (Polis 1990a). This is largely due to the “sit-and-wait” type of foraging activity adopted by most scorpions, which consists of being immobile and awaiting any prey that passes within reach. However, during the re- productive period, scoipions present certain behavioral changes. These changes occur mainly in surface activity, such as a reduction in female foraging activity and an increase in male surface activity in search of females (Benton 2001). Courtship behavior is well-studied in scor- pions. The behavior of 35 species of six fam- ilies was described, from the about 1500 spe- cies and 16 families listed in Fet et al. (2000). The courtship behavior of many bothriurid species was studied, mainly by Peretti (1995, 1997). However, there are few works on the reproductive activity of scorpions. Peretti (1997) studied the reproductive characteristics of Argentinean scorpions, describing the re- productive period, interval between mating and litter birth behavior, and period of litter birth of some bothriurid scorpions. Matthiesen (1968) studied the courtship behavior of cer- tain Brazilian scorpions during courtship, e.g., Bothriurus araguayae Vellard 1934. The genus Thestylus Simon 1880 has been little studied due to its restricted geographical distribution and low abundance. The only study on the reproduction of Thestylus was Machado & Vasconcellos-Neto (2000). They described the courtship behavior of T. auran- tiurus Yamaguti & Pinto-da-Rocha 2003 (mentioned as T. glasioui (Bertkau 1880)). According to the authors, the reproductive pe- riod is very seasonal, occurring during the hot and wet season (beginning in October), in Ser- ra do Japi (23°17'S; 47°00'W), Jundiai, SP, Brazil. Abundance and its relationship to environ- mental factors is another well studied aspect of scorpion ecology. Evidence suggests that scorpions are selective when choosing a place to build their burrows. Polis & McCormick (1986) studied desert scorpions and noticed differences of abundance among the species related to the environment. Hofer et al. (1996) presented a study relating environmental as- pects of different habitats of a Central Ama- zon Rainforest with the relative abundance of a certain scorpion species. Koch (1977, 1978) studied burrows in the genus Urodacus Peters 1861 (Urodacidae), observing a different kind of burrow for each Urodacus species and ver- ifying that environmental aspects influence burrow site location. 214 YAMAGUTI & PINTO DA-ROCHA— ECOLOGY OF THESTYLUS AURANTIURUS 215 The study presented here was conducted to assess the activity of a population of Thestylus aurantiurus (Figs. 1 & 2) throughout the course of one year (February 2000-January 2001), and the relationship between relative abundance and environmental aspects in Parque Estadual da Serra da Cantareira, Sao Paulo, SP, Brazil. METHODS Studied area* — Collections were made throughout one year in four areas of the At™ lantic Rainforest. These areas are located in the Nucleo Pedra Grande of the Parque Estad- ual da Serra da Cantareira (23°22'S; 46°36'W), Sao Paulo, SP, Brazil, in a project conducted together with Universidade Ban- deirante de Sao Paulo (UNIBAN). Area 1 named Pedra Grande (23°26'30"S; 46°38'20"W at an elevation of 1050 m), pos- sesses dense vegetation, and well-developed small and medium-sized understory strata, with lianas and bamboo groves. In area 2, named Lago das Carpas (23°25'40"S; 46°36'06"W, 700 m) possesses dense under- story, with flooded areas and bamboo groves. Area 3 named Divisa (23°25'48"S; 46°38'00"W, 1050 m), is located on the boundary between the municipal districts of Sao Paulo and Mairipora. It possesses dense vegetation, with numerous lianas and some exotic species, such as Pinus sp. Area 4, named Sede (23°27^03"S; 46°38'06"W, 650 m), is characterized by lush vegetation, with steep slopes, many lianas, a large amount of low vegetation and flooded areas. The values of monthly median temperatures and monthly total rainfall presented in this work were supplied by the Companhia de Sa- neamento Basico do Estado de Sao Paulo (SABESP). These data were recorded at Paiva Castro^s dam (23°22'11"S; 46°40'07"W), in the municipal district of Mairipora, near to the Parque Estadual da Serra da Cantareira. Ecological samplings. — Twelve monthly collections, of nine days each, were undertak- en between February 2000 and January 2001. Scorpions were collected in drift fence pitfall traps (Fig. 3). Traps were composed of groups of four buckets of 20 L each, buried with the opening nearby ground level, arranged in a “Y” shape. Each “arm” of the “Y” was 5.0 m from the central point. One bucket was bur- ied at each tip of the “Y” and one at the cen- tral point. Strips of canvas of 5.0 X 0.5 m, fixed with wooden stakes, connected each out- lying bucket to the central one in order to steer the animals into the buckets. Ten traps were placed in each area, comprising a total of 40 buckets per area. The central point of each trap was 20 m from the next in a straight line. The bottom of each bucket was perforated to prevent the accumulation of rain water. The size of the holes prevented the animals from escaping. Traps were checked every morning and covered on the last day to avoid captures out- side the sampling period. Each trap was num- bered individually, enabling the identification of scorpions according to trap, date and col- lection area. Since the traps were originally erected to capture small mammals alive, a pre- serving liquid was not used, hence animals re- mained alive until the moment of capture. Thus, there was a possibility of the scorpions having been devoured by toads, lizards or ro- dents inside the buckets. So, the number of collected scorpions may be higher than the ob- served. However, the results probably would not change, because the patterns of seasonality are clearcut in this work. This kind of sampling probably produced a different result from active collection (e.g. with ultraviolet light), but with pitfall traps, the seasonality would be more evident. Preg- nant females were identified according to Far- ley (2001), with dissections and observation of the ovariuterus, and comparison with Far- ley’s photos. Voucher specimens were depos- ited in Museu de Zoologia da Universidade de Sao Paulo (MZSP), Sao Paulo, SP, Brazil. Environmental data. — The following en- vironmental data were gathered: amount of lit- ter, density of the canopy, perimeter of closest 10 trees measured at the breast height (PBH), soil composition and elevation. All these data were recorded on the same day, in May 2002 (subsequent to the period of collection), close to the places where traps were located. The amount of litter of an area of 0.25 m^ was measured. Ten samples were collected ran- domly in each area totaling 40 samples. Each sample was placed in a stove for 48 hours and its weight was measured soon after. The den- sity of the canopy was measured using a den- siometer at 24 points in each area, totaling 96 samples. The PBH was measured at 24 points in each area, totaling 960 trees. To analyze 216 THE JOURNAL OF ARACHNOLOGY Figure 1, 2. — Thestylus aurantiurus. 1. Male, 2. Female, YAMAGUTI & PINTO-DA-ROCHA— ECOLOGY OF THESTYLUS AURANTIURUS 217 Figure 3. — Schematic representation of the drift fence pitfall traps used in sampling in P. E. Serra da Cantareira, Brazil. soil composition, samples were collected at 8 points in each area, totaling 32 samples. The percentage of organic matter and the vegeta- tional covering were measured. Statistical analysis. — In the statistical anal- ysis of the number of animals collected in each area and to verify whether environmental factors differ in the four areas, the Kruskal- Wallis test was utilized with a — 0.05. For a comparison between the number of animals collected in the higher areas and the number of animals collected in the lower areas, a Mann- Whitney test was used with a = 0.05. Before those statistical analysis, the normality and the homogeneity of variances were tested. The data were ranked in both cases. In order to verify whether there is a relationship be- tween these factors and the number of col- lected animals in each area, the Spearman Rank Correlation was utilized with n = 4. RESULTS/ Burrows of Thestylus aurantiurus. — Indi- viduals of T. aurantiurus in Parque Estadual Intervales (Sao Paulo, SP, Brazil) were ob- served foraging at the entrance of their bur- rows, in ravines near river margins, with ped- ipalps and anterior body outside (G. Machado, pers. com.), in a typical sit-and-wait foraging type. Burrowing activity of T. aurantiurus was observed in captivity, where all individ- uals constructed many burrows, apparently looking for the best place to stay. The burrows were simple, with a single vertical or oblique tunnel (1. 5-3.0 cm deep), and a horizontal re- gion at the end (about 1.5 cm long). The en- trance could be circular or semicircular in shape, with 1.0-1. 5 cm of maximum width. Seasonality. — Thestylus aurantiurus was the only scorpion species recorded in the park. Between January (summer) and July (winter). few individuals were collected (from 0-2 scorpions/month in 40 traps). Four males were collected in August, In the three subsequent months (spring), 18, 36 and 11 scorpions were collected, respectively (about 80% of the total sampled). Twelve females were collected in October (about 70% of the total sampled fe- males). In the remaining months of the year, one female was the maximum number cap- tured per month (Fig. 4). Four pregnant females were found in Oc- tober and one pregnant female in November 2000. No fecund females were found in the other months. Relative abundance in four areas. — The individuals collected possessed a heteroge- neous distribution in the four areas (K.W., H — 14.966; P = 0.002). The number of indi- viduals collected in the two higher areas (1 & 3) was significantly higher than the number of individuals collected in the two lower areas (2 & 4) (Mann-Whitney test, U = 79.500; P = 0.001). The amount of litter was different in the four areas (area 1 = 166.9 g, area 2 — 339.1 g, area 3 = 190.5 g, area 4 — 132.7 g; K.W., H - 18.00; df= 3; P = 0.0004; n = 40), but only area 2 presented a different amount of litter, the amount of the other three areas are very similar (according to the Tukey HSD test). Canopy density was different in the four areas (area 1 = 4.625, area 2 = 8.5, area 3 = 8.71, area 4 = 9.92; K.W., H = 22.46; df = 3; P < 0.0001) but it was not related to the number of scorpions collected (r^ = 0.40; n = 4; P = 0.60). The PBH was also different in the four areas (area 1 = 14.09 cm, area 2 = 18.25 cm, area 3 = 16.9 cm, area 4 = 18.86 cm; K.W., H = 18.81; df = 3; P = 0.0003) but not related to the number of scorpions col- lected (r^ = 0.21; w = 4; P = 0.79). Soil type was the same in the four areas. The number of scorpions collected was not significantly re- lated to the elevation (r^ = 0.74; n = 4\ P = 0.26). Air temperature and rainfall were not related to abundance, although the number of scorpions increased immediately after the coldest month. DISCUSSION Reproductive activity. — The population of Thestylus aurantiurus in the Parque Estadual da Serra da Cantareira presents differentiated activity throughout the year suggesting the ex- 218 THE JOURNAL OF ARACHNOLOGY Months 4 .B & ■ " temperature "# rainfall Months 5 Figures 4, 5. — 4. Seasonality of the scorpion population of R E. Serra da Cantareira, Brazil (February 2000-January 2001), with the total number of individuals collected by month. The columns represent the number of males and females collected by month. 5. Means of temperature (in °C) and total rainfall (in millimeters) by month in the P. E. Serra da Cantareira, Brazil (February 2000-January 2001). istence of a reproductive season. The repro- ductive season lasts from September to No- vember, with an activity peak in October (Fig. 4), in the beginning of the hot, wet season (Fig. 5). This can be evidenced by the fact that fertilized females were found in October and November of 2000. Ten of the 12 females collected in October were collected together with males. These fe- males may have fallen into traps while they YAMAGUTI & PINTO-DA-ROCH A— ECOLOGY OF THESTYLUS AURANTIURUS 219 were dancing with males in search of a place to deposit the spermatophore. The sit-and-wait foraging strategy of this species consists of remaining immobile, waiting for prey. This foraging strategy type does not require the scorpion to move around (Polis 1990a), lead- ing us to conclude that most of scorpions col- lected in this study were in a search of part- ners and not prey. Corey & Taylor (1987) collected scorpions in pitfall traps, close to Orlando, Florida, U.S.A. during one year, sampling every two months. Only one species was collected, the buthid Centruroides hentzi (Banks 1900). This is an errant forager but also presenting an in- crease in activity throughout the year, in July and September, apparently during the repro- ductive period. The reproductive period in scorpions varies according to the species. Some South-Ameri- can scorpions such as Tityus bahiensis (Perty 1833) do not exhibit a well-defined reproduc- tive period, instead remaining active through- out the year (Matthiesen 1968). There is in- formation on several species of the Bothriuridae. The reproductive period of Bothriurus bonariensis (C.L. Koch 1841) is from November to February, B. flavidus Krae- pelin 1911 from November to January, and Urophonius iheringii Pocock 1893 and U. brachycentrus (Thorell 1876) from May to September (Peretti 1997). However, differenc- es in the Thestylus aurantiurus reproductive period may be related to climatic differences at the different localities. The beginning of the reproductive period of the Argentinean spe- cies of Bothriurus coincides with the begin- ning of the local warm, wet season (Peretti 1997). This also occurs in the T. aurantiurus population of the Parque Estadual da Serra da Cantareira and the Parque Estadual da Serra do Japi (Machado & Vasconcellos-Neto 2000). For these populations, the beginning of the reproductive period is September to Oc- tober. There is a great difference in activity be- tween the sexes in the Thestylus aurantiurus population of the Parque Estadual da Serra da Cantareira. Many more males were captured than females (Fig. 4), probably due to the in- crease of male activity during the reproductive period. Females wait for males close to their shelters, which thus explain this low occur- rence (Benton 2001). The collection using pitfall traps produce a different result from active collection with ul- traviolet light. Individuals standing still are also found with active collection. If ultraviolet lights were used, the expected number of cap- tured scorpions would be higher and the ex- pected number of females would be closer to the number of males (since the sex ratio of Thestylus aurantiurus is 1:1). However, sea- sonality of activity probably would not be as evident as in the collection with pitfall traps. We conclude that the population of Thestylus aurantiurus in the Parque Estadual da Serra da Cantareira possesses a reproductive season from September to November (Fig. 4). Influence of environmental factors on abundance. — Individuals of the Thestylus au- rantiurus population from the Parque Estadual da Serra da Cantareira apparently prefer plac- es at a higher elevation. This can be related to the possibility of shelters in lower areas be- ing flooded in the rainy season. The two areas with a higher number of collected scorpions (1 & 3) are located at a higher elevation and farther from water sources. On the other hand, the two areas with fewer collected scorpions (2 & 4) are located in places at a lower ele- vation. These areas are close to water sources, becoming flooded in the rainy season. According to Polis (1990b), some scorpion species seek specific environmental conditions in which to build their burrows. Namibian scorpions use several places as a shelter in- cluding simple holes in the soil and under tree barks (Lamoral 1979). Harington (1978) ver- ified that Cheloctonus jonesii Pocock 1892 (Liochelidae) examines a large area before be- ginning to dig its burrow. Additionally, in Urodacus there are differences among species in the choice of place, format and structure of burrows (Koch 1978). Many researchers verified the preference of scorpions for higher elevation localities that do not flood and are far from water sources. Williams (1966) observed that burrows of An- uroctonus phaeodactylus (Wood 1863) (luri- dae) are located on steep slopes, their entranc- es being counter to surface water flow. These burrows are rarely located on the bottom of valleys, in the lower regions of drainage. Zin- ner & Amitai (1969) verified that two species of Compsobuthus Vachon 1949 (Buthidae) of Israel migrate to higher places during the rainy season. The entrances to their burrows 220 THE JOURNAL OF ARACHNOLOGY are also built to avoid the accumulation of rain water. Koch (1977) also observed that several species of Australian scorpions build their burrows only on slopes or where rain water does not accumulate. The individuals of T. au- rantiurus studied in this work were more abundant in places at a higher elevation. These places are sloping, do not flood and rain water does not accumulate. ACKNOWLEDGMENTS We thank Sergio A. Vanin, head of Depar- tamento de Zoologia of Institute de Biocien- cias of Universidade de Sao Paulo during 2003, when this study was carried out. We thank the colectors of UNIBAN: Arlei Mar- cili, Caroline C. Aires, Cristiane Fojo, Do- menica Palomaris, Fernanda A.N. Bastes, Fer- nanda Martins, Katia Brozios, Laerte B. Viola, Marcelo Timoteo, Patricia B. Bertola, Roberta Pacheco, Sandra Favorito and Sidney F. A. dos Santos. We thank Gustavo E. Kaneto, Fla- vio H.S. dos Santos, Andre do A. Nogueira, Alexandre Albuquerque da Silva, Alexandre C. Martensen, Felipe B. de Oliveira, Rodrigo H. Willemart for help in statistical analysis and Glauco Machado for help in statistical analysis and personal communication. We thank Dalmo do V Nogueira Filho, president of SABESP, for supplying the data of rainfall and temperature. We thank Thayna J. Mello for assistance in language translation of the manuscript. We thank Glauco Machado and Alfredo V. Peretti for assistance and revision of the manuscript. We also thank the other members of LAL: Ana Lucia Tourinho, Jose Paulo L. Guadanucci, Marcio B. da Silva, Marcos R. Hara and Sabrina O. Jorge for as- sistance in completing this project. LITERATURE CITED Benton, T 2001. Reproductive ecology. Pp. 278- 301. In Scorpion Biology and Research. (P. Brownell & G.A. Polls, eds.). Oxford University Press, Corey, D.T & W.K. Taylor. 1987. Scorpion, pseu- doscorpion, and opilionid faunas in three central Florida plant communities. Florida Scientist 50(3): 162-167. Farley, R. 2001. Structure, Reproduction and De- velopment. Pp. 13-78. In Scorpion Biology and Research (P. Brownell & G.A. Polls, eds.). Ox- ford University Press. Fet, V., W.D., Sissom, G. Lowe, & M.E. Braun- walder. 2000. Catalog of the Scorpions of the World (1758-1998). The New York Entomolog- ical Society. 690 pp. Harington, A. 1978. Burrowing biology of the scor- pion Cheloctonus jonesii (Arachnida: Scorpioni- da: Scorpionidae). Journal of Arachnology 5: 243-249. Hofer, H., E. Wollscheid, & T. Gasnier. 1996. The relative abundance of Brotheas amazonicus (Chactidae, Scorpiones) in different habitat types of a Central Amazon rainforest. Journal of Ar- achnology 24:34-38. Koch, L.E. 1977. The taxonomy, geographic distri- bution and evolutionary radiation of Australo- Papuan scorpions. Records of the Western Aus- tralian Museum 5(2):83-367. Koch, L.E. 1978. A comparative study of the struc- ture, function and adaptation to different habitats of burrows in the scorpion genus Urodacus (Scorpionida, Scorpionidae). Records of the Western Australian Museum 6(2): 1 19-146. Lamoral, B.H. 1979. The scorpions of Namibia (Arachnida, Scorpionida). Annals of the Natal Museum 23(3):497-784. Machado, G. & J. Vasconcellos-Neto. 2000. Sperm transfer behavior in the neotropical scorpion Thestylus glazioui (Bertkau) (Scorpiones: Both- riuridae). Re vista de Etologia 2(l):63-66. Matthiesen, EA. 1968. On the sexual behaviour of some brazilian scorpions. Re vista Brasileira de Pesquisas Medicas e Biologicas l(2):93-96. Peretti, A.V. 1995. Analisis de la etapa inicial del cortejo de Bothriurus bonariensis (C.L. Kock) (Scorpiones, Bothriuridae) y su relacion con el reconocimiento sexual. Revue Arachnologique ll(4):35-45. Peretti, A.V. 1997. Alternativas de gestacion y producion de crias en seis escorpiones argentinos (Scorpiones: Buthidae, Bothriuridae). Iheringia, Serie Zoologia, 82:25-32. Polis, G.A. 1990a. Introduction. Pp. 1-8. In The Biology of Scorpions. (G.A. Polis, ed.). Stanford University Press. Polis, G.A. 1990b. Ecology. Pp. 247-293. In The Biology of Scorpions. (G.A. Polis, ed.). Stanford University Press. Polis, G.A. & S.J. McCormick. 1986. Patterns of resource use and age structure among a guild of desert scorpions. Journal of Animal Ecology 55: 59-73. Williams, S.C. 1966. Burrowing activies of the scorpion Anuroctonus phaeodactylus (Wood) (Scorpionida: Vejovidae). Proceedings of the California Academy of Sciences 34(8):4 19-428. Zinner, H. & P. Amitai. 1969. Observations on hi- bernation of Compsobuthus acutecarinatus Si- mon and C. schmiedeknechti Vachon (Scorpion- idea, Arachnida) in Israel. Israel Journal of Zoology 18:41-47. Manuscript received 30 June 2004, revised 26 Jan- uary 2005. 2006. The Journal of Arachnology 34:221-226 OBSERVATIONS ON LOXOSCELES RECLUSA (ARANEAE, SICARIIDAE) FEEDING ON SHORT HORNED GRASSHOPPERS Jennifer Parks: University of Missouri-Rolla, Rolla MO 65401 William V. Stoecker: 1702 East 10* Street, Rolla MO 65401-4600. E-mail: wvs@ umr.edu Charles Kristensen: RO. Box 1090, Yarnell, AZ 85362 ABSTRACT. Observations on Loxosceles reclusa Gertsch & Mulaik 1940, feeding on various species of short-homed grasshoppers are presented. In this paper, prey attack strategy, duration of feeding, and behaviors surrounding feeding are reported. The spiders routinely fed on prey larger than themselves. Lightly touching prey with palps prior to feeding was always observed. The first quick bites and the first attachment sites were mostly peripheral, with later attachment sites central, on the head, thorax or abdo- men. Feeding times, typically 3-10 hours, ranged up to 23 hours 38 minutes. The first long attachment was usually on a peripheral location of the prey (antenna or leg), but subsequent long attachments were more often central. Overall, 39.5% of long attachments were on the main body of the prey (not antenna or leg). Long attachments were then frequently followed by web spinning, or uncommonly, bradykinesia. Rocking, tugging or pulling at prey between attachments was common. The slow feeding from multiple sites on the prey appears to be an efficient strategy for this sit-and-wait predator to extract maximum nourishment from the large prey. Keywords: Brown recluse spider, feeding, arachnid behavior Loxosceles reclusa Gertsch & Mulaik 1940, a species of recluse spiders found throughout the Midwestern USA, is of considerable in- terest medically because envenomation can cause significant cutaneous necrosis and, less commonly, severe systemic manifestations in- cluding hemolytic anemia and renal failure (Anderson 1997). Greater understanding of the details of feeding behavior of this species may have medical implications and this has motivated us to study in detail the feeding ac- tivities of L. reclusa on one type of prey. We studied the attack and feeding sequence of L. reclusa on various species of short-horned grasshoppers. We recorded latency of bites, duration of feeding, bite sites and movements of the spider during the feeding sequence. METHODS Fifty-six spiders (29 females, 19 males and 8 juveniles) were selected at random from a colony of 600 individually housed L. reclusa. All were captured from houses and outbuild- ings in Phelps, Dent, and Texas Counties in south central Missouri (between latitudes 37°32' and 37°56'N, and between longitudes 91°41' and 91°58'W) and had been in captiv- ity from 10 days to over two years. Spiders were fed domestic crickets in captivity (one cricket per spider every 2-3 weeks) and, be- fore our trials, prey were withheld for inter- vals varying from 3-98 days. The average in- terval between the previous feeding and the observed feeding were similar for all three groups: males: mean — 27.25 days, females: mean — 28.08 days, and immatures mean == 30.43 days. Spiders for the study were all housed individually in glass jars (5,7 cm di- ameter X 5.7 cm height) and were left in these jars for the feeding observations. No water source was provided, and the spiders were kept under room light with window light dur- ing the day and artificial light in the evenings, but no nocturnal light. Prey for this study con- sisted of short-horned grasshoppers, captured from lawns in Phelps and Dent Counties, Mis- souri. Total body sizes for these grasshoppers ranged from 7.9-19.1 mm. Biting and feeding behaviors were observed after dropping one grasshopper into the spi- 221 222 Figure 1 . — Loxosceles reclusa palpating a grass- hopper, a behavior that was present in all observed predations {n = 56). Note that the prey length is greater than that of the spider predator. der’s cage at a distance of three to five cen- timeters from the spider. The grasshoppers were partly immobilized by severing the pos- terior legs at the femorahtibial junction. This was done to allow easier capture by the spider and easier observation, but this most likely changed the number of quick bites on the pos- terior leg. Observations were made at a dis- tance of 1 m to minimize disturbance to the spider. We recorded latency to first bites (time from introduction of prey to first bite), latency to long attachment (time from introduction of prey to long attachment) and duration of long attachments. Two types of bites were ob- served: quick bites and long attachments. We define a long attachment as an attachment last- ing more than two consecutive minutes. All shorter bites are called quick bites. We re- corded location of first bite, number of quick bites, actions before and after web spinning, and locations of all long attachments. Stu- dents’ t-test statistics for the study, assuming normal distributions with unequal variances, were calculated using the PAST online statis- tics calculator (http://folk.uio.no/ohammer/ past/). Voucher specimens were deposited in the Denver Museum of Nature and Science, Denver, Colorado. RESULTS Generally, before feeding, the spider lightly touched the prey with both palps (Fig. 1) prior to delivering a first bite. This behavior oc- curred in all observed predations. The latency to the initial bite averaged 5.58 ± 9.86 min after introduction of the prey, with differences between groups not significant except for fe- THE JOURNAL OF ARACHNOLOGY !' I males vs. immatures (males vs. females, t = i 1.23, P = 0.23; males vs. immatures, t = 2.13, j P = 0.05; females vs. immatures, t = 2.14, P i' - 0.04). I Generally, L. reclusa delivered one or two j quick bites before a long attachment, but the | total number of quick bites ranged from zero j to ten (Fig. 2). Within two minutes of the first | bite, the prey ceased almost all movement and i the spider thee began a long attachment (Table | 1) . The latency to the first long attachment for females was significantly longer than the cor- responding times for juveniles {t = 2.548, P — 0.016) but not different for males {t ” ”1.650, P = 0.107). Feeding duration was longest for females (Table 1, females com- | pared with all other spiders, males and im- matures combined, t = ”3.784, P = 0.002). The quick bite sequence was extremely rap- id, with the spider darting in, biting, and jumping back, normally within a fraction of a second. Data pooled from all three spider groups show that this sequence involved one or two bites in 58% of cases (Fig. 2). These bites are delivered to easily accessible periph- eral parts of the prey, either legs or antennae (Figs. 3 & 4), allowing L. reclusa to rapidly deliver enough venom to paralyze the prey. After the first quick bite, the spider retreated quickly, as noted by Canrel (pers. comm.). Both the peripheral attack strategy and the quickness of biting and withdrawal observed in the earliest bites were observed consistent- ly. The quick bite sequence was usually fol- lowed by a retreat and wait (holding) stage that averaged 15.55 ± 22.63 minutes. The longest holding stage noted for the 56 spiders observed was 139 minutes. For some spiders, the first bite was a “long attachment” (Fig. 2) . During feeding, spinning of silk was ob- served generally before a long attachment or after a long attachment (Fig. 5). Spinning was also seen when the spider was introduced to a new jar. Web production seemed to be used to immobilize the prey for a possible addi- tional feeding, seen in 48% of the fifty-six ob- served predations. In 23.5% of cases, web spinning terminated the feeding sequence. A variation seen in one instance in this series and in one other observed instance is a slow stepping around the prey, a distinct pattern of bradykinesia confined to feeding. Frequently, the spiders walked around the jar spinning PARKS ET AI..—LOXOSCELES FEEDING 223 40 n 0123456789 10 # of bites Figure 2.— The number of quick bites inflicted by L. reclusa on a grasshopper is typically one or two, but ranged from zero to ten. Within two minutes of these quick bites, the prey ceased almost all movement. variable amounts of web after they had com- pletely finished feeding on the grasshopper. It was noted that all successful feedings (those feedings in which the spider had at least one attachment longer than two minutes) lasted at least three hours. The longest feeding reported here lasted over twenty-three hours. One other feeding lasted over 47 hours, though this was not included in this series due to the inability to observe the complete feed- ing. Disruptions were observed twice during feeding, and in each case the spiders resumed feeding after a brief pause, once at the pre- vious feeding site, and the other at a different site. DISCUSSION The preferred initial L. reclusa feeding sites of legs and antenna (Fig. 4) confirm those ob- served by Hite (1966), who noted “when feeding on grasshoppers up to 35 mm in length, the most commonly selected part is a leg or an antenna.” The preferred feeding sites cannot be generalized to all prey, as the spider appears to adapt its feeding sequence to prey morphology. As Hite noted for 1383 feedings of house flies to L. reclusa, head, abdomen, and thorax accounted for 39%, 26%, and 15% of feeding sites respectively, with legs ac- counting for only 20% of feeding sites (Hite 1966). Table 1 . — Mean latency of first quick bite and first long attachment and feeding duration, with standard deviations, for L. reclusa feeding on short homed grasshoppers. Spider Mean latency of first quick bite Mean latency of 1st long attachment Duration of feeding Immature (« = 8) Female {n = 29) Male {n = 19) 2,14 ± 1.95 min 8.00 ± 13.72 min 4.59 ± 3.64 min 15.63 ± 16.59 min 41.93 ± 45.77 min 26.00 ± 19.99 min 690.25 ± 190.76 min 737.62 ± 318.39 min 391.63 ± 178.58 min Percent occurrence 224 THE JOURNAL OF ARACHNOLOGY Figure 3. — Loxosceles reclusa feeding on grass- hopper antenna. The antenna is chosen as a site for the first long attachment in 32.1% of feedings, but is chosen as a long attachment site in only 16.1% of cases overall. The ability of L. reclusa to survive in cap- tivity for long periods of time without prey is well known and the spiders are frequently found in areas where prey is only available sporadically. Here it was observed that L. re- clusa ean take larger and potentially hazard- ous prey that exceed the weight of the pred- ator. The behavior of L. reclusa indoors in a con- fined environment could differ from L. reclusa behavior in its natural environment. Green- stone (1999) noted that starvation, generally undertaken to increase the likelihood of feed- ing, might alter metabolic rates and therefore affect feeding behavior. This argument may be less valid for L. reclusa than for some other species. Hite (1966) noted that L. reclusa feedings appear to be less frequent than other species. Hite also observed a mature female L. reclusa surviving 297 days, nearly ten months, without food or water. We observed ■ long attachment □ initial bites 0 first long feeding 45 1 antenna anterior leg posterior leg main body Location Figure 4. — Location of quick bites and long attachments of L. reclusa on prey. PARKS ET h\^.—LOXOSCELES FEEDING 225 ■ preceding web production □ following web production 60 n Action Figure 5. — Actions preceding and following web production. Long attachments are the most common actions preceding and following web production. a mature L. reclusa female similarly survive 545 days. The intervals since the last feeding, 3-98 days, were well within the feeding in- tervals that the spider may encounter in its typical confined indoor habitat. As L. reclusa appears to prefer confined areas, our indoor experimental environment may not differ sig- nificantly from the situation the spider has been in when it bites humans. The behavior of the spider under these conditions is there- fore of medical importance but it should be noted that most bites of humans are made un- der different circumstances than the predatory bites studied here. Human encroachment on L. reclusa terri- tory may have changed the natural environ- ment for this spider and it may now be true that a significant portion of all L. reclusa live indoors. Hite (1966) found 430 of 626 spiders collected in indoor locations and 196 spiders in outdoor locations. Vetter & Barger (2002) trapped 2,055 L. reclusa in a single home in Kansas. However, we do not know what nat- ural conditions outdoors allow high densities of L. reclusa. In an ongoing study, high den- sities of L. deserta in packrat dees are being investigated. One potential problem with methodology is the variation in rime since the last feeding, ranging from 3-“9S days. This variation could influence spider behavior. We could find no systematic difference in behavior as a result of time since last feeding. Figure 6 shows total duration of feeding vs. time since last feeding, which appear only modestly correlated. For the 16 feedings with duration 12 days or less, the feeding duration is 598.5 ± 269.04 min- utes, vs 621.75 ± 279.62 minutes for the 16 226 THE JOURNAL OF ARACHNOLOGY Figure 6. — Time elapsed since the last feeding vs. duration of feeding. A best-fit line shows a slow increase in feeding time with longer intervals between feedings. A minimum feeding time of about three hours was seen. feedings with duration 1 3 days or more, a dif- ference that is not significant (t = ”0.24, P = 0.81). In summary, L. reclusa feeding begins with quick bites and is followed with successively more central attachments. Females tended to take longer with all phases of feeding. The use of quick bites allows L. reclusa to take large and potentially dangerous prey. Attributes of these spiders such as the mechanics of the legs, joints, and fangs appear to be adapted for swinging in and out quickly, without phys- ically overpowering prey. The venom appears to be effective when injected at peripheral sites. The long duration of feedings for L. re- clusa that we report here, frequently exceed- ing 10 hours, are incompatible with a frequent feeding regimen. The ability to utilize large prey efficiently may be very important to the spiders in times of low prey abundance. Fur- ther explorations of the adaptive value of lon- ger feeding times, the utilization of large meals and mode of action of the venom in insects is needed. LITERATURE CITED Anderson, P.C. 1997. Spider bites in the United States. Dermatologic Clinics 15:307--311. Greenstone, M.H. 1999. Spider predation: how and why we study it. Journal of Arachnology 27: 333-342. Hite, J.M. 1966. The biology of the brown recluse spider, Loxosceles reclusa, Gertsch and Mulaik, Ph.D. thesis, Kansas State University, Manhat- tan. Vetter, R.S. & D.K. Barger. 2002. An infestation of 2,055 brown recluse spiders (Araneae: Sicari- idae) and no envenomations in a Kansas home: implications for bite diagnoses in nonendemic ar- eas. Journal of Medical Entomology 39(6):948~ 51. Manuscript received 15 May 2004, revised 30 March 2006. 2006. The Journal of Arachnology 34:227-230 THE SYSTEMATIC POSITION OF THE AMAZONIAN SPECIES OF ALBIOMIX (PSEUDOSCORPIONES, IDEORONCIDAE) Mark S. Harvey: Department of Terrestrial Invertebrates, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia. E-mail: mark.harvey@museum.wa.gov.au Volker Mahnert: Museum d’histoire naturelle, Case poste 6434, CH-1211 Geneve 6, Switzerland. ABSTRACT. The new genus Xorilbia (Pseudoscorpiones, Ideoroncidae) is established for three species from the Amazon region previously included in the genus Albiorix: the type species X. arboricola (Mah- nert), X. gracilis (Mahnert) and X lamellifer (Mahnert). The new genus bears a peculiar structure on the arolium that is only found in a few other genera of Ideoroncidae. New locality records are presented for X. gracilis and X. lamellifer, including the first record of X. gracilis from Venezuela. Keywords: Taxonomy, new genus. Neotropical, Brazil, Venezuela, Xorilbia The South Americae ideoroecid fauna con- sists of 14 species in two genera. The genus Ideoroncus Balzan 1887 contains nine species from Brazil and Paraguay (Mahnert 1984, 2001), while five species attributed to the ge- nus Albiorix Chamberlin 1930 are found in Brazil, Argentina and Chile (Mahnert 1979, 1984, 1985b). While examining the status of the various species attributed to the genus AX biorix, we have found that the three species from Amazonian Brazil attributed to the genus Albiorix differ considerably from other mem- bers of the genus, particularly in features of the arolium - the elongate, soft structure sit- uated at the distal end of the pedal tarsus be- tween the tarsal claws of all pseudoscorpions (Chamberlin 1931). In particular, the three Amazonian species of Albiorix possess a small hooked structure on the ventral surface of the arolium, first observed by Vachon (1958) for two species of Negroroncus Beier 1931, a feature that is lacking in species of Albiorix. Also, the arolium is much longer than the claws and deeply divided in all spe- cies of Albiorix, whereas the arolium is slight- ly shorter than the claws and at most only slightly divided in the Amazonian species. These features suggested to us that the Ama- zonian species are misplaced in Albiorix, and we here transfer them to a new genus Xorilbia which is described and compared with similar ideoroecid genera. The specimens mentioned in this paper are lodged in the following institutions: California Academy of Sciences, San Francisco (CAS); and the University of California, Davis, Cal- ifornia, U.S.A. (UCD). Comparative material of other ideoroecid taxa examined for this study is lodged in the Museum d’histoire na- turelle, Geneve, Switzerland (MHNG); and the Western Australian Museum, Perth, West- ern Australia (WAM). Morphological termi- nology mostly follows Chamberlin (1931) and Harvey (1992). The specimens were studied using one of two techniques. Temporary slide mounts were prepared by immersion of specimens in con- centrated lactic acid at room, temperature for several days, and mounting them on micro- scope slides with 10 or 12 mm coverslips sup- ported by small sections of 0.25, 0.35 or 0.50 mm diameter nylon fishing line. After study, the specimens were returned to 75% ethanol. Permanent slide mounts were prepared by re- moving the pedipalps, the chelicera, left leg I and left leg IV from specimens with the use of eye-scissors or small needles, and clearing overnight with 10% potassium hydroxide at room temperature. The specimens were then washed in several rinses of water and 5% ace- tic acid (to neutralize the potassium hydrox- ide), and dehydrated through a graded ethanol series. They were then transferred to Euparal essence overnight at room temperature, prior to mounting in Euparal on microscope slides using 10 or 12 mm coverslips supported by small sections of 0.25, 0.35 or 0.50 mm di- ameter nylon fishing line. All specimens were 227 228 THE JOURNAL OF ARACHNOLOGY studied using an Olympus BH-2 compound microscope and illustrated with the aid of a drawing tube. Measurements were taken at the highest possible magnification using an ocular graticule. After study the specimens were re- turned to 75% ethanol with the dissected por- tions placed in 12 X 3 mm glass genitalia mi- crovials (BioQuip Products, Inc.). Family Ideoroncidae Chamberlin 1930 Genus Xorilbia NEW GENUS Type species. — Ideoroncus arboricola Mahnert 1979. Etymology. — The generic name is an ana- gram of Albiorix and is to be treated as fem- inine. Diagnosis. — Members of Xorilbia possess a small hooked process on the ventral surface of each arolium (Fig. 1), a feature shared with Dhanus siamensis (With 1906) and with all species of Typhloroncus Muchmore 1979, Ne- groroncus Beier 1931 and Afroroncus Mah- nert 1981. It differs from these genera as fol- lows: Xorilbia and Typhloroncus have a chelal trichobothrial pattern of 22 on the fixed finger and 10, occasionally 1 1, on the movable finger (22/10 or 22/11), whereas Dhanus siamensis, Negroroncus species and Afroroncus species have a pattern of 20/10, with the exception of N. jeanneli Vachon 1958, which has a pattern of 26/12. Xorilbia has chelal teeth that are widely spaced whereas Typhloroncus has closely-spaced chelal teeth. Xorilbia has 5 tri- chobothria in the ib region and 6 trichobothria in the ist region of the fixed chelal finger, whereas species of Typhloroncus bear 4 tri- chobothria in the ib region and 7 trichobothria in the ist region. Xorilbia also differs from Typhloroncus in the presence of one pair of eyes; whereas eyes are totally absent in all species of Typhloroncus. Description. — All setae long, virtually straight and acicular. Most cuticular surfaces smooth and glossy. Pedipalps: long and slender. Fixed chelal finger with 22 trichobothria, movable chelal finger with 10 trichobothria: eb region with 1 trichobothrium; est region with 6 trichoboth- ria; ib region with 5 trichobothria; ist region with 6 trichobothria; b region with 2 tricho- bothria; and t region with 6 trichobothria; st not ventrally displaced. Venom apparatus present in both chelal fingers, venom duct ter- Fig. 1. — Xorilbia lamellifer (Mahnert): detail of tip of left tarsus IV, female from Fazenda Esteio, Brazil. The arrow indicates the ventral hooked pro- cess on the arolium. minating in nodus ramosus near est region in fixed finger and near t region in movable fin- ger. Chelal teeth widely spaced. Condyle on the chelal hand small and rounded. Chelicera: with 6 long, acuminate setae on hand; movable finger with 1 long subdistal seta; flagellum of 4 thickened blades, all blades serrate; lamina exterior absent; galea long and slender. Cephalothorax: carapace with 2 small, bulging eyes; without furrows; anterior mar- gin with 4 setae. Manducatory process with 2 long distal setae. Abdomen: tergites and sternites undivided. Pleural membrane longitudinally striate. Each stigmatic sclerite with 1 seta. Posterior max- illary lyrifissure present and sub-basally situ- ated. Spiracles simple, with spiracular helix. HARVEY & MAHNERT— AMAZONIAN SPECIES OF ALBIORIX 229 Legs: femur I and II without basal swelling; femora I and II with primary slit sensillum directed transversely; femur I much longer than patella I; suture line between femur IV and patella IV transverse; metatarsus shorter than tarsus; metatarsal pseudotactile seta sub- proximal; legs with two subterminal tarsal se- tae, each acuminate; arolium shorter than claws, slightly divided, with ventral hooked process; claws slender and simple. Remarks. — The hooked process on the ventral surface of the arolium of all legs (Fig. 1) was first noted and illustrated in two spe- cies of Negroroncus (Vachon 1958), and we have found that it is present in Dhanus sia- mensis and in species of Typhloroncus, Ne- groroncus, Afroroncus and Xorilbia. It would seem likely that the hooked process is an apo- morphic feature that defines this group as a monophyletic entity, but the close relationship between Afroroncus, which bears a ventral hook, and Nannoroncus Beier 1955 (Mahnert 1981), which lacks a hook, may negate the power of this feature to define a clade. The process is absent from species of Ideoroncus, Dhanus Chamberlin 1930 (excluding D. sia- mensis), Shravana Chamberlin 1930, Nha- trangia Redikorzev 1938, Nannoroncus, Al- biorix and Pseudalbiorix Harvey, Barba, Muchmore & Perez in press. Dhanus siamen- sis bears very little resemblance to the re- maining species of Dhanus, including the type species D. sumatranus, and will be placed in a new genus as part of a forthcoming review of the Asian members of the Ideoroncidae (Harvey unpub. data). The ideoroncids with a ventral hooked pro- cess on the arolium are widely distributed around the world with Xorilbia occurring in the Amazon basin in northern Brazil and southern Venezuela, Typhloroncus species from the West Indies and Mexico (Muchmore 1979, 1982, 1986), Negroroncus and Afroroncus from east- ern Africa (Mahnert 1981), and D. siamensis from south-east Asia (Schawaller 1994). The removal of the Amazonian species from Albiorix, and the recent transfer of two species of Albiorix to a separate genus (Har- vey et al. in press) reduces Albiorix to 1 1 spe- cies ranging from western North America to Mexico [A. anophthalmus Muchmore 1999, A. edentatus Chamberlin 1930, A. bolivari Beier 1963, A. conodentatus Hoff 1945, A. magnus Hoff 1945, A. mexicanus (Banks 1898), A. mirabilis Muchmore 1982, A, parvidentatus Chamberlin 1930, A. retrodentatus Hoff 1945], with isolated species in Argentina [A. argentiniensis (Hoff 1954)] and Chile [A. chi- lensis (Ellingsen 1905)]. Distribution. — Species of Xorilbia occur in the northern Brazilian states of Amazonas and Para, and in southern Venezuela.* Xorilbia arboricola (Mahnert 1979) NEW COMBINATION Ideoroncus arboricola Mahnert 1979:753-755, figs. 70-74; Adis et al. 1987:488. Albiorix arboricola (Mahnert): Mahnert 1984:672- 673; Mahnert 1985a:78; Mahnert & Adis 1986: 213; Mahnert et al. 1986: fig. 10; Harvey 1991: 316; Adis & Mahnert 1993: fig. 5; Mahnert & Adis 2002:379, fig. 10; Adis et al. 2002:5. Diagnosis. — Xorilbia arboricola lacks the lamelliform ridge on the fixed chelal finger that is characteristic of X. lamellifer, and the pedipalpal segments are more robust than in X. gracilis. Description. — See Mahnert (1979, 1984). Remarks. — Xorilbia arboricola occurs at several locations in Amazonas and Para where it is occasionally sympatric with X. gracilis (Mahnert 1984). Xorilbia gracilis (Mahnert 1985) NEW COMBINATION Albiorix aff. arboricola (Mahnert): Mahnert 1984: 673. Albiorix gracilis Mahnert 1985b:223-224, figs. 27- 28; Mahnert & Adis 1986:213; Harvey 1991:317. Albiorix gracilis Mahnert: Adis & Mahnert 1990: 13, figs. 2-3; Adis & Mahnert 1993:435, figs. 2- 3, 5; Mahnert & Adis 2002:379; Adis et al. 2002:5. New material examined. — VENEZUELA: Amazonas: 1 S, I 9, Alto Rio Siapa, 1°40'N, 64°35'W, 650 m, 4 February 1989, sifting leaf litter in rainforest, J. Lattke (CAS). Diagnosis.” gracilis lacks the la- melliform ridge on the fixed chelal finger that is characteristic of X. lamellifer, and the pe- dipalpal segments are more slender than in X, arboricola. Description. — See Mahnert (1985b). Remarks.”Aon7^/<2 gracilis was recorded by Mahnert (1985b) from two locations in Amazonas where it is sympatric with X. ar- boricola. The new records listed here are of two further specimens from southern Vene- zuela that generally fit the original description, although they are slightly larger than the type specimens; e.g., chela (with pedicel) length. 230 THE JOURNAL OF ARACHNOLOGY male 1.067/0.245 (= 4.36 times longer than broad) and female 1.200/0.324 (= 3.70 times longer than broad). Xorilbia lamellifer (Mahnert 1985) NEW COMBINATION Fig. 1 Albiorix lamellifer Mahnert 1985b:225— 226, figs. 29-31; Mahnert & Adis 1986:213; Harvey 1991: 317; Mahnert & Adis 2002:379. New material examined. — BRAZIL: Ama- zonas: 1 9, Fazenda Esteio, 80 km NNE. of Manaus, 2°25'S, 59°46'W, 80 m, 15 Septem- ber 1987 (UCD). Diagnosis. — The basal teeth on the fixed chelal finger of X. lamellifer are modified into a lamelliform ridge, which distinguishes this species from X. arboricola and X. gracilis. Description. — See Mahnert (1985b). Remarks. — Mahnert (1985b) described this species based upon a single female col- lected 25 km NE. of Manaus. We have ex- amined a second female from a farm situated 80 km NNE. of Manaus. ACKNOWLEDGMENTS We are grateful to Charles Griswold and Darrell Ubick (CAS) and Steve Heydon (UCD) for access to the pseudoscorpion col- lections lodged in their institutions, and to two anonymous reviewers and the editors Gail Stratton, Paula Cushing and Dan Mott for their comments on the manuscript. LITERATURE CITED Adis, J., A.B. Bonaldo, A.D. Brescovit, R. Bertani, J.C. Cokendolpher, B. Conde, A.B. Kury, W.R. Lourengo, V, Mahnert, R. Pinto-da-Rocha, N.L Platnick, J.R. Reddell, C.A. Rheims, L.S. Rocha, J.M. Rowland, P. Weygoldt & S. Woas. 2002. Arachnida at ‘Reserva Ducke’, central Amazon- ia/Brazil. Amazoniana 17:1-14. Adis, J., W.J. Junk & N.D. Penny. 1987. Material zoologico depositado nas colegoes sistematicas de entomologia do INPA, resultante do “Projeto INPA/Max-Planck”. Acta Amazonica 15:481- 504. Adis, J. & V. Mahnert. 1990. Vertical distribution and abundance of pseudoscorpion species (Arachnida) in the soil of a neotropical second- ary forest during the dry and the rainy season. Acta Zoologica Fennica 190:11-16. Adis, J. & V. Mahnert. 1993. Vertical distribution and abundance of pseudoscorpions (Arachnida) in the soil of two different neotropical primary forests during the dry and rainy seasons. Mem- oirs of the Queensland Museum 33:431-440. Chamberlin, J.C. 1931. The arachnid order Chelo- nethida. Stanford University Publications, Bio- logical Sciences 7(1): 1-284. Harvey, M.S. 1991. Catalogue of the Pseudoscor- pionida. Manchester University Press, Manchester. Harvey, M.S. 1992. The phylogeny and systematics of the Pseudoscorpionida (Chelicerata: Arachni- da). Invertebrate Taxonomy 6:1373-1435. Mahnert, V. 1979. Pseudoskorpione (Arachnida) aus dem Amazonas-Gebiet (Brasilien). Revue Suisse de Zoologie 86:719-810. Mahnert, V. 1981. Die Pseudoskorpione (Arachni- da) Kenyas. 1. Neobisiidae und Ideoroncidae. Re- vue Suisse de Zoologie 88:535-559. Mahnert, V. 1984. Beitrag zu einer besseren Kennt- nis der Ideoroncidae (Arachnida: Pseudoscorpi- ones), mit Beschreibung von sechs neuen Arten. Revue Suisse de Zoologie 91:651-686. Mahnert, V. 1985a. Pseudoscorpions (Arachnida) from the lower Amazon region. Revista Brasi- leira de Entomologia 29:75-80. Mahnert, V. 1985b. Weitere Pseudoskorpione (Arachnida) aus dem zentralen Amazonasgebiet (Brasilien). Amazoniana 9:215-241. Mahnert, V. 2001. Cave-dwelling pseudoscorpions (Arachnida, Pseudoscorpiones) from Brazil. Re- vue Suisse de Zoologie 108:95-148. Mahnert, V. & J. Adis. 1986. On the occurrence and habitat of Pseudoscorpiones (Arachnida) from Amazonian forest of Brazil. Studies on Neotrop- ical Fauna and Environment 20:211-215. Mahnert, V. & J. Adis. 2002. Pseudoscorpiones. Pp. 367-380. In Amazonian Arachnida and Myria- poda. (J. Adis, ed.). Pensoft Publishers, Sofia. Mahnert, V., J. Adis & PE Buhrnheim. 1986. Key to the families of Amazonian Pseudoscorpiones (Arachnida). Amazoniana 10:21-40. Muchmore, W.B. 1979. Pseudoscorpions from Flor- ida and the Caribbean area. 9. Typhloroncus, a new genus from the Virgin Islands (Ideoronci- dae). Florida Entomologist 62:317-320. Muchmore, W.B. 1982. Some new species of pseu- doscorpions from caves in Mexico (Arachnida, Pseudoscorpionida). Bulletin for the Association of Mexican Cave Studies 8:63-78. Muchmore, W.B. 1986. Additional pseudoscorpi- ons, mostly from caves, in Mexico and Texas (Arachnida: Pseudoscorpionida). Texas Memori- al Museum, Speleological Monographs 1:17-30. Schawaller, W. 1994. Pseudoskorpione aus Thailand (Arachnida: Pseudoscorpiones). Revue Suisse de Zoologie 101:725-759. Vachon, M. 1958. Sur deux Pseudoscorpions nou- veaux des cavemes de I’Afrique equatoriale [Ideo- roncidae]. Notes Biospeologiques 13:57-66. Manuscript received 15 March 2005, revised 29 July 2005. 2005. The Journal of Arachnology 34:231-233 SHORT COMMUNICATION ADDITIONAL NOTES ON THE POST»BIRTH DEVELOPMENT OF THE SCORPION VAEJOVIS COAHUILAE WILLIAMS (VAEJOVIDAE) W. David Sissom: Dept, of Life, Earth, & Environmental Sciences, West Texas A&M University, WTAMU Box 60808, Canyon TX 79016 Kari J* McWest: 16 Thunderbird Dr., Canyon, TX 79015 Aene L. Wheeler: 2417 Capehart Drive, Richmond, VA 23294 ABSTRACT. Fourteen specimens of Vaejovis coahuilae Williams 1968 were bom in the laboratory and reared in an incubator at a near-constant 27 °C. A single female reached maturity at the 8* instar, as previously hypothesized, but unverified, in this species. Two other specimens reached the 7* instar, but the male died at the molt and the female was not yet mature. Data on the duration and morphometries of observed instars are provided for all specimens. Keywords: Life history, instars, Vaejovidae, Vaejovis A study of the life history of Vaejovis coahuilae Williams 1968 was published earlier by Fraocke & Sissom (1984). In that study, three individuals were successfully reared to the sixth instar, with only one (a male) reaching adulthood. Based on comparisons with field-collected adults, the authors used an ex- trapolation method and a formula-based estimate to hypothesize that males of the species mature at ei- ther the sixth or seventh instar and females at either the seventh or eighth. The purpose here is to report new data that validates (at least in part) the hypoth- esis of Francke & Sissom (1984). Vaejovis coahuilae is a very common scorpion found in southeastern Arizona, much of New Mex- ico, western Texas and northern Mexico (Sissom 2000; unpub. data). In the current study, a pregnant female specimen of V. coahuilae was collected along Lea County Rd. 21, 15 mi S jet. with NM 128, New Mexico on 24 May 1990 and returned to the laboratory. At the beginning of August, the fe- male was discovered with 29 first instar offspring on her back. The molt to the second instar occurred on 5 August 1990, and the young descended from the mother's back within the next two days. After their descent, 14 offspring were transferred to in- dividual plastic widemouth specimen jars (height, 4.45 cm; inside diameter, 4,76 cm). The specimens in their jars were placed in a tabletop incubator at a near constant temperature of 27 °C. There was no internal lighting in the incubator and the specimens kept there were in near constant darkness, experi- encing light only during brief periods of mainte- nance. All specimens were fed and watered on the same schedule. Food provided included wingless Dro- sophila melanogaster Meigen, field-collected ter- mites (Reticulutermes Holmgren sp.), and small mealworm larvae (Tenebrio molitor L.), with pro- gressively larger prey being given to later instar specimens. Two circular pieces of paper towel were placed in the bottom of each container to provide shelter and water for the scorpions. Distilled water was provided at least twice weekly by pipetting sev- eral drops directly onto the paper towel; the scor- pions were frequently observed drinking from the moistened substrate. Care was taken not to over- saturate this substrate to prevent drov/ning of the young scorpions, and the paper towel was changed periodically. The lids of the jars were loosely re- placed so as to allow the humidity inside the jars to subside to normal levels within a day or two. Twelve of the individuals successfully completed the second instar, but one died during the molt. Only half of those (n = 6) survived the third instar, one more died during the fourth instar and two more died during the fifth. Two females and one male reached the seventh instar (the male dying during the molt) and, of the two females remaining, one successfully reached the eighth. The female that later died in the seventh instar was dissected and no development of the ovariuteras was observed; the eighth instar female, however, was clearly adult. 231 232 THE JOURNAL OF ARACHNOLOGY Table 1. — Growth data by instar for Vaejovis coahuilae Williams reared in an incubator at 27°C. Means, minima (min), and maxima (max) are given for raw measurements (in mm). Average growth factors (GF) between instars are also provided. Measurements included carapace length (CarL), pedipalp chela length (ChelaL), pedipalp chela width (ChelaW), metasomal segment V length and width (MetVL, MetVW), and metasomal segment IV length and width (MetIVL, MetIVW). Growth factors for the single specimen reaching the eight instar were calculated from its 7th instar sizes. Three specimens actually reached the seventh instar. The male died during the molt, and as a result, its cuticle was distorted and probably not expanded. Instar CarL ChelaL ChelaW MetVL MetVW MetIVL MetIVW 2nd {n = 12) mean 1.33 1.59 0.36 1.21 0.55 0.78 0.55 min 1.23 1.50 0.33 1.12 0.47 0.70 0.50 max 1.44 1.72 0.40 1.30 0.60 0.90 0.60 GF to 3rd 1.24 1.25 1.28 1.28 1.33 1.33 1.29 3rd {n = 6) mean 1.66 2.00 0.46 1.55 0.73 1.05 0.71 min 1.55 1.85 0.40 1.45 0.60 0.95 0.62 max 1.86 2.35 0.55 1.80 0.85 1.23 0.82 GF to 4th 1.12 1.13 1.15 1.21 1.15 1.17 1.18 4th {n - 5) mean 1.86 2.26 0.53 1.88 0.84 1.23 0.84 min 1.51 1.85 0.40 1.65 0.62 0.97 0.64 max 2.08 2.51 0.60 2.00 0.97 1.35 0.95 GF to 5 th 1.27 1.31 1.36 1.28 1.35 1.46 1.35 5th {n = 3) mean 2.37 2.95 0.72 2.41 1.13 1.79 1.13 min 2.10 2.70 0.60 2.12 0.95 1.66 0.98 max 2.60 3.20 0.82 2.70 1.30 1.95 1.25 GF to 6th 1.25 1.26 1.29 1.28 1.26 1.17 1.24 6th {n = 3) mean 2.99 3.76 0.95 3.16 1.46 2.12 1.45 min 2.75 3.50 0.85 2.89 1.32 1.97 1.31 max 3.15 3.92 1.00 3.30 1.55 2.20 1.55 GF to 7th 1.18 1.21 1.29 1.20 1.25 1.21 1.26 7th {n = 2) min 3.42 4.67 1.25 3.80 1.81 2.60 1.79 max 3.94 4.97 1.35 4.05 1.95 2.80 1.95 GF to 8th 1.35 1.36 1.55 1.36 1.37 1.31 1.35 8th {n = 1) 4.60 6.35 1.94 5.17 2.48 3.40 2.42 That adulthood in this female was reached in eight instars corroborates the earlier hypothesis by Francke & Sissom (1984) based on the theoretical and indirect methods. Whether or not individuals can mature at the seventh instar as well was not answered in the current study, but it seems likely. In natural populations of V. coahuilae, there are “small” and “large” adult males and females that could belong to different instars. Average instar duration for the incubator-reared litter is as follows: 2nd instar, 150.8 days (110.7 days, if two outliers are excluded; these individuals required 292 and 4 1 1 days to reach the 3rd instar. respectively); 3rd instar, 64.2 days; 4th instar, 95.8 days; 5th instar, 119 days; 6th instar, 422 days; and 7th instar, 835.5 days. It should be noted that in the late summer of 1992, the specimens were trans- ported from North Carolina to Texas and placed in a similar tabletop incubator soon after arrival. Per- haps this disturbance resulted in prolonged late in- stars for the three surviving specimens. Morphometric data, including measurements and growth factors for each molt are provided in Table 1. Measurements of carapace length, chela length and width, metasomal segment IV length and width, and metasomal segment V length and width were SISSOM ET AL.— SCORPION POST-BIRTH DEVELOPMENT 233 taken as shown in Sissom et al. (1990, fig. 11.1). Growth factors were calculated by dividing the di- mension of a particular structure at a given instar by its dimension at the previous instar. Growth fac- tors were similar to those reported by Francke & Sissom (1984). We wish to thank Dr. Oscar F. Francke and two anonymous reviewers for their comments on the manuscript. The specimens and exuviae are depos- ited in the entomology collection of the Department of Life, Earth, & Environmental Sciences, West Texas A&M University. LITERATURE CITED Francke, O.F. & W.D. Sissom. 1984. Comparative review of the methods used to determine the number of molts to maturity in scorpions (Arach- nida), with an analysis of the post-birth devel- opment of Vaejovis coahuilae Williams (Vaejov- idae). Journal of Arachnology 12:1-20, Sissom, W.D,, G.A. Polis & D.D. Watt. 1990. Chap- ter 11: Laboratory and field methods. Pp. 445- 461. In G. A. Polis (Ed.), The Biology of Scor- pions. Stanford University Press, Stanford, CA., 587 pp. Sissom, W.D. 2000. Family Vaejovidae. Pp. 503- 553. In Fet, V., W D. Sissom, G. Lowe, and M. Braunwalder. Catalog of the scorpions of the world (1758-1998). New York Entomological Society, 690 pp. Manuscript received 24 February 2003, revised 8 November 2004. 2006. The Journal of Arachnology 34:234--240 SHORT COMMUNICATION VARIATIONS IN WEB CONSTRUCTION IN LEUCAUGE VENUSTA (ARANEAE, TETRAGNATHIDAE) Yaee Henaut: Laboratorio de Ecologfa y Conservacion de la Fauna, E.C.O.S.U.R, Apdo, Postal 424, 77900 Chetumal, Quintana Roo, Mexico Jose Alvaro Garcfa-Ballinas: Laboratorio de Ecoetologia de los Artropodos, E.C.O.S.U.R., Apdo. Postal 36, 30700 Tapachula, Chiapas, Mexico Claude Alauzet: Laboratoire d'Ecologie Terrestre, Universite Paul Sabatier, 31062 Toulouse, France ABSTRACT. The distribution of female Leucauge venusta (Walckenaer 1841) in a coffee plantation in southern Mexico was studied in order to determine the vertical distribution of this spider. Principal com- ponent analysis clearly identified the presence of three distinct groups of L. venusta webs, based on the number of spirals/web and principally on the height at which the webs were located; most L. venusta webs (63/100) were close to the ground. Spiders on high webs (153.8 ± 3.6 cm above ground, mean ± S.E.) were significantly larger and heavier than spiders on lower webs. Large spiders had significantly larger, better developed ovaries, than smaller conspecifics, presumably indicative of sexual maturity. Sig- nificantly more insects were captured by sticky traps placed at 50 cm height than in the traps placed at 150 cm height; the most numerous captures were Diptera. However, insects caught at 150 cm were signifi.cantly larger than those caught at 50 cm above ground. We concluded that as sexual development proceeds, the spider increases the height at which the web is constructed. This vertical migration is associated with changes in web construction and the type of prey captured. These results are discussed in terms of ietraspecific competition, predation risks and sexual selection. Keywords: Web-building, prey, predation strategy, web location During foraging, a predator has to make several choices: where to eat, how much time to dedicate to eating, and what type of prey to select for capture and consumption. These choices depend on strate- gies allowing the predator to optimize its behavioral efficiency and to reduce energetic costs and time (Alcock 1993). Orb-weaving spiders are generalist predators that do not usually compete with each other for food (Wise 1993). This absence of competition may be a result of different species using different preda- tory strategies. Olive (1980) asserted that the func- tional morphology of the predator may directly af- fect the type of insect predated, as a means to avoid catching the same prey. One such strategy consists of reducing competition by building webs that dif- fer in structure and location. In this way, each spi- der species produces one type of web and employs it in one type of micro-habitat (Henaut et al. 2001). Alternatively, the web can act as a general filter and trap a large diversity of insects, and if the web is constructed at different sites, it can capture different kinds of prey. For example, Gasteracantha cancri- formis (Linnaeus 1785) builds a web in open areas whereas Cyclosa caroli (Hentz 1850) traps insects in the same type of habitat but only in enclosed areas (Ibarra-Nunez et al. 2001). In this case, selec- tion of different prey items is achieved by a com- bination of differences in web location and spider behavior (Henaut et al. 2001). These studies indi- cate that each spider species may adopt a particular strategy based on a combination of web character- istics, web location and spider behavior. Leucauge venusta (Walckenaer 1841) is a com- mon species in many habitats from the United States to South America (Levi 1981). In the field, Leucauge species generally spin inclined orb webs. The first web of the day is usually built before dawn and may be repaired or replaced during the day (Eberhard 1988). The usual sequence of orb-build- ing is to make radii and frame lines, then hub loops followed by a temporary spiral, and finally the sticky spiral. The temporary spiral is used as a bridge when moving from one radial thread to the 234 HENAUT ET AL.— WEB VARIATIONS IN LEU GAUGE 235 next during construction of the sticky spiral (Eber- hard 1987). Leucauge venusta is extremely abun- dant during the rainy season in coffee plantations in Chiapas State, Mexico (Pinkus-Rendon pers. comm.). The web of this species is constructed in semi-open sites generally between weeds or be- tween adjacent coffee bushes (Ibarra-Nunez & La- chaud 1998). The objective of the present study was to deter- mine whether a relationship exists between the body size of the female spider or characteristics of the available prey, caught by sticky traps, and the structure and placement of L. venusta webs in a coffee plantation in the south of Mexico. METHODS Field observations. — The field site was a coffee plantation in the grounds of the INIFAP (Institute Nacional de Investigaciones Forestales, Agricolas y Pecuarias) agricultural experimental station at Ro- sario Izapa located at 400 m above sea level, ap- proximately 15 km from the town of Tapachula (Chiapas, Southern Mexico) and 1 km from the bor- der with Guatemala (14° 58' N, 92° 09' W). The climate is tropical; warm and humid with a typical daily temperature range of 35 °C maximum and 23 °C minimum, and a relative humidity of approxi- mately 85%. Heavy rainfall occurs during the months of May to October (—300 mm/month) caus- ing a marked reduction in spider activity in the field. The field collection occurred between October and November (2000), at the beginning of the dry season. Adult female spiders were collected by walking between coffee plants (around 2 m height) in a 1000 m^ area of the coffee plantation at intervals of six days. The characteristics of each web were recorded for each spider collected until a total of 100 spiders and webs had been sampled. Collections were per- formed early in the morning when the webs were still clean and undamaged. Voucher specimens of L. venusta collected were deposited in the ento- mological collection of El Colegio de la Frontera Sur (ECOSUR). Characteristics of the web. — Web characteris- tics were measured directly in the field before col- lecting the corresponding occupant. We recorded the height (in cm) from the soil surface to the center of the orb and to the lowest and highest points of the web, the web diameter (in cm), the number of radii and the number of sticky spirals (from the cen- tre to the bottom of the orb and from the centre to the lower point). Spider characteristics. — The occupant of each web was collected alive in a tube and taken back to the laboratory where body length (in mm), ab- domen width (in mm) were measured with binoc- ular microscope and dry weight (in mg) of each spider was recorded. To determine the dry weight. spiders were placed in aluminium foil, killed by freezing at— 6 °C them 2 min and dried for 2 h in a temperature of 60 °C in an oven. An electronic balance was used to determine the weights (Sarto- rius Basic model BA 11 OS). In November, 30 spi- ders were collected randomly and the abdomen and ovary width were measured to determine whether a relationship existed between abdomen size and the development of the ovary. To measure the ovary size, ovaries were dissected out and the width mea- sured using a binocular microscope at the widest point of the dissected ovary Prey trapping. — To determine the abundance of different types of potential prey, 12 sticky traps were hung from randomly-selected coffee plants in the experimental area for periods of 2 days. Traps were located at 50 cm or 150 cm height above the ground and consisted of a transparent plastic board (30 X 20 cm) coated with Tangle Foot ©(The Tan- gle Foot Company, Grand Rapids MI 49504 USA) similar to those used by Eberhard (1977) and Uetz et al. (1978). Six traps were used for each height. Trapped insects were preserved in 70% ethanol and identified to order in the laboratory. Body size (length and width in mm) of each insect was also recorded as the maximum distance between head and the posterior tip of the abdomen and the max- imum width of the thorax. Statistics. — Principle Component Analysis was applied to determine the factors (web and spiders characteristics) that may separate different groups of spiders inside the study area. Web characteristics, body weight, body length and abdomen width of the spiders of the different groups were compared by Kruskal- Wallis test. The relationship between abdomen width and web characteristics of all the spiders was analysed by a non-parametric Spearman rank order correlation. The size of insects trapped at each height was compared by Mann- Whitney U test, whereas the numbers of insects of each order trapped at each height were compared using contin- gency tables (x^ test). The relationship between ab- domen and ovary width was subjected to linear re- gression analysis. RESULTS Analysis of web and spider characteristics. — Principal component analysis clearly identified the presence of three distinct groups of L. venusta webs (Fig. 1). The first group was numerous with 63 webs, the second group had 18 webs and the third group represented 19 webs (Table 1). The height of webs above the ground increased significantly from an average (± S.E.) of 54.5 ±1.8 cm in group 1 to 153.8±3.6 cm in group 3 (F = 363, d.f. = 2,97, P < 0.001). In contrast, the number of spirals/web was negatively correlated with web height (F = 5.6, d.f. — 2,91 , P = 0.004). Web diameter and the num- ber of radii were statistically similar among the 236 THE JOURNAL OF ARACHNOLOGY 3 PC2 1 2 - ♦ ♦ ♦ H1 H2®H A A A ♦ ♦ ♦ il R O l; A .4 ^ ' 0 ♦ 4 D 4 O A 4 ♦ ♦ ♦ ♦ ♦ ♦ ♦ ♦ ♦ PC1 S2o sC -1 A 1 2 3 ♦ A 4 i;? -1 - -2 - I W2 -3 - Figure 1. — Principle component analysis (n — 100) applied to the height of webs (H, HI, H2), web diameter (D), number of radii (R) per web, number of spirals (SI, S2), live weight (Wl), dry weight (W2), body length (L), and abdomen width (W). We distinguished three spider groups (circles, triangles, squares). The first axis (PCI) explains 41% of the distribution and the second axis, (PC2) explains 21% of the distribution. three groups (Table 1). The height at which the webs were located appears to be the principal factor separating the three groups; most L venusta webs (63/100) were close to the ground. The characteristics of the spiders also differed between the three groups of webs (Table 2). Spiders from web groups 1 and 2 were similar in weight (13.4-13.8 mg) whereas group 3 spiders were sig- nificantly heavier with an average dry weight of 17.3 ± 1.3 mg (F = 3.45, d.f. = 2, 97, P = 0.03). A similar relationship was also detected with body length (F = 3.55, d.f. = 2, 97, P = 0.03) and ab- domen size (F = 3.66, d.f = 2, 97, P = 0.02); group 3 spiders were larger than spiders from group 1 and group 2 webs in all respects. Significant positive correlations were detected between L. venusta abdomen width and height of webs (Spearman = 0.2, t(98) = 2.3, P = 0.02), and abdomen width and number of web spirals (Spear- man = -0.2, t(98) =-2.5, P = 0.01). The diameter of the web (Spearman = 0.07, t(98) = 0.7, P = 0.5), and the number of radii (Spearman =-0.1, t(98) =-1.2, P = 0.2), were not significantly cor- related with abdomen width. There was a significant positive correlation de- tected between abdomen size and ovary size (F = Table 1. — Height, size and structural characteristics of three groups of L. venusta webs collected in Chiapas, Mexico during the rainy season of 2001. Figures represent means ± SE. *** Mann- Whitney U- test, P < 0.001; ns, Not significant P > 0.05. Group \ {n = 63) Group 2 (n = 18) Group 3 (n = 19) P Height of web (cm) 54.5 ± 1. 8 101 ± 2.3 153.8 ± 3.6 *** Web diameter (cm) 29.1 ± 0.6 31.8 ± 1.4 30.1 ± 1.0 ns Number of radii 34.7 ± 0.4 35.1 ± 1.0 35.2 ± 0.9 ns Number of spirals 52.2 ± 1.0 47.0 ± 1.9 45.7 ± 2.2 HENAUT ET AL„— WEB VARIATIONS IN LEU GAUGE 237 Table 2. — Body weight, length and abdomen width of spiders collected from the three groups. All values are means ± SE. Mann- Whitney U-test: * E < 0.05. Group 1 {n = 63) Group 2 {n — 18) Group 3 {n ■= 19) P Weight (mg) 13.4 ± 0.6 13.8 ± 1.2 17.3 ± 1.3 Body length (mm) 5.6 ± 0.7 5.58 ± 0.1 6.1 ± 0.1 * Abdomen width (mm) 2.9 ± 0.05 2.85 ± 0.1 3.2 ± 0.09 * 22.3, d.f. = 1, 12, P < 0.001, Pearson’s correlation coefficient = 0.8); larger spiders had larger, better developed ovaries, presumably indicative of sexual maturity (Fig. 2). Insects trapped. — Significantly more insects were captured in the traps placed at 50 cm height {n — 225) than in the traps placed at 150 cm height {n = 150) (x^ = 14.99, d.f. = I, P < 0.001). The majority of arthropods captured were Diptera (207), Hymenoptera (68) or Coleoptera (55), other arthro- pods found on traps included Homoptera (17), He- miptera (2)Acari (13) and Araneae (13) (Table 3). As Diptera, Hymenoptera and Coleoptera were ob- viously the most common (88% of the trapped ar- thropods), we used these three groups for analysis. A similar number of Hymenoptera and Coleop- tera were caught on traps at 50 cm and 150 cm height (x^ = 2.1, d.f. = 1, P = 0.1 for Hymenop- tera; ^ 1.4, d.f. = 1, P = 0.2 for Coleoptera). In contrast, many more Diptera were caught at 50 cm than at 150 cm height (x^ = 5.9, d.f. = 1, P = 0.01) and they were much more numerous than oth- er insects at both 50 cm (x^ = 75.4, d.f. = 2, P < 0.001) and 150 cm height (x^ = 53.7, df = 2, P < 0.001) (Table 3). However, body length {U = 4165, P = 0.01) and width (t/ = 4210, P = 0.02) of flies trapped at 150 cm height were greater than for flies trapped at 50 cm height (Figure 3). A similar relationship was seen in hymenopterans (body length U = 313, P = 0.01; body width [/= 371.5, P = 0.01). In the case of Coleoptera, body length was greater in individ- uals trapped at 50 cm (U = 243.5, P = 0.03) but no significant difference was detected in terms of body width (C/ = 350, P = 0.7) (Fig. 3). Discussion. — The spatial distribution of the L. venusta population in a coffee plantation was stud- ied in relation to different factors including web characteristics, spider maturity and prey size. It was possible to distinguish three groups of spiders in relation to the height of their webs. The majority of webs were located close to the ground (at ap- 0.5 1.5 2 2.5 3 3.5 4 Abdomen width (mm) Figure 2. — Relationship between ovarian width and abdomen width (P < 0.0001) in Leucauge venusta. 238 THE JOURNAL OF ARACHNOLOGY Table 3. — Total number of prey trapped at two heights for the three principal groups of insects, test: ** p < 0.01; *** P < 0.001; ns Not significant P > 0.05. 50 cm height trap 150 cm height trap P Diptera 121 86 ** Hymenoptera 40 28 ns Coleoptera 32 23 ns Total 193 137 P proximately 50 cm height), but a part of the pop- ulation constructed webs at heights of —100 cm or — 150 cm above the ground. Webs at different heights had similar diameters and number of radii but the number of spirals was reduced in webs at 150 cm height giving the webs a more open struc- ture. Spiders occupying webs at —150 cm were larger than conspecifics occupying lower webs. Body size was shown to be positively correlated with ovarian size in a sub-sample of these spiders. Apparently, sexual development may influence the choice of web site. Web design may also be influenced by sexual development, or may be relat- ed to physical stresses experienced when webs are constructed high above the ground, such as air cur- rents and movement of plant supporting structures. Traps placed at two different heights indicated that the majority of potential prey were dipterans and that more insects were captured at 50 cm than at 150 cm height. Diptera and Hymeeoptera, the pre- ferred prey of L. venusta (Henaut et al. 2001), caught at 150 cm height were, however, signifi- cantly larger than those caught at 50 cm height. Chacon and Eberhard (1980) found similar results using artificial webs: smaller insects were most fre- quently caught in the lowest traps. Ibarra-Nunez et al. (2001) also reported that the most common in- sects caught in Leucauge spp. webs are Hymenop- tera and Diptera, and that Coleoptera were not im- portant prey items. The web location appears related to the sexual development of the spider. Observations in the field confirmed that L. venusta nymphs build webs close to the ground and that immature spiders may be very abundant (Pinkus-Rendon pers.comm.). Our results suggest that young adult spiders also build webs close to the ground, but as sexual develop- ment proceeds, the spider increases the height at which the web is constructed. This vertical migra- tion is also associated with changes in web con- struction and the type of prey captured, with the possible effect of reducing intraspecific competition between L, venusta adults and juveniles. Another possibility is that sexually maturing spiders seek the larger prey that can be caught at higher sites in and between plant canopies. In the same way, a more open web structure with a larger mesh size may be an adaptation to target the capture of larger prey, as reported in Argiope species (Uetz et al. 1978). Uetz and Hartsock (1987) found that webs of Mi- crathena gracilis (Walckenaer 1805) show selectiv- □ 150 cm 50 cm Length Width Length Width Length | Width Dipterans Hymenopterans Coleopterans Figure 3. — Comparison of body length and width of Diptera, Hymenoptera and Coleoptera caught on sticky traps placed at 50 and 150 cm above the ground in a coffee plantation (Mean ± SE). Mann-Whitney U test: * P < 0.05, ** P < 0.01, ns Not significant P > 0.05. HENAUT ET AL.— WEB VARIATIONS IN LEU GAUGE 239 ity for prey sized greater than 3 mm even if poten- tial prey are smaller. The same phenomen of prey specialization may be also occur in L. venusta and in this case be reinforced by web relocation. The dynamics of habitat choice in spiders are complex and difficult to analyze. Spider behavior represents a compromise between many needs dur- ing the life cycle and the choices available may of- ten be limited (Riechert & Gillespsie 1986). Spatial distribution may be influenced by the competition for suitable web sites. Dispersal can occur on two levels, first haphazardly by ballooning on air cur- rents followed by a controlled and active site selec- tion (Richert 1970). This active dispersal has been described by Kronk and Riechert (1979) wherein, similar to the results of our study with L. venusta, the sexual maturity of females induces migration to sites in which prey are more abundant. There are, however, alternative interpretations of these results. First, larger spiders may exclude smaller conspecifics from high locations by aggres- sion (Danielson-Frangois et al. 2002). Second, larg- er spiders that spin looser webs higher in the coffee canopy may also competitively exclude smaller spi- ders that construct webs with closer spirals. Third, predatory wasps may also influence the vertical dis- tribution of spiders; larger spiders experience a sig- nificantly diminished risk of predation (Blackledge & Wenzel 2001). Finally, the vertical migration dur- ing the search for females could be a serious con- straint upon male mating success in orb wearing spiders (Moya-Larano et al. 2002). This suggests that female L. venusta may migrate upwards as a means of selecting especially vigorous males as ma- tes. In conclusion, the variability in web construction in L. venusta is correlated with sexual maturity and predatory opportunities perhaps dictated by the en- ergy required for the development of eggs and re- productive activities. Constructing an open web in a site with an abundance of large prey may repre- sent savings in terms of silk production and a re- duction in intraspecific competition with immature conspecifics. These observations illustrate how web site selection may reflect physiological needs as- sociated with spider reproduction in habitats with heterogeneous predation opportunities. ACKNOWLEDGMENTS Thanks to Jesus Pablo Chavez for technical as- sistance, Javier Valle Mora for statistical advice, Guillermo Ibarra-Nunez, Jean-Paul Lachaud and Trevor Williams for comments and corrections to the manuscript. INIFAP (Rosario Izapa) kindly per- mitted access to the field site. This work was finan- cially supported by CONACyT project number 28869 N. LITERATURE CITED Alcock, J. 1993. Animal Behavior. An Evolutionary Approach, Fifth Edition, Sinauer Associates Inc. Publishers Sunderland, Massachusetts. 625 pp. Anderberg, M.R. 1973. Cluster Analysis for Appli- cations. Academic Press, New York. 359 pp. Blackledge, TA. & J.W. Wenzel, 2001. Silk medi- ated defense by an orb web spider against pred- atory mud-dauber wasps. Behaviour 138:155- 177. Chacon, P & WG. Eberhard. 1980. Factors affect- ing numbers and kinds of prey caught in artificial spiders webs, with considerations of how orb web trap prey. Bulletin of the British Arachnol- ogical Society 5:29-38. Danielson-Fran§ois, A., C.A. Fetterer & P.B. Small- wood. 2002. Body condition and mate choice in Tetragnatha elongata (Araneae, Tetragnathidae). Journal of Arachnology 30:20-30. Eberhard, W.G. 1977. Artificial spider webs. Bul- letin of the British Arachnological Society. 4: 126-128. Eberhard, W.G. 1987. Effects of gravity on tem- porary spiral construction by Leucauge mariana (Araneae: Araneidae). Journal of Ethology 5:29- 36. Eberhard, W.G. 1988. Memory of distances and di- rections moved as cues during temporary spiral construction in the spider Leucauge mariana (A., Araneidae). Journal of Insect Behavior 1:51-66. Henaut, Y, J. Pablo, G. Ibarra-Nunez & T Williams. 2001. Retention capture and consumption of ex- perimental prey by orb-web weaving spiders in coffee plantations of Southern Mexico. Entomo- logia Experimentalis et Applicata 98:1-8. Ibarra-Nunez G. & J.P. Lachaud. 1998. Comple- mentarite spatiale de la predation due aux arai- gnees et aux fourmis en plantation de cafe au Mexique. IV Conference Internationale Franco- phone d’Entomologie, Saint-Malo, France, 5-9 July 1998. 1 pp. Ibarra-Nunez, G., J.A. Garcia, J.A. Lopez & J.P. Lachaud. 200 1 . Prey analysis in the diet of some ponerine ants (Hymenoptera: Formicidae) and web-building spiders (Araneae) in coffee plan- tations in Chiapas, Mexico. Sociobiology 37: 723-755. Kronk, A.E. & S.E. Riechert. 1979. Parameters af- fecting the habitat choice of a desert wolf spider, Lycosa santria Chamberlin and Ivie. Journal of Arachnology 7:155-166. Levi, H.W. 1981. The American orb- weaver genera Dolichognatha and Tetragnatha north of Mexico (Aranaeae: Araneidae, Tetragnathinae). Bulletin of the Museum of Comparative Zoology 149: 271-318. Moya-Larano, J., J. Halaj & D.H. Wise. 2002. Climbing to reach female: Romeo should be small. Evolution 56:420-425. 240 THE JOURNAL OF ARACHNOLOGY Olive, C.W. 1980. Foraging specializations in orb- weaving spiders. Ecology 61:1133-1144. Richert, C.J.J. 1970. Aerial dispersal in relation to habitat in eight wolf spiders species (Pardosa, Aranea, Lycosidae). Oecologia 5:200-214 Riechert, S.E. & R.G. Gillespsie. 1986. Habitat choice and utilization in web-building spiders. Pp. 23—48. In Spiders: Webs, Behavior and Evolution. W.A. Shear, ed., Stanford University Press. Uetz, G.W., A.D. Jonson & D.W. Schemske. 1978. Web placement, web structure, and prey capture in orb-weaving spiders. Bulletin of the British Arachnological Society 4:141-148. Uetz, G.W. & S.P. Hartsock. 1987. Prey selection in an orb-weaving spider: Micrathena gracilis (Araneae:Araneidae). Psyche 94:103-116. Wise, D.H. 1993. Spiders in Ecological Webs. Cam- bridge University Press. Cambridge, UK. 328 pp. Manuscript received 6 December 2002, revised 2 February 2005. 2005. The Journal of Arachnology 34:241-243 SHORT COMMUICATION NEST SITE FIDELITY OF PARAPHIDIPPUS AURANTIA (SALTICIDAE) Kailen A. Mooney^: University of Colorado, Department of EEB, Boulder, CO 80309-0034, USA. Jon R. Haloin: Center for Population Biology, University of California, Davis, CA 95616, USA. ABSTRACT. We investigated the nest building behavior of Paraphidippus aurantia (Lucas 1833) (Sal- ticidae) following the experimental destruction of their nests. We located 61 nests on 52 pine saplings (43 saplings with one nest, nine with two nests) and carefully displaced all spiders and destroyed their nests. On saplings with two spiders, we removed one spider. Of the 52 nests in which the resident spider was left in place, 29 new nests were constructed in the identical location as the nests we removed. Of the 9 nests in which the resident spider was removed, no new nests were constructed. There were no nests constructed in new locations. Despite other suitable nest site locations, P. aurantia showed extreme nest site fidelity following the disturbance. Keywords; Nest guarding, anti-predator strategy, jumping spider, retreat Jumping spiders (Salticidae) build small, com- pact nests out of silk (Richman & Jackson 1992). Adult and juvenile spiders occupy nests when they are not foraging, adult females lay eggs in nests and spiderlings may remain in nests for several days after hatching. Thus, nest sites may have a strong influence on spider success at foraging, avoiding predation and reproduction. Once constructed, nests may be destroyed by abiotic factors (e.g. rain or wind-blown vegetation) and biotic factors (e.g. grazing vertebrates or predators). While a great deal of attention has been given to spider habitat selec- tion and site fidelity with respect to food availabil- ity (Edgar 1971; Kronk & Riechert 1979; Morse & Fritz 1982; Janetos 1986; Riechert & Gillespie 1986), relatively little is know of the responses of spiders to nest destruction. We studied nest site fidelity of Paraphidippus au- rantia (Lucas 1833) (Salticidae) in response to the destruction of its nest. Paraphidippus aurantia builds its nests at the bases of needle clusters on ponderosa pine (Pinus ponderosa Laws. var. sco- pulorum) sapling at the Manitou Experimental For- est (LF.S.D.A. Forest Service, Rocky Mountain Ex- periment Station) in Woodland Park, Colorado USA (39° 06' 02" N, 105° 05' 32" W, elevation 2400 m). Voucher specimens from this work have been de- ^ Current address: Department of Ecology & Evo- lutionary Biology, Cornell University, Ithaca, NY 14853. posited at the Denver Museum of Nature and Sci- ence, Denver, Colorado, USA. On 22 July 2000 and 24 July 2001 we selected 52 small ponderosa pine saplings (< 2 m) with oc- cupied spider nests {n = 22 in 2000, « = 30 in 2001). Forty-three of these saplings had a single occupied nest, while nine saplings had two occu- pied nests. In the later case, the two nests were nev- er on the same sapling branch. The 61 nests (43 saplings with one nest and nine samplings with two nests) were built on branch tips at the bases of nee- dle clusters located at varying heights and aspects (i.e. cardinal directions) in the sapling canopies. Each sapling canopy offered many (> 20) potential nest-building sites that to our eye did not in any respects from those supporting nests. Except for oc- cupied nests, there were no other P. aurantia or nests on the experimental saplings. The weather on the days of the nest destruction was clear to partly cloudy, and it neither rained nor was it particularly windy. We coaxed the spiders from their nests using puffs of air from a rubber bulb until the spiders emerged. We waited until the spiders had traveled at least 20 cm before carefully removing all visible silk threads from the pine nee- dles with our thumbs and forefingers. We continued to observe the displaced spiders for at least 60 sec- onds. The spiders typically remained motionless during nest removal and this subsequent observa- tion period. In no instance did the spider jump from the branch or flee more than 50 cm during the time 241 242 THE JOURNAL OF ARACHNOLOGY Table L — Fate of Paraphidippus aurantia nest sites following nest destruction. Spider left Spider in place removed {n - 52) {n = 9) No nest built 45% 100% Nest built on original site 55% 0% Nest built on new site 0% 0% of our observation. For the nine saplings with two nests, we collected one of the two spiders, but re= moved both nests. The nest sites can thus be divid- ed into two groups, (1) those where the occupying spider was left in place (n = 52), and (2) those where the occupying spider was collected (n = 9), We placed flagging on the ground immediately below each nest site to mark its location. We then monitored the nest building activity on the entirety of each sapling on each of the following two days, and at three to four day intervals thereafter, for a total of 33 days in 2000 and 34 days in 2001. Be- cause we did not mark the displaced spiders, we do not have direct evidence that the spiders observed on subsequent days were the same individuals we displaced. While we do not know the life stage or sex of the displaced spiders, we have these data on 60 spiders collected from the branches of trees sur- rounding the experimental saplings at the time of the experiment: 22% were adult females, 78% were juveniles, and there were no adult males (Mooney unpub. data). Forty-two saplings had evidence of nest con- struction on the day following nest destruction. Thirteen of these 42 nests were abandoned by the second day, leaving 29 saplings with nest sites un- der active construction for two or more days. No additional nest construction began after these first two days. Furthermore, when nest constraction was not initiated within these first two days, we never again observed P, aurantia on the saplings. In 2001 we destroyed 20 newly rebuilt nests 21 days after the first experimental destruction, and four of those nests were rebuilt a second time. Thus, in total we observed 46 instances of new nest constraction fol- lowing removal. The most notable result from our study was that every new nest was constructed in precisely the same locations as destroyed nests of spiders that we had left in place (Table 1). In one particular case we observed that a nest that originally spanned sev- eral needles and a flake of bark was again con- structed to incorporate the bark and needles. No nests were constructed on the nine nest sites from which we removed the spiders, and no nests were constructed elsewhere on the saplings (Table 1). While we did not mark spiders, these results pro- vide strong, indirect evidence that the same spiders whose nests we destroyed also built the new nests; had a previously undetected spider or an immigrant spider built these new nests, some of the new nests would have been constructed on those nine sites. It is unlikely the nine nest sites of removed spiders were neglected by chance alone (x^d) = 219, P < 0.0001). Our results also show that this extreme level of nest site fidelity was not for lack of other suitable nest sites on the saplings. Nine saplings originally supported two spiders, yet the nest sites of the re- moved spiders were never re-used by those spiders we did not remove. In addition, to our eye there v/ere many unused sites on each tree that were in- distinguishable from those actually utilized (see above). Paraphidippus aurantia thus showed extremic nest site fidelity, despite (1) their previous nests having been destroyed at those sites and (2) alter- nate, suitable nest sites apparently being available within the area the spiders would routinely travel during foraging. There are at least two possible ex- planations to this behavior. First, there may be some benefits to re-using a familiar nest site such as (a) more rapid nest reconstruction, (b) improved for- aging surrounding the already familiar habitat of an existing nest site, or (c) improved predator avoid- ance in familiar habitat. Second, the benefit of switching nest sites is predicted to be lower in hab- itats where risk of future nest destruction is ho- mogeneously distributed (Switzer 1993). Sources of threats from predators may be homogeneous within a sapling. For instance, birds (Gunnarrson 1993; Riechert & Hedrick 1990) and ants (Halaj et al. 1997; Eubanks 2001) are both significant predators of spiders, but there is no reason to believe their effects would vary among nest sites within a single sapling. Future work should experimentally test these hypotheses for P. aurantia and investigate whether similarly high nest site fidelity is exhibited by other salticids. In addition, the mechanisms by which the spider recognizes and chooses a partic- ular site for nesting is of interest and deserves fur- ther attention. This research was supported by funds provided by the Rocky Mountain Research Station, U.S. De- partment of Agriculture Forest Service and by the University of Colorado Undergraduate Research Opportunities Program. Mark Gillilan provided ex- tensive field assistance on this project in 2000. Pau- la Cushing identified P. aurantia and provided background on salticid natural history. Robert Jack- son, Yan Linhart, Ken Keefover-Ring and two anonymous reviewers gave helpful criticisms of an earlier draft of this manuscript. Brian Geils, Wayne Shepperd and Steve Tapia (USDA Rocky Mountain Research Station) provided logistical assistance. MOONEY & HALOIN— NEST SITE FIDELITY 243 LITERATURE CITED Edgar, W.D. 197L Life-cycle, abundance and sea- sonal movement of wolf spider, Lycosa {Pardo- sa) lugubris, in central Scotland. Journal of An- imal Ecology 40:303-322. Eubanks, M.D. 2001. Estimates of the direct and indirect effects of red imported fire ants on bio- logical control in field crops. Biological Control 21:35-43. Gunnarsson, B. 1983. Winter mortality of spruce- living spiderS' — effect of spider interactions and bird predation. Oikos 40:226-233. Haiaj, J., D.W. Ross & A.R. Moldenke. 1997. Neg- ative effects of ant foraging on spiders in Doug- las-fir canopies. Oecologia 109:313-322. Janetos, A.C. 1986. Web-site selection: are we ask- ing the right questions? Pp. 9-22. In Spiders— webs, behavior, and evolution. (W.A. Shear, ed.). Stanford University Press, Stanford, USA. Kronk, A.E. & S.E. Riechert. 1979. Parameters af- fecting the habitat choice of a desert wolf spider, Lycosa santrita Chamberlin and I vie. Journal of Arachnology 7:155-166. Morse, D.H. & R.S. Fritz. 1982. Experimental and observational studies of patch choice at different scales by the crab spider Misumena vatia. Ecol- ogy 63:172-182. Richman, D. & R.R. Jackson. 1992. A review of the ethology of jumping spiders (Araneae, Sal- ticidae). Bulletin of the British Arachnological Society 9:33-37. Riechert, S.E. & R.G. Gillespie. 1986. Habitat choice and utilization in web-building spiders. Pp. 23-48. In Spiders — Webs, Behavior, and Evolution. (W.A. Shear, ed.). Stanford University Press, Stanford, USA. Riechert, S.E. & A.V. Hedrick. 1990. Levels of pre- dation and genetically based antipredator behav- ior in the spider, Agelenopsis aperta. Animal Be- haviour 40:679-687. Switzer, P.V. 1993. Site fidelity in predictable and unpredictable habitats. Evolutionary Ecology 7: 533-555. Manuscript received 15 June 2003, revised 10 June 200L 2006. The Journal of Arachnology 34:244-246 SHORT COMMUNICATION A NEW MASTOPHORA FROM ARGENTINA AND THE MALE OF MASTOPHOMjk VAQUERA (ARANEAE, ARANEIDAE) Herbert W. Le¥i: Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138-2902. E-mail: levi@fas.harvard.edu ABSTRACT. A new species of the genus Mastophora is described from Argentina. The male of M. vaquera is described from Cuba. Keywords! Taxonomy, new species, Mastophora After a new revision is published, curators check- ing their collections often find new species belong- ing to the newly revised genus. Because my interest is in revisions and in adequately illustrating previ- ously poorly described species, rather than in de- scribing new species, I leave these to others. But the specimen of Mastophora Holmberg 1876, re- cently found from Argentina is exceptional in ap- pearance, and I feel it should be described along with the male of M. vaquera Gertsch 1955, not pre- viously known. Mastophora adults live in trees, attached to a silken substrate, on branches or leaves sometimes on berries or leaf buds mimicking bird droppings. They have unusual predatory methods. The adults are nocturnal, and give off odors that attract specific male moths, which are caught by swinging a viscid globule on a silken thread toward the prey that has been attracted by the scent. While moths often es- cape from orb webs by shedding scales, the globule attaches to the moth. The animals are very difficult to find, often being located only by finding egg sacs attached to branches. Minute males are rarely found and usually have to be raised from egg sacs (Levi 2003). The methods used are the same as those used in Levi (2003). Abbreviations: AMNH, American Mu- seum of Natural History, New York, U.S.A.; MACN, Museo Argentine de Ciencias Naturaies, Buenos Aires, Argentina; MNHNC, Museo Nacion- al de Historia Naturaies, San Antonio de los Banos, Havana, Cuba; MCZ, Museum of Comparative Zo- ology, Cambridge, Massachusetts, U.S.A. TAXONOMY Family Araneidae Simon 1895 Genus Mastophora Holmberg 1876 Mastophora comica new species Figs. 1-7 Material examined. — Female holotype from Punta Indio, 35°16'S, 57°14'W, Buenos Aires Prov- ince, Argentina, 17 November 1991, M. Ramirez (MACN). Etymology,— The specific name is an adjective referring to the clown-like appearance of the spider. Diagnosis, — The shape, coloration (Figs. 2, 3) and the triangular plate between the slits of the pos- terior face of the epigynum (Fig. 6) are diagnostic and separate M. comica from all other species. Description. — Female holotype: Carapace light to dark brown with three white stripes (Figs. 1-3). Chelicerae yellowish with brown mottling. Labium, endites, sternum dark brown. First coxae brown, others yellowish; legs yellowish first, second and fourth femora with proximal black ring and distal gray band; tibiae with an anterior diagonal black stripe (Fig. 3). Dorsum of abdomen with a black oval on each side and indistinct, gray patches on three anterior swellings, and some black spots (Figs. 3, 4); venter yellowish (Fig. 4). Abdomen wider than long (holotype in poor condition). Total length 7.5 mm. Carapace 3.1 mm long, 3.0 wide, 1.7 wide at constriction behind cephalic region. First femur 3.6 mm, patella and tibia 4.4, metatar- sus 3.2, tarsus 0.8. Second patella and tibia 3.1mm, third 1.7, fourth 2.7. Remarks. — The male is unknown, and no other specimens apart from the holotype have been found. Mastophora vaquera Gertsch 1955 Figs. 8-11 Mastophora vaquera Gertsch, 1955: 240, figs. IS- IS (female holotype from Torriente, Matanzas, Cuba, in AMNH, examined); Levi, 2003: 342, figs. 142-152, 452; map 2G. Specimens examined. — CUBA: San Antonio de los Banos, La Habana, from edge of citrus planta- tion, Sept. 1984, imm. $, adult d, R. Regalado (MCZ and MNHNC). 244 UEWl—MASTOPHORA 245 Figures 1-7. — Mastophora comica new species, female: 1, 2. Carapace and chelicerae: 1. Frontal; 2. Lateral; 3, 4. Carapace and abdomen: 3. Dorsal, with left legs; 4. Lateral, with chelicera; 5-7. Epigynum: 5. Ventral; 6. Posterior; 7. Posterior cleared; 8-11. M. vaquera Levi, male: 8. Dorsal; 9-11. Left palpus: 9. Mesal; 10. Ectal; 11. Apical. Scale bars = genitalia 0.1 mm, all others 1 mm. 246 THE JOURNAL OF ARACHNOLOGY Diagnosis. — As in other male Mastophora, the palpus shows a very large extended median apoph- ysis (Figs. 9-11), and a small sclerotized, pointed embolus near the tip of the bulb (Figs. 9, 11). Most of the visible portion of the bulb containing the wide duct is the tegulum. Mastophora vaquera dif- fers by having a heavier curl at the end of the long median apophysis than other species (Figs. 9-11). Description. — ^Male (from San Antonio de los Banos, La Habana).* Carapace, chelicerae, labium, endites, sternum, legs beige. Carapace with a rect- angular white mark extending and covering two- median tubercles and the four horns (Fig.8). Dor- sum of abdomen white, anterior edge darker (Fig. 8); venter beige with two lateral gray bands that approach each other posteriorly and surround the spinnerets. Palpal patella with no macroseta. First coxa without hook. Unlike female, first femora without tubercles. Abdomen without humps. Total length 1.5 mm. Carapace 0.65 mm long, 0.61 wide, 0.41 wide behind lateral eyes. First femur 0.55 mm, patella and tibia 0,65, metatarsus 0.36, tarsus 0.23. Second patella and tibia 0.50 mm, third 0.30, fourth 0.41. Length of first patella and tibia about equal to width of carapace. Variation: Total length of males L3-L5 mm. I thank Cristian Grismado, who found the new Mastophora species when sorting collections of the Museo 7\rgentieo de Ciencias Naturales and letting me describe it, Cristina Scioscia for permitting the loan of the specimen, and Giraldo Alayon for find- ing the male of the Cuban species. Loma R. Levi polished the writing. LITERATURE CITED Gertsch, W.J. 1955. The North American bolas spi- ders of the genera Mastophora and Agatostichus. Bulletin of the American Museum of Natural History 106:223-254. Levi, H.W. 2003. The bolas spiders of the genus Mastophora. Bulletin of the Museum of Com- parative Zoology 157:309-382. Manuscript received 16 March 2004, revised 11 May 2004. 2005. The Journal of Arachnology 34:247 SHORT COMMUNICATION A REPLACEMENT NAME FOR IRACEMA PEREZ^MILES 2000 (ARANEAE, THERAPHOSIDAE) Fernando Perez-Miles: Seccion Entomologia, Facultad de Ciencias, Igua 4225, 11400 Montevideo, Uruguay. ABSTRACT. Maraca is proposed as a new name for Iracema Perez-Miles 2000 because it is preoc- cupied by Iracema Triques 1996 (Pisces). Two new combinations are established. Keywords I Iracema, new name The genus Iracema was described by Perez-Miles (2000) for a new species of theraphosid spider from Brazil, Iracema cabocla Perez-Miles 2000, unav/are that the name Iracema had been previously used for a Neotropical freshwater fish (Triques 1996). A sec- ond species of Theraphosidae, Paraphysa horrida Schmidt 1994, has been attributed to Iracema by Bertani (2003). To remove the generic homonymy, the replacement name Maraca is here proposed for Iracema Perez-Miles 2000 (Araneae) with two in- cluded species, Maraca cabocla (Perez-Miles 2000), NEW COMBINATION, and Maraca horrb da (Schmidt, 1994) NEW COMBINATION. Maraca (feminine) is a noun in apposition taken from the type locality of Maraca cabocla (“Mara- ca”). I thank Cristiano Moreira, Paulo Henrique Fran- co Lucinda and Rogerio Bertani who advised me of this problem. LITERATURE CITED Bertani, R. & Da-Silva, S.C. 2003. Notes on the ge- nus Iracema Perez-Miles, 2000 with the first de- scription of the male of I. horrida (Schmidt, 1994) (Araneae: Theraphosidae). Zootaxa 362:1-8. Perez-Miles, F. 2000. Iracema cabocla new genus and species of a theraphosid spider from Ama- zonic Brazil (Araneae, Theraphosidae). Journal of Arachnology 28:141-148. Triques, M.L. 1996. Iracema caiana a genus and species of electrogenic neotropical freshwater fish (Rhamphicthyidae: Gymnotiformes: Ostar- iophysi: Actinopterygii). Revue Frangaise de Aquariologie 23:91-99. Manuscript received 15 March 2004, revised 2 April 2004. 247 2005. The Journal of Arachnology 34:248-253 SHORT COMMUNICATION AN EXTREMELY LOW GENETIC DIVERGENCE ACROSS THE RANGE OF EUSCORPIUS ITALICUS (SCORPIONES, EUSCORPIIDAE) Victor Fet: Department of Biological Sciences, Marshall University, Huntington, West Virginia 25755-=2510, USA Benjamin Ganteebein: Department of Genetics, University of Cambridge, Downing Street, Cambridge CB2 3EH, UK Ay§egiil Karata§ and Ahmet Karata§: Department of Zoology, Nigde University, Nigde, Turkey ABSTRACT. Little or no genetic divergence is detected using mitochondrial 16S rDNA sequence com= parisons across the entire geographic range of the scorpion Euscorpius italicus (Herbst 1800) from Swit- zerland, Italy, Slovenia, Greece and Turkey. This is consistent with known absence of patterns of allozymes and morphological variation. Euscorpius italicus is found almost exclusively in human habitations. Its sister species, E. naupliensis, exhibits much higher genetic diversity within southern Greece. We suggest that the natural populations of the thermophilic E. italicus underwent a bottleneck during the glaciations, and that its modern range could be a result of dispersal with humans. Keywords: Scorpions, genetic distance, DNA, 16S rRNA, biogeography A large, conspicuous scorpion Euscorpius itali- cus (Herbst 1 800) has been known to arachnologists for 200 years and to humankind for millennia. It is commonly found in many localities in Italy and Greece, being an especially common species in hu- man habitations (Crucitti 1993; Braunwalder 2001). This species is found from the French Riviera to the northern and eastern shores of the Black Sea. Euscorpius italicus prefers a xeric microclimate (Birula 1917a, 1917b; Braunwalder 2001; Fet et al. 2001). In Italy, this species is locally very abundant and usually synanthropic. To the north of Italy it is limited by the southern Alpine valleys in Italy and Switzerland (Crucitti 1993; Braunwalder 2001); in Turkey and the Caucasus it also does not venture into high mountains (Birala 1917a, 1917b), The species’ elevational preference seems to range from 0-500 m, while reported well-isolated “island” populations above 500 m could be attributed to re- cent human-mediated range expansion (Braunwald- er & Tschudin 1997). Several subspecies were de- scribed in this species (Birula 1917a; Birala 1917b; Caporiacco 1950) but are currently not recognized (Kinzelbach 1975; Vachon 1981; Bonacina 1982; Fet & Sissom 2000)(Fig. 1). A detailed redescrip- tion and taxonomic history of E. italicus was re- cently published by Gantenbein et ah (2002), who also demonstrated the separate species status (well defined by both morphological and molecular cri- teria) for E. naupliensis (C. L. Koch 1837) from southern Greece, for many years considered a syn- onym of E. italicus. In order to assess the species structure of E. it- alicus, we used comparative analyses of the mito- chondrial 16S ribosomal RNA gene, a molecular marker that has been recently applied to resolve the species-level phylogeny of several species of Eus- corpius (Gantenbein et al. 1999, 2000, 2001, 2002; Fet et al. 2002, 2003); for the detailed DNA anal- ysis procedures and phylogenetic tree-building al- gorithms, see Gantenbein et al. (1999, 2000). Seven mtDNA sequences (ca. 400 base pairs each) were aligned using ClustalX 1.81 (Thompson et al. 1997). Two new DNA sequences, obtained for the present study, were deposited in GenBank (http:// www.ncbi.nlm.iiih.gov) under accession numbers: EiTUl (AY371536) Samugiiney Village, Bulancak, Giresun, Turkey, 40°56'N, 38°15'E, 17 February 2003 (coll. A. Karata§), and TiSMl (AY371535) Silvi Marina, Abrazzo, Italy, 42°34'N, 14°05'E, 20 June 2000 (coll. F. Kovarik). Voucher specimens are deposited in the Unites States National Museum (USNM), Smithsonian Institution, Washington, DC, USA. Four additional DNA sequences of E. italicus 248 FET ET Al..~EUSCORPIUS ITALICUS LOW DIVERGENCE 249 14' 21 28^ 35' 49' 42" 35" Figure 1. — Origin of population samples of Euscorpius italicus and E. naupliensis (outgroup; dark circle). published earlier by our research group and its col- laborators (Gantenbein et al. 1999, 2002) were ex- tracted from the GenBank online database. Their abbreviations, accession numbers and geographic origin were: EiBR (AJ389378), Brissago, Ticino, Switzerland, 46°07'N, 08°43'E, 25 May 1996 (coll. B. Gantenbein); E/TOl (AJ298067), Tortoreto, Abmzzo, Italy, 42°47'N, 13°55'E, 7 October 1997 (coll. M. Bellini); EiMVl (AJ506152), Metsovo, Epirus, Greece, 39°46'N, 21°10'E, 13 May 2001 (coll. V Fet); EiSLl (AJ5 12752), Brje, Dobravlje, Aidovscina, Slovenia, 45°46'N, 13°50'E, 7 August 2000 (coll. B. Sket). As an outgroup, we used E. naupliensis: EnZAl (AJ506153), Zakynthos Island, Greece, 37°46^N, 20°46'E, 20 August 1999 (coll. K. Palmer) (Gantenbein et al. 2002). For estimation of within- species variation in spe- cies with moderate genetic variation, the application of networks and cladistic methods has been pro- posed to be the most efficient (Posada & Crandall 2001). Recently developed methods allow evalua- tion of the limits of parsimony (Templeton et al. 1992, 1995). Therefore, we calculated a statistical network that only connects haplotypes with a 95 % confidence limit using the program TCSalpha vl.Ol (Clement et al. 2000). From the length of the DNA sequences we estimated the maximum number of steps that haplotypes can differ from each other for a 95% confidence limit. This statistical cladistic analysis revealed a very weak detectable geograph- ic structure across the entire range of E. italicus. The statistical cladogram in Fig. 2 connects haplo- types of up to 7 mutation steps, whereas gaps are treated as the “fifth” base pair. The estimated level of divergence ranging from zero to four base pair substitutions (i.e. from 0-1.2 % uncorrected “p”) is in a dramatic contrast with the elaborate, deep geographic structure detected using the same mi- tochondrial gene fragment in the congeneric species E. germanus (C.L. Koch 1837) and E. alpha Ca- poriacco 1950 (Gantenbein et al. 2000), E. nau- pliensis (C.L. Koch 1837) (Gantenbein et al. 2002), E. tergestinus (C.L. Koch 1837) (Fet et al. 2002), and E. sicanus (C.L. Koch 1837) (Fet et al. 2003); in each of the listed taxa, within-species divergenc- es were up to 5%. The relatively poor genetic di- versity of the mtDNA marker clearly supports the complete absence of nuclear variation at allozyme loci among Swiss populations of E. italicus (Gan- tenbein et al. 1998) compared with populations of other congeneric species (Gantenbein et al. 2001). The network (Fig. 2) also is consistent with the hy- pothesis of artificial transplantation, which is an im- portant issue for phylogeographic studies on scor- pions (Gantenbein & Largiader 2002). The Swiss haplotype (EiBRl) is identical with the Slovenian haplotype (EiSLl), which supports a very recent transplantation from the east into the region of 250 THE JOURNAL OF ARACHNOLOGY Figure 2. — Maximum Parsimony tree connecting isolated mtDNA haplotypes of a fragment of the 16S rRNA gene. Connections are exclusively drawn with a confidence limit of 95% (i.e., < 7 steps) as calculated according to the method of Templeton et al. (1992). Large circles represent haplotypes; small circles represent hypothetical intermediate haplotypes that connect the haplotypes with each other. Network is not drawn proportionally to genetic distance. See text for the designation of haplotypes. Outgroup shown as a dark circle. Northern Italy and Switzerland. On the other hand, the haplotypes from Italy (EiSMl/TOl) and from Greece (EiMVl) are each connected to each other with only a single mutational step. Bimla (1917a, 1917b) characterized in detail the geographic distribution of E. italicus, describing clearly two disjunct parts of the E. italicus range, “western” (Europe) and “eastern” (Anatolia and Caucasus). This idea still holds as the species has never been found in the eastern part of the Balkan Peninsula (Gantenbein et al. 2002). Bimla (1917a, 1917b) considered the eastern part of the range (a narrow strip along the southern and eastern coasts of the Black Sea) reduced compared to the western, due either to southward increase of the Black Sea basin, or aridization of the climate in Anatolia. Morphologically, E. italicus from the “western” and “eastern” parts of the range are the same spe- cies (Gantenbein et al. 2002). Thus a strong case can be made for recent, even historical time, dis- persal of E. italicus between the “western” and “eastern” portions of its range. From the current network of sampled haplotypes one could speculate that Anatolia might have served as the refuge for E. italicus during the last Pleistocene glaciations. Evidence that Anatolia might have been an important refuge for plants and animals has been recently found for the gall wasp (Andricus quercustozae) by Rokas et al. (2003) who reported a higher within-population diversity in the Anatolian populations than in the European ones. We cannot infer any conclusions about genetic di- versity of populations; for this, many more E. it- alicus populations and specimens per population need to be genotyped for the orthologous mtDNA fragment. The low genetic divergence between the haplotypes, however, supports a recent (postglacial) range expansion of this species. Similar low genetic diversity across a wide range was found in the con- generic species E. flavicaudis, which is not closely related to E. italicus (—10% sequence divergence between species; Gantenbein et al. 1999). In E. flav- icaudis, the combined analysis of multilocus allo- zyme data and mtDNA sequence data also revealed a low diversity, which can be interpreted as the ev- idence of rapid range expansion, most likely by hu- man transplantation (Gantenbein et al. 2001). Eus- corpius flavicaudis is known to be an invasive species since it has been recently reported from places obviously outside its natural habitat, e.g., south of England (Benton 1991) and Umguay (Tos- cano-Gadea 1998), where it manages to survive and reproduce. Hewitt (1996, 1999) lists several exam- ples for rapid natural spreading of animals from glacial refuges into Europe, with dispersal rates of —300 m per year and higher. Hewitt (1990) esti- mated that flightless grasshoppers like Chorthippus parallelus spread from southern Europe to England at a rate of about 300 m per year. In scorpions, however, much lower annual dispersal rates have been determined, which range between 1-30 m, males having a higher dispersal rate (Polls et al. 1985). If we assume as lower dispersal rates for scorpion species than for the flightless grasshop- pers, we have to conclude that E. italicus and E. flavicaudis populations were both spread through human civilization. It is also very likely that these two species had two different glacial refuges: E. italicus, in Anatolia and E. flavicaudis probably in the south of Italy, which has been identified as a FET ET Al^.—EUSCORPIUS ITALICUS LOW DIVERGENCE 251 main refuge for many other species (Taberlet et al. 1998). Unlike other species of Euscorpius, E. italicus was never reported from any of the Aegean islands, or from any Mediterranean islands such as Baleares, Sicily, Sardinia, Corsica or Malta; it has been only recorded from the offshore islands in the Adriatic Sea (Dalmatian coast of Croatia) and Ionian Sea (Corfu, Greece) (Gantenbein et al. 2002). At the present time, this species appears to be successfully dispersing with human assistance, since in parts of its range it is almost or exclusively synanthropic, being found only in human habitations or ruins but not in the wild (Cracitti 1993). Braunwalder (2001) documented that in only 33 records out of 1,031 records in southern Switzerland, E. italicus has been found in decidedly natural habitats. Another sign of its probable dispersal with humans is the fact that this species, like E. flavicaudis, establishes new reproducing populations, often remotely dis- junct from its continuous range. As examples we can mention established populations in lower Don, Russia (Zykoff 1912); in Sion, Valais, Switzerland (Braunwalder 2001); in Ljubljana, Slovenia (Fet et al. 2001); and even in Yemen (Birala 1937) and Iraq (Fet & Kovank 2003). Records from south- western Romania (Mehadija, Oravitza; Birula 1917a, 1917b; confirmed by Vachon 1981) proba- bly also refer to introduced populations. Single specimens of E. italicus have been found in many localities well outside the main range (Fet & Graodis 1987; Fet & Sissom 2000; Gantenbein et al. 2002). Moreover, at least within Europe, the ac- tive transplantation of Euscorpius with the human peddlers of “scorpion oil” (an infusion of olive oil with live scorpions, allegedly of medicinal value) has been possible until recently (Komposch et al. 2001). This frequent anthropochory and synanthro- py, absence from most islands, and high morpho- logical and genetic similarity of the studied popu- lations from Switzerland, Italy, Slovenia, Greece and Turkey, all suggest that the dispersal of E. it- alicus (likely from glacial refugia) might not be an ancient event. Further investigation of multiple pop- ulations could determine an exact refugial origin and possible ways of dispersal of E. italicus. We thank Michael E. Soleglad for his valuable insights, help, and enthusiasm in the study of Eus- corpius. We thank Marco Bellini, Frantisek Kova- nk, Kevan Palmer, and Boris Sket for providing specimens. We are grateful to Elizabeth V. Fet and W. Ian Towler for their skilled assistance in the lab. B.G. was supported with an SNF-IHP grant 83EU065528. LITERATURE CITED Benton, T.G. 1991. The life history of Euscorpius flavicaudis (Scorpiones, Chactidae). Journal of Arachnology 19:105-110. Birula, A. (Byalynitsky-Birula, A. A.). 1917a. Arachnoidea Arthrogastra Caucasica. Pars 1. Scorpiones. Zapiski Kavkazskogo Muzeya (Me- moires du Musee du Caucase), Imprimerie de la Chancellerie du Comite pour la Transcaucasie, Tiflis, A(5), 253 pp. (in Russian). English trans- lation: Byalynitskii-Biralya, A. A. 1964. Arthro- gastric Arachnids of Caucasia. 1. Scorpions. Is- rael Program for Scientific Translations, Jerusalem, 170 pp. Birala, A. (Byalynitsky-Birula, A. A.). 1917b. Fau- ne de la Russie et des pays limitrophes foiidee principalement sur les collections du Musee Zoologique de F Academic des Sciences de Rus- sie. Arachnides (Arachnoidea). Petrograd, 1(1): XX, 227 pp. (in Russian). English translation: By- alynitskii-Birulya, A. 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Mesobu- thus gibbosus (Scorpiones: Buthidae) on the is- land of Rhodes — Hybridisation between Ulysses’ stowaways and native scorpions? Molecular Ecology 11:925-938. Gantenbein, B., M.E. Soleglad & V. Fet. 2001. Eus- corpius balearicus Caporiacco, 1950, stat. nov. (Scorpiones: Euscorpiidae): molecular (allo- zymes and mtDNA) and morphological evidence for an endemic Balearic Islands species. Organ- isms, Diversity & Evolution 1:301-320. Gantenbein, B., M.E. Soleglad, V. Fet, P. Crucitti & E.V. Fet. 2002. Euscorpius naupliensis Caporiac- co, 1950 (Scorpiones: Euscorpiidae): elevation to species level justified by molecular and mor- phology data. Revista Iberica de Aracnologia 6: 13-43. Hewitt, G.M. 1990. Divergence and speciation as viewed from an insect hybrid zone. Canadian Journal of Zoology 68:1701-1715. Hewitt, G.M. 1996. Some genetic consequences of ice ages, and their role in divergence and speci- ation. Biological Journal of the Linnean Society 58:247-276. Hewitt, G.M. 1999. 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Euscorpius flavicaudis (de Geer, 1778) in Uruguay: first record from the New World. Newsletter of the British Arachnol- ogical Society 81:6. Vachon, M. 1981. Remarques sur la classification FET ET M^.—EUSCORPIUS ITALICUS LOW DIVERGENCE 253 sous-specifique des especes appartenant au genre Euscorpius Thorell, 1876 (Scorpionida, Chactidae). Comptes-Rendus 6eme Colloque d’Arachnologie d' Expression Frangaise (Colloque International Eu- ropeen), 1981 (Modena-Pisa). Atti della Societa Toscana di Scienze Natural!, Memorie (B), 88(suppl.):193--203. Zykoff, W.P. 1912. Ueber das Vorkommen von Skorpionen im Dongebiet. Zoologischer Anzeig- er 39:209-211. Manuscript received 1 7 September 2003, revised 14 April 2004. 2005. The Journal of Arachnology 34:254-257 SHORT COMMUNICATION DISPERSAL BY UMMIDIA SPIDERLINGS (ARANEAE, CTENIZIDAE): ANCIENT ROOTS OF AERIAL WEBS AND ORIENTATION? William G. Eberhard: Smithsonian Tropical Research Institute, and Escuela de Biologia, Universidad de Costa Rica, Ciudad Universitaria, Costa Rica Keywords; Mygalomorph ballooning behavior, orientation It is well known that many araneomorph spiders disperse by ballooning (e.g. Decae 1987; Suter 1999), but similar dispersal abilities of mygalo- morph spiders are much less well established. Pre- vious publications are easily summarized. The most complete observations are those of Coyle (1983, 1985) of spiderlings of Sphodros sp. (prob. S. at- lanticus Gertsch & Platnick 1980) (Atypidae), and Ummidia sp. (Ctenizidae). Spiderlings of both spe- cies moved along bands of silk lines, and launched themselves into the air after dangling at the ends of draglines. Other descriptions of mygalomorph bal- looning did not provide details on how spiders took to the air. Baerg (1928) carefully observed move- ments of Ummidia carabivora (Atkinson 1886) (originally described in the genus Pachylomerus) spiderlings from their mother’s burrow along wide silk trails to elevated sites, but did not witness the spiders taking off. Enock (1885) saw that Atypus piceus (Sulzer 1776) spiderlings followed ascend- ing silken cords to upwardly projecting objects, from which they were “blown off into midair . . . until they became attached to other sticks” (p. 394). Muma & Muma (1945) also observed silk bands produced by the spiderlings of Sphodros rufipes (Latrielle 1829) (= Atypus bicolor Lucas); they stated that the spiderlings dispersed by ballooning, but gave no details. Cutler & Guarisco (1995) ob- served a group of spiderlings of S. fitchi Gertsch & Platnick 1980 and apparent ballooning attempts at the top of a small tree. Main (1957) suggested that Conothele malayana (Doleschall 1859) (Ctenizidae) spiderlings balloon, but only on the basis of ob- serving large numbers of fine threads of silk pro- duced by spiderlings held in collecting tubes. This note reports an observation of dispersal and bal- looning by spiderlings of the another ctenizid, an undetermined species of Ummidia. Observations were made in San Antonio de Es- cazu, San Jose Province, Costa Rica (el. 1325 m; approximately 9° 51' N, 84° 10' W). Observations with the naked eye were complemented using a 2x headband magnifier. Some individual silk lines were extremely fine and difficult to see; checks for unseen lines were made by moving an object where a line might have been, and noting whether this movement produced tugs on the spider or nearby silk lines. A voucher (mature female) specimen will be deposited in the Museum of Comparative Zo- ology, Cambridge, MA 02138, USA. At about 09:00 on the cool rainy season morning of 17 Oct. 2002 with only intermittent sunshine and weak, erratic wind (a very fine, weak, intermittent drizzle fell briefly at 12:30, but it did not rain until after 15:00), I noted a small ball of spiderlings at the tip of a long thin leaf growing at the edge of the deck of my house (Figs. 1, 2). A file of spider- lings toiled slowly up the edge of the leaf toward the ball. Below the ball of spiders, I traced a more or less straight trail of shiny silk downward along the edge of the leaf, across an open space of about 10 cm to the rock wall supporting the deck, and then across this irregular surface (Fig. 3) for about 70 cm to its end, the edge of the extremely well- camouflaged ctenizid burrow lid covered with green moss (Fig. 5). In places the trail ran along the sur- face of the wall as a wide band, about the width of a spiderling’s body. In others, where it bridged cracks and spaces of up to 7 cm between rocks, it narrowed to a single thick, aerial line (Fig. 3). Spi- derlings were scattered along this trail. The burrow entrance was about 10 cm above the ground, in de- tritus in a shallow indentation in the wall. Spider- lings were emerging one by one from under the lid, which was hinged at the top and slightly open, and climbing up along the trail. Judging by their rates of movement, they must have been emerging for at least 30 min previously. My tentative attempt to lift the lid was answered by a sharp tug that closed and held it tight, showing that the female resident was evidently holding the inner side of the lid (see Bond & Coyle 1995). The silk trail also continued in the other direction beyond the ball, A few flimsy lines radiated from 254 EBERHARD— BALLOONING IN UMMIDIA 255 Figures 1--5. — Dispersal by Ummidia sp. spider- lings. 1. aerial lines to post of railing; 2. spiderling (right) leaves a ball of spiderlings to walk along an aerial portion of the trail; 3. relatively straight silk trail followed by spiderlings across uneven rock face; v/here the trail was aerial, the band narrowed to a single, thick thread (arrow); 4. horizontal por- tion of the trail of silk along the top of the railing (arrow indicates fork where some spiders went to the right, others to the left); 5. the trail (single ver- tical line, arrow) ends on an adult female’s burrow lid, which is camouflaged with green moss. the tip of the leaf, forming aerial bridges to other leaves 10-20 cm away, and a longer and stronger, approximately horizontal line about 30 cm long connected the leaf tip to the post of the railing of the deck (Figs. 1,2). From here it ran horizontally along the railing for about 6 m (Fig. 4) until it went straight up the corner of the house for about 4 m to the underside of the roof, where several thin lines bridged to the underside of the eaves. Several lines that streamed downwind from the eaves waved about in the light breeze as if their tips (which I was not sure I could see) were free; the nearest object in that direction was some 10 m away. It is possible that these were lines which had broken when the spider was several m from this takeoff site. An estimated 50-100 spiderlings were seen at different points along the trail during the next two hours, nearly all moving away from the burrow. The spiderlings dispersed aerially, both from near the ball and from under the eaves. Most of the spi- deriings in the bail dispersed in the space of 10--15 min around 10:45. No obvious change in wind strength or intensity was noted at this time, but the wind was so light that a subtle change could have gone unnoticed. The last spiderling was seen at about 11:30, In each of five cases in which I ob- served take-off behavior from the beginning, the spiderling first descended on a dragline that was attached to a horizontal aerial line, and then glided smoothly upward and laterally, moving in the same direction as the wind. The longest glide I was able to follow took the spider just over 5-10 m until the line it was on became entangled in a bush. Another glide, in which I lost sight of the spider when it v/as about 5 m above the ground, probably took the spider at least 10 m, judging by the direction in which it was moving and the closest objects in that direction. Much of the spiderlings’ behavior near the ball was apparently tentative. Spiders moved back and forth on the more or less horizontal lines, and as- cended and descended vertical draglines. Most de- scents were followed by ascents of the same line rather than by glides. The draglines were too thin to be easily observed directly, and only occasion- ally, when lighting and background conditions were appropriate, did I succeed in seeing the silk as the spider descended. Nevertheless, the spider’s move- ments (slow descent straight downward with the spider facing downward, with its legs more or less spread and moving little if at all; legs never moving as if walking along a line), left no doubt that they were descending at the tips of lines they were pro- ducing. The spinnerets were spread, at least in some cases, as the spider descended. Spiders produced drag lines as they walked along horizontal lines. Some spiders kept their spinnerets spread as they walked, and in these cases it was clear that the spider produced at least two lines, and probably more. When the spinnerets were directed rearward, these lines apparently merged into a sin- gle thread. Coyle (1985) reported that each spider- ling of Ummidia sp. produced a band of numerous fibers as it moved. In no case did I see a spider break and reel up a line as it walked along it (as is typical of araneoid spiders— Eberhard 1982, 1990; Griswold et al. 1998). Nor did a spiderling ever slide tarsus IV along the dragline as it emerged, or break and reel up aerial lines as it moved along them, other traits that are common in araneomorph spiders as they ascend draglines and produce span- ning lines (Eberhard 1986, 1987). I was not able to decipher with certainty how spiders initiated the horizontal aerial lines along which they walked, or the lines with which they ballooned. As in Sphodros and Ummidia (Coyle 1983, 1985), ballooning was preceded by descent on a dragline. But in no case did the spider give any sign that this dragline broke as it glided up and 256 THE JOURNAL OF ARACHNOLOGY away, as described in these other species (Coyle 1983, 1985). The spider’s upward and lateral glid- ing movement was observed carefully: it was very smooth, and was not interrupted by any perceptible jerk that would be produced when a line broke. Nor did the spider move its legs as if reeling in lines, as occurs in some araneomorphs (Eberhard 1987). These details suggest that no lines v/ere broken. It was as if the spider smoothly lengthened its drag- line while being pulled by another airborne length of silk. One long (>1 m) horizontal line was established by a ballooning spider and then used by several other spiders, supporting the idea that at least these early stages of flight during ballooning did not in- volve breaking the drag line. The new horizontal line ran in just the direction in which the first spider glided away several minutes previously. This line was not present before this spider glided away, be- cause I had walked past this spot several minutes earlier and would have broken any lines there. In a second case, a line was apparently formed by one spider during the time it hung more or less motionless at the tip of a vertical line. The spider was first observed dangling at the end of a dragline. I passed my hands through the air at its sides and between the spider and my ov/n body without hav- ing any effect on its position, thus confirming that there were no unseen lines running from the spider or its dragline in these directions. Nevertheless, in the following minute, during which the spider re- mained at the tip of its dragline, a line was formed that connected the spider or the dragline near it to my body (perhaps 30 cm away): each time I moved, the spider was displaced. A minute or so later, this spider then glided smoothly away out of sight in a slightly different direction. These observations of Ummidia sp. ballooning are compatible with two different hypotheses re- garding the initiation of ballooning lines: the spin- neret spreading idea of Blackwell (fig. 1C in Eber- hard 1987); and the “second line” method (fig. 2 in Eberhard 1987). It is not clear whether my in- ability to confirm the third “dragline breaking” technique for initiating ballooning lines, which was proposed by Coyle (1983, 1985) for other myga- lomorphs, was due to limitations in the resolution of my observations imposed by my general inability to see the lines the spiderlings were producing, to my inability to follow spiders for longer distances (perhaps they break their draglines after having moved several meters through the air), to differenc- es between species in the process of ballooning, or to imprecisions in previous descriptions. If, as in the observations reported here, the extra ballooning lines (in addition to the dragline) were difficult to see in the Sphodros and Ummidia species observed by Coyle, his observations are consistent with both the spinneret-spreading and the second line hypoth- eses, as well as the dragline breaking hypothesis (F. Coyle pers. comm.). Resolution of this uncertainty will unfortunately have to await further lucky oc- casions when ballooning behavior by mygalomorph spiders can be observed again. Perhaps the most useful technique to employ in such a situation would be to lightly dust the lines with cornstarch or talcum powder, to make additional fine lines vis- ible. Several details of pre-ballooning dispersal by mygalomorphs merit comment. In both genera that have been observed, the spiderlings migrate as a group from their mother’s burrow to the ballooning site, forming a strong band of silk (Baerg 1928, Fig. 3). Spiderlings of the theraphosid Brachypeima va- gans (Ausserer 1875) also migrate in single file on the ground, perhaps also following a band of silk (Reichling 2000). Such mass movement, and the resulting formation of compact aerial silk highways, is very unusual in araneomorph spiders. I know of only one other case; the highways produced when colonies of the social theridiid Achaearanea wau Levi 1982 migrate (Lubin & Robinson 1982). The general pattern for dispersing araneomorphs seems to be for each spiderling to strike out on its own. Spiderlings may benefit from moving as groups; following lines established by nest mates may fa- cilitate rapid movement to ballooning sites. The ability (and readiness) of Ummidia sp. spi- derlings to walk upside down along aerial cables (Figs. 1, 2) was surprising. Such dexterity in walk- ing under aerial lines may thus be a very ancient trait, and it could have been important in facilitating the evolution of aerial webs in other groups. How did these mygalomorph spiderlings orient? Perhaps a partial answer is related to a further re- markable detail of the highways: the trails are quite extraordinarily straight (Figs. 3, 4), Baerg (1928), who observed about 30 different trails of U. cara- bivorus ranging from 10-68 feet long, also noted that trails were “a straight line to the nearest tree of considerable size. A tree less than 6 inches in diameter is usually ignored, even if it is much near- er than some larger tree.” Coyle (pers. comm.) has also seen a straight 4 m trail of Ummidia across a grassy lawn to the base of a small holly tree. The trail of the theraphosid B. vagans may also be rel- atively straight; the text description says it “snaked its way” along a road, but the accompanying photo shows a straight line of spiderlings (Reichling 2000). If one makes the apparently reasonable as- sumption that the path of a trail reflects the path followed by the first spiderlings to emerge from the maternal burrow, it seems likely that these animals must have used some sort of landmark orientation or a sun compass to maintain such straight trajec- tories when crossing irregular terrain, an ability documented in some araneomorphs (Corner 1973, 1986; Tongiorgi 1959). Such an ability is surprising EBERHARD— BALLOONING IN UMMIDIA 257 in spiders that probably seldom venture from their burrows once they are established, and that are gen- erally thought to depend largely on substrate vibra- tions rather than visual stimuli to orient in other contexts (Coyle 1986). One possibility is that this possibly ancient orientation ability may have evolved to enable males to search more effectively for females, instead of simply wandering randomly (Bell 1991). Male S. abboti Walckenaer search for females during the day, and may orient visually to- ward tree trunks (Coyle & Shear 1981). The search- ing behavior of mature male mygalomorphs (and for that matter, of mature male spiders in general) seems to be little known, and would repay further study. I thank Jason Bond for identifying the spider, and Fred Coyle and an anonymous reviewer for many helpful comments on the ms. and references. LITERATURE CITED Baerg, W.J. 1928. Some studies of a trapdoor spider (Araneae: Aviculariidae). Entomological News 39:1-4. Bell, W. 1991. Searching Behaviour. New York, Chapman and Hall, Bond, J. & EA. Coyle. 1995. Observations on the natural history of an Ummidia trapdoor spider from Costa Rica (Areneae, Ctenizidae). Journal of Arachnology 23:157-164. Coyle, EA. 1983. Aerial dispersal by mygalomorph spiderlings (Araneae, Mygalomorphae). Journal of Arachnology 11:283-295. Coyle, EA. 1985. Ballooning behavior of Ummidia spiderlings (Araneae, Ctenizidae). 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In Proceeding of the Ninth International Congress of Arachnology, Panama. (W. G. Eber- hard, Y. D. Lubin & B. Robinson, eds.) Wash- ington. DC, Smithsonian Institution Press. Eberhard, W.G. 1987. How spiders initiate airborne lines. Journal of Arachnology 15:1-9. Eberhard, W.G. 1990. Early stages of orb construc- tion by Philoponella, Leucauge, and Nephila spi- ders (Araneae: Uloboridae and Araneidae). Jour- nal of Arachnology 18:205-234. Enock, E 1885. The life-history of Atypus piceus, Sulz. Transactions of the Entomological Society of London 1885:389-420. Gorner, P. 1973. Beispiele einer Orientierung ohne richtende Aussenreize. Fortschritte der Zoologie 21:20-45. Gorner, P. 1986. Adjustment of the optical reference direction in the optical orientation of the funnel- web spider Agelena labyrinthica Clerck. Pp. 109-112. In Proceedings of the Ninth Interna- tional Congress of Arachnology, Panama 1983 (W. G. Eberhard, Y. D. Lubin & B. Robinson, eds.) Washington, DC, Smithsonian Institution Press. Griswold, C.E., J.A. Coddington, G. Hormiga & N. Scharf. 1998. Phylogeny of the orb-web building spiders (Araneae, Orbiculariae: Deinopoidea, Ar- aneoidea). Zoological Journal of the Linnean So- ciety 123:1-99. Lubin, YD. & M.H. Robinson. 1982. Dispersal by swarming in a social spider. Science 216:319- 321. Main, B. 1957. Occurrence of the trap-door spider Conothele malayana (Doleschall) in Australia (Mygalomorphae: Ctenizidae). Western Austra- lian Naturalist 5:209-216. Muma, M.H. & K.E. Muma. 1945. Biological notes on Atypus bicolor Lucas. Entomological News 56:122-126. Reichling, S.B. 2000. Group dispersal in juvenile Brachypelma vagans (Araneae, Theraphosidae). Journal of Arachnology 28:248-250. Suter, R.B. 1999. An aerial lottery: the physics of ballooning in a chaotic atmosphere. Journal of Arachnology 27:281-293. Tongiori, P. 1959. Effects of the reversal of the rhythm of nocturnal illumination on astronomical orientation and diurnal activity in Arctosa vari- ana C. L. Koch (Araneae — Lycosidae). Archivo Italiano de Biologia 97:251-265. Manuscript received 9 September 2003, revised 15 March 2004. 2005, The Journal of Arachnology 34:258-260 SHORT COMMUNICATION REGURGITATION AMONG PENULTIMATE JUVENILES IN THE SUBSOCIAL SPIDER ANELOSIMUS CF. STUDIOSUS (THERIDHDAE): ARE MALES FAVORED? Carmen Viera/^^ Soledad Ghione^’^ and Fernando G, Costa^: ^Seccion Entomologia, Facultad de Ciencias, Igua 4225; ^Laboratorio de Etologia, Ecologia y Evolucion, IIBCE, Av, Italia 3318, Montevideo, Uruguay. E-mail: cviera@fcien.edu.oy ABSTRACT, Regurgitation from adult females towards juveniles is a well known phenomenon in social spiders. However, occasional observations in Anelosimus cf. studiosus from Uruguay showed the occun rence of food transfer also between large juveniles. We experimentally tested if well fed penultimate females were capable of regurgitating fluids to starved males, and if well fed penultimate males were capable of regurgitating fluids to starved females. Other isolated and starved penultimate males and fe- males were used as controls. Starved males and females of the experimental groups significantly increased their body weight, whereas body weight decreased in controls. Males increased their weight more than females. We conclude that both well fed penultimate males and females can feed other starved subadults, but when given access to members of the opposite sex, males benefit than females. This bias in the regurgitation exchange among subadults could contribute to accelerate the maturation of males. Keywords: Social spider, inter-juvenile regurgitation, Anelosimus cf. studiosus, Uruguay Although some solitary species feed their spider- lings by regurgitation, this maternal behavior is considered the first step in the subsocial pathway to social life in spiders (Foelix 1996). Sociality evolved in a few families of spiders in which the juveniles depend on maternal regurgitation feeding (Kullmann 1972; Brach 1977; Buskirk 1981; Darchen & Delage-Darchen 1986; Foelix 1996). This phenomenon has been frequently described in the theridiid genus Anelosimus (Brach 1977; Chris- tenson 1984; Vasconcellos-Neto et al. 1995), which contains both “non-territorial, permanent-social” species such as A. eximius Keyserling 1884, and “non-territorial, periodic-social” species such as A. studiosus (Hentz 1850), following Aviles (1997). However, inter-juvenile regurgitation in spiders has yet to be described. In laboratory conditions, we observed regurgitation from penultimate females to soliciting penultimate males in the subsocial Ane- losimus cf. studiosus (reported in an abstract, Viera et al. 2001). In this paper, we indirectly tested the food transfer and sexual bias among penultimate juveniles by weighing individuals before and after they had access to well fed individuals. This anal- ysis demonstrates an additional means of coopera- tion among spiders. Anelosimus cf. studiosus, taxonomically close to Anelosimus studiosus (Agnarsson pers. comm.) were collected as subadults in Montevideo, Uru- guay (34°53H5"S, 56°08'33"W) during June 2001, from several nests located in low branches of a sin- gle tree. In the laboratory, they were reared in social groups (mixed from different nests) in large petri dishes (8.7 cm diameter and 1,4 cm height), until they reached the penultimate stage. They were fed various fly species (Musca sp. and Drosophila spp.) ad libitum. Penultimate individuals were recognized by size and secondary sexual characters. For the experi- ment, spiders were confined in small petri dishes 3 cm diameter and 1 cm height, without water, and were weighed before the experiment and 24 h later at the end of the experiment. A scale of 0.1 mg of accuracy was used. For the experiments, one spider per dish was deprived of food for 6 days (starved spider). Four experimental groups were carried out simultaneously. In group A (« = 80), a starved pen- ultimate male was maintained with four satiated penultimate females. In group B (« = 40), a starved penultimate female was maintained with four sati- ated penultimate males. In group C (« = 20), a single starved penultimate male was maintained isolated, as a control for group A. In group D (« = 20), a single starved penultimate female was main- tained isolated, as a control for group B, When one or more individuals molted or died during the ex- 258 VIERA ET AL.— INTER JUVENILE REGURGITATION IN A SOCIAL SPIDER 259 Table 1. — Spider weights in the four experimental groups (in mg) after a 24 hour period. Only starved individuals from groups A and B were weighed. Mean weight changes were calculated from the individual differences for each group; relative weight changes were calculated in relation to the initial weight. Experimen- tal groups N Initial weight Mean ± SD Final weight Mean ± SD Weight changes Mean ± SD Relative weight changes (%) Mean ± SD Group A 63 2.314 ± 0.439 2.451 ± 0.438 0.135 ± 0.118 6.164 ± 5.314 Group B 34 2.597 ± 0.521 2.682 ± 0.527 0.085 ± 0.110 3.505 ± 4.865 Group C 20 2.585 ± 0.574 2.540 ± 0.529 -0.045 ±0.110 -1.392 ± 4.357 Group D 20 2.720 ± 0.884 2.690 ± 0.895 -0.030 ± 0.130 -1.198 ± 5.484 periment, that trial was discarded. Room tempera- ture during the period of study averaged 18.7 °C (± 2.5 SD; range 13.5-23.0). The non-paired Student /-test was used to compare difference in weight gain between groups. Voucher specimens were deposited in the Arachnological collection of the Faculty of Sciences, Montevideo, Uruguay. Starved individuals of groups A and B increased their weight, whereas the weight decreased in the control groups C and D (Table 1). Mean weights changes showed statistically significant differences between males from groups A and C (/ = 6.39, P < 0.001); between females from groups B and D (/ = 3.32, P < 0.01); and between males from group A and females from group B (/ = 2.49, P < 0.02); but not between the control groups (t = 0.39, P > 0.60). We estimated the mean weight of these re- gurgitations by adding the daily mean loss of weight per spider (caused by metabolic expenditure, defecation, water loss, silk generation) of the groups C and D plus the mean increment in weight observed in the starved spiders in A and B. Then, we estimated the weight of the regurgitations re- ceived by penultimate males of group A as 0.180 mg, which represents the 7.73 % of their initial weight, whereas females from group B gained 0.115 mg, representing 4.42 % of their initial weight. The increment of body weight in starved penul- timate males and females can only be attributed to the transfer of food from well fed conspecifics, be- cause no other significant source of weight gain, food or water, was available in the experimental pe- tri dishes. These gains in weight seem to be im- portant, taking into account that they occur in only a 24 h period. The increment in group A, where one male was with 4 well fed females, was greater compared to B, where one female was with 4 well fed males in agreement with previous observations. Penultimate females could be better “donors” of regurgitations than penultimate males, or that males could be better “beggars” than the females. We conclude that immature A. cf. studiosus, at least in the penultimate stage, share food among juveniles helping the starving individuals of both sexes, and equalizing the food distribution in the colony. Regurgitation among juveniles could have an important role in the colony, because generally, the mother dies when juveniles are at the fourth or fifth stage (Viera et al. 2002; Viera et al. submitted). Food transfer in the field could be especially sig- nificant for males, considering that they received more food than females in this experiment and in the field, there are two females per male in this species (Viera et al. 2001; Viera et al. submitted). Regurgitation from subadult females could have an important role in determining the early maturation of males observed in this and other Anelosimus spe- cies, possibly reducing inbreeding (Viera et al. 2001; Bukowski & Aviles 2002). Furthermore, it could also provide competitive advantages for mat- ing, as was pointed out by Henschel et al. (1995) and Schneider & Lubin (1997) for Stegodyphus spp. (Eresidae). We thank Marco Antonio Benamu, Fernando Nieto and Rosario Porras for their help in the lab- oratory work. We also thank the Department of Bio- chemistry of the IIBCE for allowing us to use the precision scale, Anita Aisenberg for improving the English and, two anonymous reviewers for their suggestions. LITERATURE CITED Aviles, L. 1997. Causes and consequences of co- operation and permanent-sociality in spiders. Pp. 476 — 498. In The evolution of social behavior in insects and arachnids. (Choe, J. & B. Crespi eds.) Cambridge University Press, Cambridge. Brach, V. 1977. Anelosimus studiosus (Araneae: Theridiidae) and the evolution of quasisociality in theridiid spiders. Evolution 31:154-161. Bukowski, T.C. & L. Aviles. 2002. Asynchronous maturation of the sexes may limit close inbreed- ing in a subsocial spider. Canadian Journal of Zoology 80:193-198. Buskirk, R.E. 1981. Sociality in the Arachnida. Pp. 281-367. In Social insects (2) (H.R. Hermann ed.). Academic Press, New York. Christenson, T.E. 1984. Behaviour of colonial and 260 THE JOURNAL OF ARACHNOLOGY solitary spiders of the theridiid species Anelosi- mils eximiiis. Animal Behaviour 32:725-734. Darchen, R. & B. Delage-Darchen. 1986. Societies of spiders compared to the societies of insects. Journal of Arachnology 14:227-238. Foelix, R.E 1996. Biology of Spiders. Second Edi- tion, Oxford University Press, Oxford. Henschel, J.R., Y.D. Lubin & J. Schneider. 1995. Sexual competition in an inbreeding social spi- der, Stegodyphiis dwnicola (Araneae: Eresidae). Insect Sociaux 42:419-426. Kullmann, E.J. 1972. Evolution of social behavior in spiders (Araneae; Eresidae and Theridiidae). American Zoologist 12:419-426. Schneider, J.M. & Y. Lubin. 1997. Infanticide by males in a spider with suicidal maternal care, Stegodyphiis lineatiis (Eresidae). Animal Behav- iour 54:305-312. Vasconcellos-Neto, J., A.L.T. Souza, E.S.A. Marques & FEE Ferraz. 1995. Comportamiento social de Anelosimus eximius (Theridiidae: Araneae). Anais de Etologia 13:217-230. Viera, C., M.A. Benamu & EG. Costa. 2002. Fen- ologia y desarrollo de la araha social Anelosimus studiosus (Araneae, Theridiidae) en Uruguay. Actas 3’' Encuentro de Aracnologos del Cono Sur: 64. Viera, C., M.A. Benamu, S. Ghione, F. Nieto, R. Porras & EG. Costa. 2001. Trofalaxia entre su- badultos de la araha social Anelosimus studiosus (Araneae, Theridiidae); un sesgo a favor de los machos. Actas VI Jornadas de Zoologia del Uru- guay: 70. Manuscript received 30 June 2003, revised 24 March 2004. 2006. The Journal of Arachnology 34:261-265 SHORT COMMUNICATION ACTIVITY OF JUVENILE TARANTULAS IN AND AROUND THE MATERNAL BURROW Cara ShilMngton: 316 Mark Jefferson, Department of Biology, Eastern Michigan University, Ypsilanti, MI 48197. E-mail: cara.shillington@emich.edu Brian McEwen: Ypsilanti, MI 48198 ABSTRACT. Despite their notoriety, little is known about tarantulas in their natural environment. Here we describe activity of juvenile tarantulas (Brachypelma vagans) in and around the maternal burrow as well as emergence and dispersal behavior. Juveniles remain within the natal burrow for several weeks and undergo at least one molt after emerging from the egg sac. Small numbers of juveniles are active at night and emerge along with the adult female where they remain close to the entrance of the burrow. Most juvenile activity outside the burrowed occurred in the early morning shortly after sunrise when the female was no longer active or visible at the burrow entrance. We also observed juveniles dispersing en masse from the maternal burrow. Spiderlings moved away from the burrow in lines, following one behind each other. Keywords: Juvenile dispersal, natal burrow, tarantulas Cooperation and coordinated movement and ac- tivities are common behaviors among social spiders (see reviews in D’Andrea 1987; Aviles 1997; Uetz & Hieber 1997). However, these types of behaviors have also been observed in solitary spiders which, during early developmental stages, often undergo a brief gregarious phase (Gundermann et al. 1986; Horel et al., 1996; Reichling 2000, 2003; Jeanson et al. 2004). Here we report on activities and be- haviors of juvenile tarantulas, Brachypelma vagans (Ausserer 1875) still living in the maternal burrow and also describe their unique aggregative dispersal (see also Reichling 2000, 2003). Study site. — Our field site was located on a pri- vate dairy ranch in Puebla, Mexico, 0.8 miles west the town Venustiano Carranza. At this site in May 2003, we monitored 117 tarantula burrows in the mowed lawn (approximately 0.5 hectares) imme- diately surrounding the family ranch house. We conducted field observations as part of an ongoing study of the life history of tarantulas. Females with egg sacs were observed in mid-April (pers. Comm.), and on 16 May 2003, we found juveniles still within the natal burrows along with the female (ji = 6). These burrows were closely monitored for a two week period. All observations were made from approximately 50 cm from the burrow en- trance and substrate vibrations caused by our move- ments were kept at a minimum to avoid disturbing the animals. If tarantulas were startled they quickly retreated into their burrows but typically reappeared within 5-10 minutes if there were no further dis- turbances. The majority of observations were per- formed using a red flashlight but white light was sometimes used briefly for greater clarity. Females were visible within their burrows soon after sunset (~ 8:02pm). As sit-and-wait predators, they remained around the burrow entrance for most of the night and were often motionless for more than an hour at a time. In a typical “waiting pos- ture” females positioned themselves halfway out of the burrow with their legs on the 1-2 cm silk collar around the entrance (Fig. 1) (see also Minch 1978). Juveniles appeared at the burrow entrance at least 30 minutes after the female’s nightly emergence. During nocturnal observations, the maximum num- ber of juveniles visible was 15. They always emerged from the buiTOw on the side that was not occupied by the female (Fig. 1). Although they ac- tively moved around and often climbed over each other, there was little physical contact with the fe- male. Juveniles were active throughout the night and constantly changed positions on the silk collar and/or moved in and out of the burrow. They were approximately 6 mm in length (including legs) and, unless they were moving, were sometimes difficult to see in the vegetation surrounding the burrow. Be- cause of the constant movement in and out of the burrow, we could not determine if the same indi- viduals were active throughout the night. In addi- tion, because of their small size and the low light conditions, it was not possible to determine if they 261 262 THE JOURNAL OF ARACHNOLOGY SHILLINGTON & MCEWEN— SPIDERLING ACTIVITY AND DISPERSAL 263 Figure 3. — Exoskeletons from juveniles around the entrance of the natal burrow. White arrows indicate some of the individual exoskeletons. were actively involved in prey capture although we assumed this was the purpose of their emergence from the burrow. Females often remained at the burrow entrance throughout the night and retreated into their bur- rows around sunrise (7:05 am) and were not visible during daylight hours. The end of the nocturnal for- aging period was signaled when the female started laying a thin silk covering over the burrow en- trance. Surprisingly, juveniles often remained active and visible for up to one hour after the female had retreated. They were able to move easily through the web covering laid by the female over the bur- row entrance and during these times we observed as many as 64 individuals around the burrow en- trance (Fig. 2). We suggest that the presence of the female may limit juvenile activity at night. The silk network around the burrow provides an important chemotactic cue for orientation (Minch 1978) and juveniles probably remain in contact with this net- work at all times. When the female forages at night, she occupies a substantial portion of the silk collar so less area is available for juvenile activity. Dispersal of juveniles. — Dispersal of juveniles was observed from only one of the six burrows. On 24 May 2004, the female emerged shortly after sun- set and removed the silk covering from the burrow entrance. She remained inside the burrow with her first pair of legs and pedipalps at the burrow lip. However, at approximately 8:30 pm she disap- peared into the burrow and many juveniles sudden- ly started to emerge. Although we were unable to count individuals because of the large numbers we estimated that there were > 100 juveniles. Although clutch sizes for B. vagans have not been reported, it is not unusual for tarantulas to produce more than 100 offspring in a single egg sac (see summary in Punzo & Henderson 1999). Because of the large number of individuals, many juveniles moved off the web collar lining the burrow but they remained around the burrow entrance. Within a few minutes of this mass emergence, individuals started to move Figure 1 . — Female and juvenile tarantulas at the burrow entrance. The double-headed arrow indicates the silk collar and the three single-headed arrows point to three of the spiderlings around the burrow. Figure 2. — Juveniles around the natal burrow after sunrise. Note the thin silk strands across the burrow entrance. 264 THE JOURNAL OF ARACHNOLOGY Figure 4. — Female burrow with egg sac at entrance. The egg sac was discarded from the burrow after all juveniles had dispersed. away from the bun'ow, starting with the individuals at the outer edge. Instead of dispersing randomly in all directions, juveniles left the burrow in three lines, following one behind each other. Similar lines of juvenile B. vagcms have been observed in Belize (Reichling 2000, 2003). We followed the longest line which initially had 52 individuals in a single column. There was no discernable silk trail but ju- veniles closely followed the path of the individuals ahead of them. At a distance of over 3 meters from the burrow, the line suddenly forked. This started when a single individual left the main column and headed in a different direction. At random intervals, other individuals also left the main column and in- stead followed the new path. At this point, we con- tinued to follow the longest column of individuals and used flags to indicate the path that they trav- eled. Over a 2.5 hour period, several additional “forks” occurred and the number of individuals in the ob.served column was eventually reduced to three. Because of their small size, these individuals were quickly lost in the grass. The distance traveled by these three individuals while we were following them was 14.3m, however; the maximum distance from the maternal burrow was 9m. Over the 2.5 hour period, the path curved around and seldom followed a straight-line direction away from the na- tal burrow. There did not appear to be any specific directionality to the movement nor was it influenced by the slope of the terrain. Instead “leaders” ap- peared to choose the easiest path through the veg- etation. The next morning (25 May 2003) we observed many juvenile exoskeletons around the natal bur- row (Fig. 3). Presumably, the female had discarded them from the burrow although we did not observe this behavior. The small exoskeletons were only visible around the burrow entrance for approxi- mately one day. Because of their light weight, we assumed they were dispersed by air currents or crushed by the female’s movements around the en- trance within a very short period of time. Later that same evening, we observed seven additional juve- niles emerging from the burrow soon after sunset. Their behavior was similar to that of their siblings the night before. They sat around the lip of the bur- row for only a few minutes and then started to move away. Interestingly, they started along the same path as the column we had followed the pre- vious night which suggests they were able to detect a chemical or tactile cue laid down by their siblings. Finally, on the morning of 26 May 2003, we ob- served the egg sac at the entrance to the female’s burrow (Fig. 4). After removing the egg sac from the burrow, we did not see any other juveniles in or around the burrow although juveniles were still present at the other five burrows. Unfortunately we did not observe emergence and dispersal of juve- SHILLINGTON & MCEWEN— SPIDERLING ACTIVITY AND DISPERSAL 265 niles from the other burrows although by the end of May 2003, juveniles had disappeared from two additional burrows. The emergence and dispersal from the natal bur- row occurs very suddenly and from our observa- tions we were unable to predict when these behav- iors would occur. More information is needed to better understand and explain the gregarious phase in these typically solitary animals and to identify the mechanisms underlying this type of collective dispersal. As suggested by Reichling (2000) these behaviors may explain aggregations of tarantula burrows in their natural environment and may allow spiderlings to cluster in a more favorable environ- ment (Jeanson et al. 2004). We would like to thank the Alagon family for allowing us access to the site and providing such wonderful accommodations. We also thank George Odell for his help and support. Partial funding for this research was provided by Eastern Michigan University. LITERATURE CITED Aviles, L. 1997. Causes and consequences of co- operation and permanent-sociality in spiders. Pp. 476-498. In The Evolution of Social Behaviour in Insects and Arachnids. (J.C. Choe & B.J. Crespi, eds). Cambridge University Press, New York, New York. D’ Andrea, M. 1987. Social behaviour in spiders (Arachnida, Areaneae). Monitore Zoologico It- aliano (Nuova Serie), Monografia 3:1-156. Gundermann, J.L., Horel, A., & Krafft, B. 1986. Experimental manipulation of social tendencies in the subsocial spider Coelotes terrestris. Insec- tes Sociaux 40:219-229. Jeanson, R., Deneubourg, J.-L., & Theraulaz, G. 2004. Discrete dragline attachment induces ag- gregation in spiderlings of a solitary species. An- imal Behaviour 67:531-537. Horel, A., Krafft, B., & Aron, S. 1996. Processus de socialization et preadaptations comportemen- tales chez les araignees. Bulletin de la Societe Zoologique de France 21:31-37. Punzo, E and Henderson L. 1999. Aspects of the natural history and behavioral ecology of the ta- rantula, Aphonopelma hentzi (Girard 1854) (Or- thognatha: Theraphosidae). Bulletin of the Brit- ish Arachnological Society 11:121-128. Reichling, S.B. 2003. Tarantulas of Belize. Krieger Publishing Company, Malabar, Florida. Reichling, S.B. 2000. Group dispersal in juvenile Brachypelma vagans (Araneae, Theraphosidae). Journal of Arachnology 28:248-250. Uetz, G.W & Hieber, C.S. 1997. Colonial web- building spiders: balancing the costs and benefits of group-living. Pp. 458 — 475. In The Evolution of Social Behavior in Insects and Arachnids. (J.C. Choe & B.J. Crespi, eds). Cambridge Uni- versity Press, New York, New York. Manuscript received 27 January 2005, revised 21 February 2006. 2006. The Journal of Arachnology 34:266-268 SHORT COMMUNICATION TYPES OF SHELTER SITES USED BY THE GIANT WHIPSCORPION MASTIGOPROCTUS GIGANTEUS (ARACHNIDA, UROPYGI) IN A HABITAT CHARACTERIZED BY HARD ADOBE SOILS Fred Punzo: Department of Biology, University of Tampa, 401 W. Kennedy Blvd., Tampa, Florida 33606 USA. E-mail: fpunzo@ut.edu ABSTRACT. Shelter site selection by Mastigoproctus giganteus in an atypical microhabitat in the north- ern Chihuahuan Desert characterized by hard adobe soils is described for the first time. The majority of the 321 whipscorpions (70.4%) were found within rock crevices during periods of highest daytime ambient temperatures, as compared to those found under plant debris (4.4%) or inside small mammal holes (25.2%). The percentage of available crevices, holes or plant debris that were occupied by whipscorpions was 41.5, 3.8 and 7.3%, respectively. Most occupied crevices (66.7%) were in the shade. Depths of occupied crevices ranged from 6.4-36.7 cm. Crevice widths ranged from 0.7-2. 9 cm. Whipscorpions used crevices whose height above the surface of the ground ranged from 6.5 cm-l.l m. No whipscorpions were observed at the ground surface, even in shaded areas, between 0645 and 1910 hr (CST). Keywords: Retreat, rock crevices Whipscorpions (Arachnida, Uropygi) are found worldwide, from southeastern Asia, Indonesia, Aus- tralia, New Zealand, India, the whole of Africa and Europe, as well as North and South America (Po- cock 1895; Haupt 2000). The posterior 3 pairs of legs are used for walking whereas the anterior pair are modified as sensory organs which allow then to detect and respond to chemical and tactile stimuli (Geethabali & Moro 1988). Thelyphonids possess an attenuated, multi-segmented tail, and typically have median and lateral eyes (Shultz 1990). The genus Mastigoproctus (Arachnida, Uropygi, Thelyphonidae), with 14 species, occurs in Cuba, the Antilles, Mexico, southern regions of the United States, and South America (Haupt 2000). Depend- ing on the species, these arachnids typically inhabit mesic habitats and can be found beneath logs, leaf litter, and within burrows (Cloudsley-Thompson 1991). Some species have adapted to xeric condi- tions and can be found in more arid woodlands and forests in Columbia, Brazil, and desert regions in Mexico and North America (Rowland & Cooke 1973). These arachnids are well known for their ability to spray defensive, vinegar-like secretions from their pygidial glands (Schmidt et al. 2000). The giant whipscorpion Mastigoproctus gigan- teiis (Lucas 1835) is a common representative of the arachnid fauna of the northern Chihuahuan De- sert (Punzo 2001). It is a nocturnal predator that feeds on a wide variety of arthropod prey (Punzo 2000a). It typically seeks shelter during daylight hours beneath surface plant debris, in shallow bur- rows, or within rock crevices (Punzo 2000b). In Big Bend National Park (BBNP), located in the Big Bend region of far west Texas (Brewster County; northern region of the Chihuahuan Desert), M. gi~ ganteus is most commonly found in microhabitats associated with sand-loam soils characterized by soil hardness values ranging from 7. 2-8. 3 kg/cm^, and least likely to be found in areas where hard, adobe soils predominate (penetrometer readings: 37-41 kg/cm2; Punzo 2000a, 2001). Burro Mesa (31°47'N, 103°18'W; elevation: 870- 917 m) is located in the west-central region of the Park (Maxwell et al. 1967). Although this site is characterized by hard, adobe soils (38-40 kg/cm^) and an abundance of rocks and small boulders, M. giganteus does occur at this location (Punzo 2001). Because plant surface debris is sparse at this site, nymphs and adults of M. giganteus typically seek shelter from harsh summer daytime temperatures within rock crevices. The purpose of this study was to identify types of shelter sites and analyze specific physical features of rock crevices used by M. gi- ganteus. The study sites consisted of three 30 m transects chosen at random within a 1 .0 km radius of Burro Mesa. Whipscorpions were hand-collected (be- tween the hours of 1200 and 1500 hr) during June and July of 2002 by walking slowly through the 266 PUNZO— SHELTER SITES USED BY WHIPSCORPION 267 area during daylight hours. This time period is char- acterized by the highest ambient temperatures at this site (36.9-41.2 °C). For each animal collected I recorded total body length, the air temperature (held 1 cm above substrate where animal was ini- tially observed), and type of shelter site where it was found. Rock crevices were inspected using a high-inten- sity hber optic illuminator (Model ER-59-2242, Wards, Rochester, NY). This provided adequate il- lumination of the deepest crevices. When an animal was observed within a crevice, a Im wooden ruler was slowly inserted into the crevice until its tip touched the animal in question, and the distance from the crevice opening to the animal was record- ed. In order to determine the species encountered the animal was then gently prodded to the surface using a plastic rod, 75 cm in length. I also recorded the following measurements as- sociated with rock crevices occupied by M. gigan- teus: (1) depth of the crevice; (2) width of the crev- ice; (3) height of the crevice from the ground and (4) whether the occupied crevice was found in the shade, open sun or sun-shade mosaic. Additionally, I counted the number of crevices and other potential shelter sites (plant debris, occupied and abandoned mammal burrows) within the transects, and searched under plant debris and within burrows for the presence of whipscorpions. Burrows ranged in depth from 12-48 cm. Voucher specimens (SR- 67815-67821) have been deposited in the Inverte- brate Collections at Sul Ross State University (Al- pine, TX), and at the University of Tampa. Mean monthly air temperatures (1200 CST) at study sites were 37.8 °C ± 0.14 SE (June) and 38.7 °C ± 0.08 (July). A total of 456 whipscorpions were found. Thirty-nine of these were tritonymphs (8,5%) ranging in body length from 35-40 mm; 231 (50.7%) were adult males (44-52 mm); and 186 (40.8%) were females (48-57 mm). No proto- or deutonymphs were found. The majority of whipscorpions were found with- in rock crevices (n = 321; 70.4%; 26 nymphs, 152 males, 143 females) as compared with those found under surface plant debris (n = 20; 4.4%; 3 nymphs, 12 males, 5 females) and within holes in the ground (n = 115; 25.2%; 29 nymphs, 39 males, 47 females) (Chi square: = 46.73, P < 0.05). Along transects there were 773 crevices, 1564 holes and 516 clumps of plant debris. The percentage of available crevices, holes or plant debris that were occupied by whipscorpions was 41.5, 3.8 and 7.3%, respectively. The most common plant debris shel- tering whipscorpions were fallen leaves or stems of lechuguilla {Agave lechuguilla), sotol {Dasylirion leiophyllum), blind prickly pear (Opuntia rufida), and rat-tail cactus (Coryphantha pottsii). Holes ranged from 8-37 cm in depth. All whipscorpions found within crevices, bur- rows or under plant debris were alone and those found in rock crevices had their entire bodies within the crevice. Most of the crevices with animals were in the shade (66.7% or 201 out of 301), as com- pared to crevices with animals in sun-shade mozaic (20.9% or 63 out of 301) and open sun (12.2% or 37 out of 301). Depths of crevices occupied by whipscorpions ranged from 6.4-36.7 cm (mean: 18.23 ± 5.41 SE). Width of crevices ranged from 0.7-2. 9 cm (mean: 1.64 ± 0.44 SE). Whipscorpions used cracks in sur- face rocks whose height above the surface of the ground ranged from 6.5 cm-1.1 m (mean: 1 1.32cm ± 2.58 SE). These results show that individuals of M. gigan- teus prefer to use rock crevices at these study sites, where hard adobe soils predominate, even though there are over twice the number of holes present. Out of 1564 holes that were located, only 115 (3.8%) contained a whipscorpion. Most of the holes examined were occupied by rodents {n = 863; 55.1%) or shrews {n = 121; 7.7%) which indicates that small mammals are capable of excavating bur- rows, even in the presence of hard soils. Mastigo- proctus giganteus, in contrast, may not only lack this ability, but may avoid burrows occupied by small mammals such as grasshopper mice, deer mice and shrews, animals known to include arthro- pods in their diets (Schmidly 1977; Punzo 2003). Previous studies on the efficacy of different types of shelter sites to protect desert arthropods from high daytime summer temperatures have indicated that ambient temperatures immediately below plant debris are typically higher than those associated with crevices and burrows (Cloudsley-Thompson 1975; Crawford 1981). Thus, during periods of highest ambient temperature, seeking refuge under plant debris may not allow ectotherms to adjust body temperatures within the preferred range (Pun- zo 2000b). This may explain why only a small per- centage of whipscorpions (4.4%) were found under plant debris at Burro Mesa. This appears to apply to other large arthropods as well. Out of 516 clumps of plant debris, only 11 (2.1 %) were occupied by scorpions (Vaejovidae), 8 (1.5%) by solifugids (Er- emobatidae), 7 (1.3%) by wolf spiders (Lycosidae), 6 (1.1%) by male tarantulas (Theraphosidae), and 9 (1.7%) by centipedes (Scolopendromorpha). No whipscorpions were found at the ground sur- face, even in shaded areas, between 0645 and 1910 hr (CST). This is in agreement with the nocturnal activity patterns reported for M. giganteus at other sites within BBNP (Punzo 2000a) as well as other desert areas (Cloudsley-Thompson 1991). The only arthropods regularly observed actively moving over the ground surface between 1 200 and 1 500 hr were harvester ants (Formicidae: Pogonomyrmex spp.), velvet ants (Mutillidae: Dasymutilla spp.), and the desert millipede (Orthoporous ornatus). 268 THE JOURNAL OF ARACHNOLOGY I thank G. Stratton, R Cushing, D. Mott and anonymous reviewers for critical comments on an earlier version of the manuscript, L. Ludwig, J. Bot- trell, C. Fisher and K. Smart for assistance in lo- cating and observing animals in the field, and the University of Tampa for providing a Faculty De- velopment Grant which made much of this work possible. LITERATURE CITED Cloudsley-Thompson, J.L. 1975. Adaptations of ar- thropods to desert environments. Annual Review of Entomology 20:261-283. Cloudsley-Thompson, J.L. 1991. Ecophysiology of Desert Arthropods and Reptiles. Springer, Hei- delberg. Crawford, C.S. 1981. Biology of Desert Inverte- brates. Springer, Berlin. Geethabali, G. & S.D. Moro. 1988. The general be- havioural patterns of the Indian whipscorpion Thelyphomis indicus. Revue Arachnologique 7: 189-196. Haupt, J. 2000. Biologic der GeiBelskorpione (Uro- pygi, Thelyphonida). Memore Societa Entomo- logia Italia 78:305-319. Maxwell, R.A., J.T. Lonsdale, R. Hazzard & J. Wil- son. 1967. Geology of Big Bend National Park, Brewster County, Texas. University of Texas Publication No. 6711, Austin, Texas. Pocock, R.I. 1895. Whipscorpions and their ways. Knowledge 18:272-274. Punzo, F. 2000a. Diel activity pattern and diet of the giant whipscorpion Mastigoproctus giganteus (Lucas) (Arachnida, Uropygi) in Big Bend Na- tional Park (Chihuahuan Desert). Bulletin of the British Arachnological Society 11:385-387. Punzo, E 2000b. Desert Arthropods: Life History Variations. Springer, Heidelberg. Punzo, F. 2001. Geographic variation in male courtship behavior of he giant whipscorpion Mastigoproctus gigan- teus (Lucas) (Arachnida, Uropygi). Bulletin of the British Arachnological Society 12:93-96. Punzo, F. 2003. Natural history and ecology of the desert shrew Notiosorex crawfordi from the northern Chihauhaun Desert, with notes on cap- tive breeding. Mammalia 67:541-550. Rowland, J.M. & J.A. Cooke. 1973. Systematics of the Arachnid order Uropygida (= Thelyphonida). Journal of Arachnology 1:55-71. Schmidly, D. J. 1977. The Mammals of Trans-Pecos Texas. Texas A & M University Press, College Station, Texas. Schmidt, J.O., F.R. Dani, G.R. Jones & E.D. Mor- gan. 2000. Chemistry, ontogeny, and role of py- gidial gland secretions of the vinegaroon Masti- goproctus giganteus (Arachnida: Uropygi). Journal of Chemical Ecology 54:67-83. Shultz, J.W. 1990. Evolutionary morphology and phylogeny of Arachnida. Cladistics 6:1-38. Manuscript received 29 August 2004, revised 14 April 2005. 2006. The Journal of Arachnology 34:269-272 SHORT COMMUNICATION FIRST CASE OF MATERNAL CARE IN THE FAMILY CRANAIDAE (OPILIONES, LANIATORES) Glauco Machado: Museu de Historia Natural, Institute de Biologia, CP 6109, Universidade Estadual de Campinas, 13083-970, Campinas, SP, Brazil. E-mail: glaucom@unicamp.br Joseph Warfel: 537 Boston Road #1, Billerica, Ma 01821, U.S.A. ABSTRACT. In this paper, we provide the first observations of maternal care for the harvestman family Cranaidae. Adult females of two species, Santinezio serratobialis Roewer 1932, which belongs to the group curvipes, and Santinezia sp., which is probably a new species of the group gigantea, were found in association with egg clutches. Since the microhabitats used for oviposition by these species are very similar, we believe that maternal care may be a synapomoiphic trait of the genus Santinezia. Keywords: Evolution, Gonyleptoidea, Santinezia, subsocial behavior The Cranaidae comprises 75 genera and 143 spe- cies of large-bodied harvestmen (Kury 2003). The family is distributed in the northern region of South America, along the Andes and Amazon Basin up to Panama and Venezuela (Pinto-da-Rocha & Kury 2003). So far, there is no information on the biology of the cranaids, perhaps because they occur in a biome where few studies on harvestmen have been done (but see Friebe & Adis 1983). In this paper, we provide the first behavioral data for the family, describing maternal care in two species, namely Santinezia serratobialis Roewer 1932 and Santine- zia sp., which probably is a new species. Two females of S. serratobialis were found car- ing for offspring during a field trip to Trinidad, con- ducted in July 1999 by the second author. Obser- vations were made at two sites. Mount St. Benedict (10°39'N; 6ri4'W) and Paria Springs (10°46'N; 61°14'W). The Mount St. Benedict site is located 10 km northeast of the capital Port of Spain, and field observations were done in a small forest frag- ment. The Paria Springs site is located near the vil- lage of Brasso Seco, in the northern coastal moun- tains of Trinidad, and field observations were made along an isolated road cut. The batches were pho- tographed in the field so that it was possible to count the number of eggs in the laboratory and also to identify the harvestmen. Only two cranaids are known to occur in Trinidad and Tobago (cf. Kury 2003): Santinezia serratoti- bialis, which is one of the most common harvest- men species on the island (R. Pinto-da-Rocha pers. comm.), and Phareicranaus calcariferus (Simon 1879), which was described from Colombia but was also recorded for the Tucker Valley, nearly 15 km south of Port of Spain (Goodnight & Goodnight 1974). Comparisons between our photos and indi- viduals of the former species collected in Trinidad and deposited in the Museu de Zoologia da Univ- ersidade de Sao Paulo, Brazil, allowed us to identify the guarding females as S. serratotibialis. However, since the individuals were not collected, there are no voucher specimens. Individuals of S. serratotibialis at Mount St. Ben- edict site were found at the bottom of a small ra- vine, a short distance off a trail inside the forest. One female was observed resting on 38 white, re- cently laid eggs within a small sheltered damp pocket, high up on a steep overhanging embank- ment. These eggs were very large compared to the guarding female, with diameters ranging from 23- 26% of the dorsal scutum length of the female. At Paria Springs, another guarding female was found near the bottom of a small creek bed within a small damp pocket among tree roots exposed on the steep slope (Fig. 1). There were 70 dark eggs and three early-hatched nymphs (Fig. 1). Both females were found in a stereotyped position, similar to that de- scribed for guarding females in other harvestman species (e.g., Gnaspini 1995; Machado & Oliveira 1998, 2003). Although they were not seen groom- ing or protecting the offspring against predators, we assume that the behavior described here corre- sponds to a case of maternal care. An analysis of the harvestmen collection of the Museum of Comparative Zoology (MCZ), Harvard, USA, revealed another case of maternal care in a cranaid species. One female of Santinezia sp. was 269 270 THE JOURNAL OF ARACHNOLOGY MACHADO & WARFEL— MATERNAL CARE IN THE FAMILY CRANAIDAE 271 collected in Valle del Cava (ca. 1800 m), above Felidia, western Cali, Colombia. The individual was collected in January 1977, and the collecting label stated that the female was “guarding the eggs”. The female was found on the eggs in a typical resting position and the eggs were attached to the roof of a small natural cavity in a ravine along a road cut bordering the forest (W.G. Eberhard pers. comm.). According to the collecting label, the eggs num- bered 103 and there was a sample of 21 large eggs in the vial containing the female. Unfortunately, the eggs were not well preserved, thus it was not pos- sible to measure their diameter accurately. Recently, Pinto-da-Rocha & Kury (2003) pub- lished a phylogenetic hypothesis for the genus San- tinezia, dividing it into three monophyletic groups: group curvipes (11 spp.), group festae (2 spp.), and group gigantea (8 spp.). The species studied here are representatives of two of these groups: S. ser- ratobialis belongs to the group curvipes and San- tinezia sp. belongs to the group gigantea (Pinto-da- Rocha & Kury 2003; A.B. Kury pers. comm.). Until more information on the other species of the genus become available it is not possible to know if the maternal behavior in these two species is ho- mologous. Since the microhabitats used for ovipo- sition by these species are very similar, we hypoth- esize that subsocial behavior is a synapomorphic trait of the genus Santinezia and predict that study of congeneric species will reveal further cases of maternal care. The genus Santinezia shows several morpholog- ical convergences with the genus Goniosoma (Gon- yleptidae), which is endemic of the Atlantic Forest (Kury 2003). According to Pinto-da-Rocha & Kury (2004), only details of leg armature and the male genitalia betray their far remote common ancestry. Species of both genera are large-bodied harvest- men, with glossy teguments, stout and long legs bearing few spines, robust and heavily armed ped- ipalps, and area II projecting into I until it touches the scutal groove (Fig. 2). In this study we add oth- er convergent traits relating to behavior: females in the two genera lay large eggs and care for the off- spring until the nymphs hatch (Figs. 1-2). More- over, some species of Goniosoma may also lay eggs on damp pockets in ravines and on rocks along river banks (Machado 2002; Fig. 2). As studies on harvestmen behavior have ad- vanced, several cases of parental care have been described (review in Machado & Raimundo 2001). Maternal care is present in at least five families of the suborder Laniatores, including representatives of the infra-orders Grassatores (Cosmetidae, Cran- aidae, Gonyleptidae and Stygnopsidae) and Insidia- tores (Triaenonychidae). The cranaids belong to the superfamily Gonyleptoidea, which embraces the great majority of cases of maternal care in the order (nearly 80% of the total). All families comprising the Gonyleptoidea have an almost exclusively pan- tropical distribution, being most common in wet, warm environments, such as forests and .caves (Shear 1982). Therefore it is possible that maternal care in harvestmen is a convergent behavioral trait adopted by some lineages in response to similar ecological pressures (Machado & Raimundo 2001). One important question to be investigated in the future is why this behavior has evolved in some species, and not in others. The hypothesis first put forth by Wilson (1971) postulates that intense pre- dation on eggs by conspecifics and ants, as well as the high risk of fungal attack in tropical rain forests may have been the major forces favoring the evo- lution of parental care in arthropods. Although this hypothesis may explain why maternal care is so fre- quent among the tropical Gonyleptoidea, it does not provide an answer to question raised above. More recently, Tallamy & Wood (1986) proposed that the answer to this question involves many interacting factors, such as morphological and physiological characteristics of the species, the presence of some behavioral pre-adaptations and phylogenetic con- straints. Accordingly, maternal care in arthropods is expected to evolve when females (1) live long enough to benefit the offspring after oviposition, (2) are able to defend the offspring against predators, and (3) are constrained to semelparity (sensu Tal- lamy & Brown 1999). The morphological and be- havioral convergence between goniosomatines and cranaids may provide phylogenetically independent data to test these predictions and thus may consti- tute appropriate starting point for studies on the evolution of maternal care in harvestmen. We thank Bill Eberhard for additional informa- tion on Santinezia sp., Adriano B. Kury and Ricar- do Pinto da Rocha for identifying the species and for comments on the manuscript, Paula Cushing for helping us during the fieldwork and for providing the coordinates for the Trinidad sites, Bruno A. Buzatto for the photo of the Goniosoma female, Jim Figures 1—2. — 1. Female of the cranaid havestman Santinezia serratotibialis caring for prior hatching eggs and some early hatched nymphs on a small damp pocket among tree roots in Trinidad (photo by J. Warfel); 2. Female of the gonyleptid havestman Goniosoma sp. caring for recently laid eggs on a quite similar microhabitat in Parque Estadual Intervales, Sao Paulo state, southeastern Brazil (photo by B.A. Buzatto). Note that, despite phylogenetic distance, these species are morphologically very similar. Scale bars = 1 cm. 272 THE JOURNAL OF ARACHNOLOGY Costa and two anonymous reviewers for critically reading the manuscript. GM is supported by grants from Funda^ao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP, proc. # 02/00381-0). LITERATURE CITED Friebe, B. & J. Adis. 1983. Entwicklungszyklen von Opiliones (Arachnida) im Schwarzwasser- Uberschwemmungsald (Igapo) des Rio Tucuma Mirim (Zentralamazonien, Brasilien). Amazoni- ana 8:101-110. Gnaspini, P, 1995. Reproduction and postembryonic development of Goniosoma spelaeum, a caver- nicolous harvestman from southeastern Brazil (Arachnida: Opiliones: Gonyleptidae). Inverte- brate Reproduction and Development 28:137- 151. Goodnight, J.C. & M.L. Goodnight. 1974. Studies on the phalangid fauna of Trinidad. American Museum Novitates 1351:1-13, Kury, A.B. 2003. Annotated cataloue of the Lan- iatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologfa, monographic volume 1, pp. 5-337. Machado, G. 2002. Maternal care, defensive behav- ior, and sociality in neotropical Goniosoma har- vestmen (Arachnida: Opiliones). Insectes So- ciaux 49:388-393. Machado, G. & PS. Oliveira. 1998. Reproductive biology of the neotropical harvestman Gonioso- ma longipes (Arachnida, Opiliones, Gonylepti- dae): mating and oviposition behaviour, brood mortality, and parental care. Journal of Zoology 246:359-367. Machado, G. & PS. Oliveira. 2003. Maternal care in the neotropical harvestman Bourguyia albior- nata (Arachnida: Opiliones): oviposition site se- lection and egg protection. Behaviour 139:1509- 1524. Machado, G. & R.L.G. Raimundo. 2001. Parental investment and the evolution of subsocial behav- iour in harvestmen (Arachnida: Opiliones). Ethology, Ecology and Evolution 13:133-150. Pinto-da-Rocha, R. & A.B. Kury. 2003. Phyloge- netic analysis of Santinezia with description of five new species (Opiliones, Laniatores, Cranai- dae). Journal of Arachnology 31:173-208. Shear, W.A. 1982. Opiliones. In: Parker, S. P. (ed.) Synopsis and Classification of Living Organisms. New York, Mcgraw-Hill. 2 v. Tallamy, D.W. & W.P. Brown. 1999. Semelparity and the evolution of maternal care in insects. An- imal Behavior 57:727-730. Tallamy, D.W. & T.K. Wood. 1986. Convergence patterns in subsocial insects. Annual Review of Entomology 31:369-390. Wilson, E.O. 1971. The Insect Societies. Cam- bridge, Belknap Press. Manuscript received 3 September 2004, revised 17 June 2005. 2006. The Journal of Arachnology 34:273-278 SHORT COMMUNICATION FIRST UNEQUIVOCAL MERMITHID^-LINYPHIID (ARANEAE) PARASITE-HOST ASSOCIATION David Penney: Earth, Atmospheric and Environmental Sciences, The University of Manchester, Manchester, M13 9PL, UK. E-mail: david.penney@manchester.ac.uk Susan P* Bennett: Biological Sciences, Manchester Metropolitan University, Manchester, Ml 5GD, UK. ABSTRACT. The first description of a Mermithidae-Linyphiidae parasite-host association is presented. The nematode is preserved exiting the abdomen of the host, which is a juvenile Tenuiphantes species (Araneae, Linyphiidae), collected from the Isle of Mull, UK. An updated taxonomic list of known mer- mithid spider hosts is provided. The ecology of known spider hosts with regard to the direct and indirect life cycles of mermithid worms suggests that both occur in spiders. Keywords: Aranimermis, Isle of Mull, Linyphiidae, Mermithidae, Nematoda Nematode parasites of spiders are restricted to the family Mermithidae but are not uncommon (Poinar 1985, 1987) and were first reported almost two and a half centuries ago (Roesel 1761). How- ever, given the difficulty of identifying and rearing post-parasitic juvenile mermithids, they have re- ceived inadequate systematic treatment (Poinar 1985). In addition, the complete life history is known for only one species of these spider parasites (Poinar & Early 1990). Poinar & Welch (1981) sup- ported the use of the genus Agamomermis Stiles, 1903 for previously described mermithids that could not be placed in existing taxa and that were considered species inquirendae. This is the case for all spider mermithids described prior to 1986 (Poin- ar 1987). Currently three extant species of spider mermithid parasites are recognized: Aranimermis aptispicula Poinar & Benton 1986, A. actereki Gu- farov & An 1987 (spider host species unknown) and A. giganteus Poinar & Early 1990. In addition, the fossil species Heydenius araneus Poinar 2000 (a genus restricted to Tertiary fossil nematodes [Poinar 2003]) has been described from a crab spi- der (Thomisidae) in Baltic amber. Poinar (1985, 1987) provided lists of spider spe- cies with records of mermithid nematode parasit- ism. However, many of these taxa have now re- ceived taxonomic revisions and or transfers. In addition, a number of subsequent reports of para- sitism have been published (Gafurov & An 1987; Poinar & Early 1990; Camino & de- Villalobos 1998; Matsuda 1999; Poinar 2000; Allard & Rob- ertson 2003; lida & Hasegawa 2003; Vandergast & Roderick 2003; Ahtiainen et al. 2004). We provide an updated and taxonomically correct list in Table 1. Here we describe the first Mermithidae-Liny- phiidae parasite-host association and discuss the ecology of known spider hosts with regard to the life cycles of mermithid worms. This paper concerns three spider specimens, one with a worm in situ and two that are presumed to have been parasitized, but from which the worms have emerged and are lost. The specimens were col- lected during May 2004, in pitfall traps containing 50 ml of 70% alcohol, from a hazel forest in the Tireragan Estate on the Isle of Mull, UK. The spi- ders belong to the linyphiid genus Tenuiphantes but the female with the worm in situ cannot be identi- fied to species because it is a juvenile. In the spec- imen with the mermithid, the anterior and posterior regions of the worm have exited the abdominal cav- ity just anterior to the epigastric furrow and close to the pedicel (Eig. 1), but at least one coil can be observed interiorly, as a distortion beneath the ab- dominal integument, which is devoid of white gua- nine pigmentation. The worm is pale white/cream with a diameter of 0.13 mm and an approximate length (assuming only one coil exists in the spider) of 15.6 mm. The body length of the spider is 2.14 mm. The two other specimens are both female Ten- uiphantes tenebricola (Wider 1834) but no worms are visible, although the abdomens of both are se- verely damaged at the same point at which the worm is emerging in the other specimen. We con- sider it probable that both of these specimens were parasitized as well. One of the specimens has an emaciated, disk-shaped abdomen (similar to that of the specimen with the worm in situ), which gives 273 274 THE JOURNAL OF ARACHNOLOGY Table 1. — Spider hosts of mermithid worms: * A. aptispicula, ** A. giganteus, remainder species in- quire ndae. Family Species Reference Comments Agelenidae Agelenopsis oregonensis Cham- berlin & Ivie 1935 in Poinar (1987) Amaurobiidae Eurocoelotes inermis (L. Koch 1855) in Poinar (1987) as Coelotes i. Antrodiaetidae Atypoides riversi O.R-Cambridge 1833 in Poinar (1987) Anyphaenidae Wulfila albens (Hentz, 1847)* in Poinar (1987) as W. alba Araneidae Aculepeira ceropegia (Walckenaer 1802) in Poinar (1987) as Araneus ceropegius Araneidae Araneus diadematus Clerck 1757 in Poinar (1987) Araneidae Verrucosa arenata (Walckenaer 1842)* in Poinar (1987) Ctenidae Leptoctenus byrrhus Simon 1888 Poinar (2000) as Ctenus bryrrbus Cybaeidae Argyroneta aquatica (Clerck 1757) in Poinar (1987) Gnaphosidae Cesonia bilineata (Hentz 1847)* in Poinar (1987) Gnaphosidae Gnaphosa lucifuga (Walckenaer 1802) in Poinar (1987) Hexathelidae Porrhothele antipodiana (Wal- ckenaer 1837)** Poinar & Early (1990) as P. a. (Dipluridae) Idiopidae Misgolas borealis (Forster Poinar & Early as Cantuaria b. (Cteni- 1968)** (1990) zidae) Linyphiidae? Micryphantes bicuspidatus C.L. Koch 1838 in Poinar (1987) nomen dubium (Plat- nick 2004) Lycosidae Alopecosa inquilina (Clerck 1757) in Poinar (1987) as Tarentula i. Lycosidae Alopecosa trabalis (Clerck 1757) in Poinar (1987) as Lycosa vorax Lycosidae Arctosa alpigena (Doleschall 1852) Poinar (2000) possible mermithid Lycosidae Geolycosa patellonigra Wallace 1942 in Poinar (1987) Lycosidae Hygrolycosa rubrofasciata (Ohlert 1865) Ahtiainen et al. (2004) Lycosidae Pardosa agrestis (Westring 1861) in Poinar (1987) Lycosidae Pardosa amentata (Clerck 1757) in Poinar (1987) as Lycosa saccata Lycosidae Pardosa furcifera (Thorell 1875) in Poinar (1987) Lycosidae Pardosa glacialis (Thorell 1872) in Poinar (1987) Lycosidae Pardosa hortensis (Thorell 1872) in Poinar (1987) Lycosidae Pardosa lugubris (Walckenaer 1802) in Poinar (1987) Lycosidae Pardosa milvina (Hentz 1844) in Poinar (1987) as P. nigropalpis and P. scita Lycosidae Pardosa palustris (Linneaus 1758) in Poinar (1987) as Lycosa tarsal is Lycosidae Pardosa pseudoannulata (Boesen- berg & Strand 1906) lida & Hasegawa (2003) Lycosidae Pardosa riparia (C.L. Koch 1833) in Poinar (1987) Lycosidae Pardosa sphagnicola (Dahl 1908) in Poinar (1987) as Lycosa riparia s. Lycosidae Pardosa suwai Tanaka 1985 Matsuda (1999) Lycosidae Pardosa vancouveri Emerton 1917 in Poinar (1987) Lycosidae Rabidosa rabida (Walckenaer 1837) in Poinar (1987) as Lycosa scutulata Lycosidae Schizocosa saltatrix (Hentz 1844) in Poinar (1987) as Lycosa versimilis Lycosidae Sosippus floridanus Simon 1898 in Poinar (1987) Nemesiidae Stanwellia kaituna (Forster Poinar & Early as Aparua k. (Dipluri- 1968)** (1990) dae) PENNEY & BENNETT— MERMITHID-LINYPHIID PARASITE-HOST 275 Table 1. — Continued. Eamily Species Reference Comments Oxyopidae Oxyopes sertatus L. Koch 1877 Okochi (1969) Oxyopidae Peucetia viridans (Hentz 1832) in Poinar (1987) Philodromidae Tibellus oblongus (Walckenaer 1802) in Poinar (1987) Salticidae Habronattus signatus (Banks 1900) Vandergast & Roderick (2003) Salticidae Myrmarachne formicaria (De Geer 1778) in Poinar (1987) as Salticiis formicarius Salticidae Phidippus borealis Banks 1895 in Poinar (1987) Salticidae Phidippus clarus Keyserling 1885 in Poinar (1987) Salticidae Phidippus johnsoni (Peckham & Peckham 1883) in Poinar (1987) Salticidae Phidippus putnami (Peckham & Peckham 1883) in Poinar (1987) Salticidae Sitticus floricola palustris (Peck- ham & Peckham 1883) in Poinar (1987) as Sitticus p. Stiphidiidae Cambridgea foliata (L. Koch 1872) in Poinar (1987) Tetragnathidae Tetragnatha anuenue Gillespie 2002 Vandergast & Roderick (2003) Tetragnathidae Tetragnatha brevignatha Gillespie 1991 Vandergast & Roderick (2003) Tetragnathidae Tetragnatha praedonia L. Koch 1878 Okochi (1969) Tetragnathidae Tetragnatha quasimodo Gillespie 1991 Vandergast & Roderick (2003) Theridiidae Enoplognatha ovata (Clerck 1757) in Poinar (1987) as The rid ion ova turn and T redimitum Thomisidae Diaea dorsata (Eabricius 1777) in Poinar (1987) Thomisidae Misumenops tricuspidatus (Fabri- cius 1775) Okochi (1969) Thomisidae Xysticus deichmanni Soerensen 1898 in Poinar (1987) Thomisidae Xysticus durus (Soerensen 1898) in Poinar (1987) Thomisidae Xysticus funestus Keyserling 1880 in Poinar (1987) Zoridae Zora maculosa Roewer 1951 in Poinar (1987) as Z. maculata O.P.-C; nomen dubium (Plat- nick 2005) the impression of having been host to a worm. It has a normal degree of guanine pigmentation. The second specimen is not emaciated and has almost no guanine pigmentation. It cannot be ruled out that the specimens were damaged upon sorting the pit- fall trap contents, but as no other spiders (including many smaller species) were damaged, we consider this unlikely. Alternatively, they may have been hosts to non-mermithid parasites, such as acrocerids or phorids. Unfortunately, the pitfall contents are no longer available to check for emerged worms. It is probable that the specimen with the worm also be- longs to T. tenebricola, given that six other indi- viduals (three males and three females) were also collected at the same time from the same locality, whereas only one female of each of the following species was collected: T. alacris (Blackwall 1853), T. cristatus (Menge 1866) and T. mengei (Kulczyn- ski 1882). Poinar (1985, 1987) cited von Siebold (1848) as having identified an unknown mermithid worm in the spider Micryphantes bicuspidatiis (listed in Ta- ble 1 under Linyphiidae?). At the time of von Sie- bold’s paper (his description consisted of only five lines and no figures), only six spider families had been established, Linyphiidae was not erected until 1859. The genus Micryphantes C.L. Koch 1833 and M. bicuspidatus are both nomina dubia (Platnick 2005). Therefore, the new specimen described here represents the first described record of a mermithid- 276 THE JOURNAL OF ARACHNOLOGY 1 : Figure 1 . — The mermithid-carrying spider. Note the absence of abdominal guanine pigmentation, arrows point to the internal coil of the worm visible as a distortion in the integument. Scale lines =1.0 mm. linyphiid parasite-host association. Furthermore, the claims of Allard & Robertson (2003) of having identified mermithids in Pardosa milvina (Hentz) (Lycosidae) for the first time are unjustified, be- cause they were previously reported by Montgom- ery (1903) under the junior synonyms Pardosa ni- gropalpis Emerton 1885 and Pardosa scita Montgomery 1902. Being host to the worm carries a physiological cost for the spider in addition to its ultimate demise upon the emergence of the parasite. Infection signs generally start with a reduction or absence of di- gestive structures, and other organs may also be re- duced in extreme cases (Poinar 1985). It is inter- esting to note the reduced amount of guanine deposition in two of the above specimens, and in- deed the total absence of it in the specimen with the worm. Guanine is a crystalline purine excretory product, which accumulates in specialized, periph- eral cells of the digestive diverticula lying directly beneath the hypodermis (Oxford & Gillespie 1998) and has a white, blocky appearance. Although pat- terns of guanine deposition can vary intraspecifi- cally and throughout an individuals’ development (Oxford 1 998), there is usually a distinct pattern of guanine pigmentation present in Tenuiphantes Saar- isto & Tanasevitch 1996 specimens (DP pers. obs). In the case of mermithid-carrying individuals how- ever, the worm may be compromising the spider to such an extent, that the spider is receiving insuffi- cient nourishment to produce enough of these met- abolic waste products for pigmentation purposes. Further research would be required to confirm this hypothesis. At the time of emergence from the spider host, mermithid worms are mature third stage postpar- asitic juveniles. Thus, the individual described above cannot be identified to species because di- agnosis is based on adult characters. Mermithid life cycles are either direct or indirect. Direct life cycles are characterized by direct penetration of the spider host through the integument by the infective stage larva following emergence from the egg. Indirect life cycles involve a paratenic host (or a host in which significant development does not occur) in addition to the developmental (spider) host, which is infected by ingesting the infective stage of the parasite (Poinar 1985). The direct cycle is by far the most common among mermithids studied to date, however, it has been suggested that spider mermithids first undergo an indirect life cycle in- volving an aquatic paratenic host (Poinar 1987). Reasoning for this was based on observations of the life cycle of the spider mermithid Aranimermis ap- tispicula as follows: “adults were found in an aquatic habitat, whereas parasitized spiders were found in a variety of foraging habitats. Parasitized spiders were observed to enter the water and the nematodes were seen to emerge from the hosts’ bodies. These observations support an indirect type of cycle, but further studies are required to substan- tiate this” (Poinar 1987). Poinar (1987) did not rule PENNEY & BENNETT— MERMITHID-LINYPHIID PARASITE-HOST 277 out that some spider mermithid species may also have a direct life cycle. The life cycle of A. actereki presumably has an indirect life cycle because all worms studied (12 males, two females and four post-parasitic larvae) were collected from the bot- tom of a freshwater spring (Gufarov & An 1987). A. giganteus from New Zealand does have an in- direct life cycle involving aquatic invertebrates (Poinar & Early 1990). Given that most aquatic insect larvae have winged adults, the ecology of the spider hosts may provide insights that support or refute Poinar’s idea regarding the indirect life cycle of A. aptispicula. For example, winged, flying insects are more likely to be consumed by web spinning or foliage/flower hunting spiders than they are by non-web spinning, ground hunters or burrowing, sit and wait predators, which can be expected to feed primarily on non- flying, mainly fully terrestrial prey and are thus, more likely to be hosts to mermithids with a direct life cycle. Interestingly, the ecology of all but one of the spiders reported by Poinar & Benton (1986) as hosts of A, aptispicula (Gnaphosidae, Cesonia bilineata; Thomisidae, Misumenops sp. and Tmarus sp.; Salticidae, Phidippus sp.; Araneidae, Verrucosa arenata; Amaurobiidae, Wadotes sp. and Any- phaenidae, Wulfila albens), supports the idea of an indirect life cycle for this parasitic worm. The po- tential problem species of those listed is C bilinea- ta (Gnaphosidae). These are fast moving, agile hunters usually found under loose leaf litter at ground level and males are often found in pitfall traps. However, they have been collected by sweep- ing low vegetation and have also been collected from malaise traps (Platnick & Shadab 1980), which are primarily designed for catching flying in- sects. The large number of non-v/eb spinning, cursorial Lycosidae (Table 1; 38% of all species, excluding nomina dubia) with undescribed mermithids, sug- gests that a direct life cycle may be involved in some instances. Admittedly, some lycosids are common by freshwater, such as Hygrolycosa rub- rofasciata and Pardosa pseudoannulata (known to be mermithid hosts, see Table 1) and may be hosts to worms with indirect life cycles. However, lycosid genera such as Pirata and the pisaurid genus Do- lomedes, which are encountered almost exclusively near freshwater are unknown as mermithid hosts. The mermithid life cycle type in relation to the spi- der host Argyroneta aquatica also poses interesting questions, as this spider spends its entire life under water. Clearly, much work needs to be done before we can fully understand these interesting host-par- asite relationships, but a knowledge of the ecology of the host spiders can provide helpful clues in re- solving these. We thank G.O. Poinar Jr. (Oregon State Univer- sity) for his comments on the manuscript, J. Dunlop (Museum fiir Naturkunde, Berlin) for providing old German literature and D. Logunov (Manchester Museum) for translating a Russian paper. DP ac- knowledges a Leverhulme Trust grant to P. Selden and SB thanks L, Lace (Manchester Metropolitan University) for assistance and advice. LITERATURE CITED Ahtiainen, J.J., R.V. Alatalo, R. Kortet & M.J. Ran- tala. 2004. Sexual advertisement and immune function in arachnid species (Lycosidae). Behav- ioural Ecology 15:602-606. Allard, C. & M.W. Robertson. 2003. Nematode and dipteran endoparasites of the wolf spider Par- dosa milvina (Araneae, Lycosidae). Journal of Arachnology 31:139-141. Camino, N.B. & L.C. de Villalobos. 1998. First oc- currence of a mermithid (Nematoda: Mermithi- dae) parasitizing a spider (Arachnida: Araneida) in Argentina. Re vista de la Sociedad Entomolo- gica Argentina 57:6. Gafurov, A.K. & P.N. An. 1987, A new species of mermithid Aranimermis actereki sp. n. (Mermi- thidae, Nematoda) from the Kirghizia. Izvestiya Akademii Nauk Kirgizskoi SSR Khimiko Tekhn- ologicheskie Biologicheskie Nauki 1987:79-82, [In Russian]. lida, H. & H. Hasegawa. 2003. First record of a mermithid nematode emerging from the wolf spider Pardosa pseudoannulata (Araneae: Ly- cosidae). Acta Arachnologica 52:77-78. Matsuda, M. 1999. Two intersexual spiders of the family Lycosidae from Japan. Bulletin of the Hi- gashi Taisetsu Museum of Natural History 21:5- 54. [In Japanese]. Montgomery, TH. 1903. Studies on the habits of spiders, particularly those of the mating period. Proceedings of the Academy of Natural Sciences of Philadelphia 55:80-90. Okochi, T. 1969. Reports on parasites of spiders. Kishidaia 9:2-. [In Japanese]. Oxford, G.S. 1998. Guanine as a colorant in spi- ders: development, genetics, phylogenetics and ecology. Pp. 121-131. /« Proceedings of the 17th European Colloquium of Arachnology, Edin- burgh, 1997. (P.A. Selden, ed,). British Arach- nological Society, Bucks. Oxford, G.S. & R.G. Gillespie. 1998. Evolution and ecology of spider coloration. Annual Review of Entomology 43:619-643. Platnick, N.I. 2005. The world spider catalog, ver- sion 5.5. American Museum of Natural History, online at http://research.amnh.org/entomology/ spiders/catalog/INTRO 1 .html. Platnick, N.I. & M.U. Shadab. 1980. A revision of the spider genus Cesonia (Araneae, Gnaphosi- dae). Bulletin of the American Museum of Nat- ural History. 165:335-386. Poinar, G.O, Jr. 1985. Mermithid (Nematoda) par- 278 THE JOURNAL OF ARACHNOLOGY asites of spiders and harvestmen. Journal of Ar- achnology 13:121-128. Poinai; G.O. Jr. 1987. Nematode parasites of spi- ders. Pp. 299-308. In Ecophysiology of Spiders. (W. Nentwig, ed.). Springer Verlag, Heidelberg. Poinar, G.O. Jr. 2000. Heydenius araneus n. sp. (Nematoda: Mermithidae), a parasite of a fossil spider, with an examination of helminths from extant spiders (Arachnida: Araneae). Invertebrate Biology 119:388-393. Poinar, G.O. Jr. 2003. Trends in the evolution of insect parasitism by nematodes as inferred from fossil evidence. Journal of Nematology 35:129- 132. Poinar, G.O. Jr, & C.L.B. Benton Jr. 1986. Arani- mermis aptispicula n.g., n.sp. (Mermithidae: Nematoda), a parasite of spiders. Systematic Par- asitology 8:33-38. Poinar, G.O. Jr. & J.W. Early. 1990. Aranimermis giganteus n. sp. (Mermithidae: Nematoda), a par- asite of New Zealand mygalomorph spiders (Ar- aneae: Arachnida). Revue de Nematologie 13: 403-410. Poinar, G.O. Jr. & H.E. Welch. 1981. Parasites of invertebrates in the terrestrial environment. Pp. 947-954. In Review of Advances in Parasitolo- gy. (W. Slusarski, ed.). Polish Scientific Publi- cations, Warsaw. Roesel, A.J. 1761. Insectenbelustigung, Johann Jo- seph Fleishmann, Niimberg. Vandergast, A.G. & G.K. Roderick. 2003. Mermi- thid parasitism of Hawaiian Tetragnatha spiders in a fragmented landscape. Journal of Inverte- brate Pathology 84:128-136. von Siebold, C.TE. 1848. Ueber die Fadenwurmer der Insekten. Entomologische Zeitueg 9:290- 300. Manuscript received 16 November 2004, revised 21 April 2005, 2006. The Journal of Arachnoiogy 34:279-280 SHORT COMMUNICATION THREE HOMONYMOUS GENERIC NAMES IN ARANEAE AND OPILIONES Hiiseyin Ozdikmee: Department of Biology, Faculty of Science and Arts, University of Gazi, Ankara 06500, Turkey. E-mail: ozdikmen@gazi.edu.tr. Adriano Brilhante Kury: Dept. Invertebrados, Museu Nacional, Ueiversidade Federal do Rio de Janeiro, Quinta da Boa Vista ^/n, Sao Cristovao 20.940-040, Rio de Janeiro, RJ, Brazil ABSTRACT. Three junior homonyms were detected among the Arachnida and the following replace- ment names are proposed; Neoarminda for Arminda Roewer 1949 (Opiliones); Alpazia for Lapazia Roewer 1949 (Opiliones); and Araneotanna for Tanna Berland 1938 (Araneae). Accordingly, three new combi- nations are herein proposed for the respective type species. All three genera are monotypic. Keywords: Araneae, Opiliones, homonymy, replacement names, Bolivia, Brazil, New Hebrides While recently researching the “Nomenclator Zoologicus” (Neave 1939-1950) three homony- mous arachnid generic names were found. As far as it could be ascertained from our sources (Kury 2003 for the Opiliones and Platnick 2004 for the Ara- neae), all these homonymies have been hitherto un- detected. The opportunity is here taken to provide replacement names for them in accordance with the International Code of Zoological Nomenclature (1999). Order Opiliones Family Gonyleptidae Genus Neoarminda NEW NAME Remarks. — The name Arminda Roewer 1949 was proposed as a monotypic genus of Opiliones in the Phalangodidae, Tricommatinae (Roewer 1949b: 144) for the species Phalangodella colatinae Soares & Soares 1946 from Brazil. The generic name Ar- minda is preoccupied by Arminda Krauss 1892 (Or- thoptera, Caelifera, Catantopidae) (Krauss 1892: 168), Therefore, Neoarminda NEW NAME is here proposed as a replacement name. The follov/ieg new combination is established: Neoarminda cola- tinae (Soares & Soares 1946), NEW COMBINA- TION. Gonyleptoidea incertae sedis Genus Alpazia NEW NAME Remarks, — The genus Lapazia Roewer 1949 was proposed as another monotypic genus of Opi- liones in the Phalangodidae Phaiangodinae (Roewer 1949a: 14) for the species Lapazia minima Roewer 1949 from Bolivia. Kury (2003:25) removed this genus from the Phalangodidae and left it as incertae sedis. The generic name Lapazia is preoccupied by Lapazia Ferris 1937 (Insecta, Homoptera, Diaspi- didae) (Ferris 1937:68). Consequently, Alpazia NEW NAME is here proposed as a replacement name. The following new combination is estab- lished: Alpazia minima (Roewer 1949), NEW COMBINATION. Order Araneae Family Salticidae Genus Araneotanna NEW NAME Remarks. — The genus Tanna Berland 1938 was proposed as a monotypic genus of Araneae in the Salticidae for the species Tanna ornatipes Berland 1938 from the New Hebrides (Berland 1938:141). The generic name is preoccupied by Tanna Distant 1905 (Insecta, Homoptera, Cicadidae) (Distant 1905:61). Therefore, Araneotanna NEW NAME is here proposed as a replacement name. The follow- ing new combination is established: Araneotanna ornatipes (Berland 1938), NEW COMBINATION. We wish to thank Max Moulds (Australian Mu- seum, Sidney) and Penny Gullan (University of California at Davis) for their assistance in locating some of the entomological literature, and the editor Mark Harvey for substantially revising the manu- script. LITERATURE CITED Berland, L. 1938. Araignees des Nouvelles Hebri- des, Annales de la Societe Entomologique de France 107:121-190. 279 280 THE JOURNAL OF ARACHNOLOGY Distant, W.L. 1905. Rhynchotal notes, XXIX. An- nals and Magazine of Natural History, (7)15:58- 70. Ferris, G.F. 1937. Atlas of the Scale Insects of North America. Stanford University Press, Palo Alto, California, Series 1, Vol. 1:68. International Code of Zoological Nomenclature. 1999, The International Trust for Zoological No- menclature, London. Fourth edition. Krauss, H.A. 1892 [1891]. Systematisches Ver- zeichniss der canarischen Dermapteren und Or- thopteren mit Diagnosen der neuen Gattungen und Arten. Zoologischer Anzeiger 15:163-171. Kury, A.B. 2003. Annotated catalogue of the Lam iatores of the New World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Zaragoza, vol. especial monogrMco, n° 1:1-337. Neave, S.A. 1939-1950. Nomenclator Zoologicus. The Zoological Society of London, London. Platnick, N.I. 2004. The world spider catalog, ver- sion 5.0, American Museum of Natural History, online at http://research.amnh.org/entomology/ spiders/catalog/index . html Roewer, C.F. 1949a,. Uber Phalangodiden 1. Subfam. Phalangodinae, Tricommatinae, Samoinae. Wei- tere Weberknechte XIII. Senckenbergiana, Frankfurt 30:11-61. Roewer, C.F. 1949b. Einige neue Gattungen der Phalangodidae. Veroffentlichungen aus dem Mu- seum fiir Natur--, Vdlker- u. Handelskunde in Bremen, Reihe A: Naturwissenschaften 1:143- 144, Manuscript received 20 July 2004, revised 13 March 2005. 2006. The Journal of Arachnology 34:281-284 OBITUARY LE TEMPS MARCHE SI VITE— IN MEMORY OF KONRAD THALER Christoph Muster: Universitat Leipzig, Institut fiir Biologic II, Molekulare Evolution und Systematik der Tiere, TalstraBe 33, D-04103 Leipzig, Germany, E-mail: muster@rz.uni-leipzig.de Jason A. Dunlop: Institut fiir Systematische Zoologie, Museum fiir Naturkunde der Humboldt-Universitat zu Berlin, InvalidenstraBe 43, D- 101 15 Berlin, Germany. It is hard in the moment of sorrow to mea- sure the degree of loss, but European arach- nologists must come to terms with the passing of one of their most influential figures. On the June, 2005, Konrad Thaler died suddenly and unexpectedly at the age of 64 during a student excursion in the Stubaier Alps. With him we have lost someone who has left his mark on a whole generation of zoogeogra- phers, taxonomists, mountain ecologists and entomologists and who was described at his funeral by a long-time friend as a scientist of “enthusiastic heart and rational words”. Konrad Thaler was born on December 19'*’, 1940 in Innsbruck, Austria and stayed true to his Tyrolean mountains throughout his life. 281 282 THE JOURNAL OF ARACHNOLOGY After attending school in Innsbruck, he re- ceived his leaving certificate in 1958, spent two years in military service and began his studies in zoology and botany at the Univer- sity of Innsbruck in 1959/60. His professors, H. Janetschek, O. Steinbock, H. Gams and W. Larcher, were important figures in the study of Alpine biogeography. His 1967 dissertation was (in translation): “On the spider fauna of Northern Tyrol (excluding Linyphiidae and Micryphantidae. Prelude to a catalog of the large spiders of North Tyrol)”. Subsequently, linyphiids would become one of his favorite groups. Field-experience and much material for future revisions was gathered in the six years he spent at the Alpine Research Station in Obergurgl, before taking on an assistant post at the University of Innsbruck in 1970. He submitted his 1978 ‘Habiiitation' thesis on “The taxonomy and zoogeography of Alpine spiders” and since 1983 led the department of Terrestrial Ecology and Taxonomy at the In- stitute of Zoology and Limnology of Inns- bruck University. He was a council member of the “Centre International de Documenta- tion Arachnologique” (CIDA) from 1986- 1989, CIDA (later ISA) correspondent for Austria, and President of the Austrian Ento- mological Society from 2002-2005. Konrad Thaler died at the peak of his pro- ductivity. Until the very end he worked tire- lessly each day, almost as if he knew how little time he had left. The bare facts are clear: between 1963 and 2005 he authored or co- authored more than 220 journal articles. There was a continual increase in his yearly output: on average one a year during his time in Ob- ergurgl (1964-1970), three as a university As- sistant (1970-1978), and seven a year since his 'Habilitation' in 1978. Since 2002 alone he published 40 papers! Additionally, there were popular science articles, often in the journal of the Austrian Alpine Society, plus abstracts and book reviews. From 1973-2005 he supervised 41 diploma theses and 10 PhDs; mostly faunistic and ecological, or taxonomic and morphological projects. As well as arach- nids, he supervised numerous studies of myr- iapods and beetles. Thanks to his careful re- cord-keeping, we know he gave exactly 100 presentations at Austrian and international meetings; the last four days before his death on “Areal forms of invertebrates in the east- ern Alps”. It is hard to pick out individual research highlights. His work on the arachnids near Lunz in Austria (Thaler 1963), published when he was only 23, remains of great value as the first, and until recently, the only record of males of the parthenogenetic harvestman Megabunus lessertL Characteristic would be the serial publications “Uber wenig bekanete Zwergspineen aus dee Alpen”, which was published over nine issues; as well as “Frag- menta Faunistica Tiroleesia”. Here, Konrad attempted, as part of partial inventory of the North-Tyrol fauna, to make what little was known about the less-familiar invertebrate groups successively accessible; thus impres- sively demonstrating the breadth of his knowl- edge. The reprints of part 17 (Thaler 2005) were posted on the day of his death. In his last years he was particularly keen to produce summary works, such as the faunistic synop- sis of North-Tyrol spiders (Thaler 1998) and a review of the ecology of high- Alpine species (Thaler 2003). Also important was his edito- rial work on the “Diversity and biology of spiders, scorpions and other arachnids” which included papers from long-term collaborators and showed Austria as a working environment for arachnologists (Thaler 2004). For a full measure of the merit of his life’s work one should compare our state of knowledge at the end of his studies in the “Contributions to the spider fauna of North-Tyrol” (Thaler 1992, 1994, 1995, 1997a, b, 1999) with how things were before he started, when the spiders were . an unhappy picture of insufficient fau- nistic research.” (translated from Holdhaus 1954). A full inventory of Austrian spiders (begun by Thaler & Knoflach 2002, 2003, 2004) was sadly not to be completed in his lifetime. Konrad’s taxonomic work included the au- thorship of two genera (Carnielia, Mysmeni- ola), 11 species and one subspecies of spider, and one harvestman species. Of these, 48 he collected himself, and none have so far proved to be synonyms. His new taxa spanned 17 families, predominantly Linyphiidae (42 spe- cies) and Amaurobiidae (12 species), with a geographical concentration in the Alps and the Mediterranean. Twenty-six species from vari- ous animal groups bear his name, including twelve spiders, four flies, a tardigrade and an oligochaete worm. It’s obvious that such productivity could MUSTER & DUNLOP—THALER OBIT 283 only be achieved through great personal and passionate commitment to research. For Kon-= rad, science was his life-work {labor vincit omnia). Insiders knew one could in, variably meet Konrad in the institute seven days a week, so long as he wasn’t on excursion. He had the good fortune with his second wife Barbara to find an equally enthusiastic and tal- ented comrade-in-arms. Their years together were a particularly productive phase of co- operative activity, during which the Mediter- ranean arachnids became a further focus of research. Although he enjoyed considerable interna- tional recognition, his achievements were not always recognized by his owe institute. Here, he was often accused of failing to keep up with the latest trends or buzz- words. It is not that he rejected, for example, molecular meth- ods, but simply felt that . the state of knowledge achievable by AonventionaT means was far from being reached. . . In- deed it was through conventional methods that Konrad became a leading figure of 20* cen- tury arachnology. His death means, regretta- bly, a further substantial loss of taxonomic ex- pertise among the German-language universities. It can only be hoped that those in authority recognize the consequences of this before it is too late. Everyone who visited Konrad in his office was impressed by the concentration of litera- ture, in particular the many originals of stan- dard works and a rich collection of compara- tive material. They were also astounded by their host’s memory. Konrad could recognize almost every Central European spider, without the use of literature, and when he said ‘T ha- ven’t seen anything like that before.” you knew you had found something special. But most of all, people remember his courtesy and helpfulness, his stimulating inquisitiveness, constant ability to enthuse and his many words of encouragement. As an example, be- tween two stressful meetings he was asked to check the identification of a Troglohyphantes male and got up from the microscope with the words “Thanks for the nice view”. No one left his room without a better understanding, a constructive thought, or feeling more moti- vated. He always had an open door for his students and it is no accident that shortly after his death many of them offered thanks on the university homepage for his remarkable per- sonal contact and the enthusiasm and devotion he brought to his teaching. Administrative duties meant that despite his discipline and industry, time for research be- came increasingly scarce. He often wrote of a “Mountain of paper in front of the micro- scope.” and the “Lure of the mountains for arachnological collecting”. At the 8* meeting of the German- speaking arachnologists in Salzburg, he mused about whether we should go into the Alps, simply to enjoy the distinc- tive fauna or the landscape per se. For Kon- rad, life without the mountains was impossible to imagine. He felt happiest at 3,000 m; where the motto might have been: concentrate on that which is most important. Longer collect- ing trips were made to the Caucuses, Pyrenees and Atlas mountains, where the local guides were said to have whispered “he marches like an Arab”. His student trips into the Alps were legendary, and up to the very end he would be walking way ahead of the younger partic- ipants, especially on critical passes. The ex- cursions took place in all weathers, often with the resume that this enabled one to better un- derstand the requirements of Alpine animals. It seems fate that Konrad died during his final regular student excursion, only a few months before he was due to retire, when he would have had more time for fieldwork and his own projects. His friends, colleagues and students must now take over his legacy and try to . write at least one new line each day”. For advice and information we are very grateful to Barbara Knoflach. A German ver- sion of this obituary has been published in the Arachnoiogische Mitteilungee 30 (2005), which also includes a complete bibliography of Konrad Thaler’s publications by this date. We thank the editors for permission to offer this translation. LITERATURE CITED Holdhaus, K. 1954. Die Spuren der Eiszeit in der Tierwelt Europas. Abhandlungen der zoologisch- botanischen Gesellschaft in Wien 18:1-493. Thaler, K. 1963. Spinnentiere aus Lunz (Niederos- terreich) nebst Bemerkungen zu einigen von Kul- czynski aus Niederosterreich gemeldeten Arten. Berichte des naterwisseeschaftlich-medizinisch- en Vereins in Innsbruck 53:273-283. Thaler, K. 1992. Beitrage zur Spinnenfauna von Nordtirol — 1. Revidierende Diskussion der “Ar- achniden Tirols” (Anton Ausserer 1867) und 284 THE JOURNAL OF ARACHNOLOGY Schrifttum. Veroffentlichungee des Museum Fer- dinandeum (Innsbruck) 7 1(1991): 155-189. Thaler, K. 1994. Beitrage zur Spinnenfauna von Nordtirol — 2: Orthognathe, cribellate und hap- logyne Familien, Pholcidae, Zodariidae, Mime- tidae und Argiopiformia (ohne Linyphiidae s.L) (Arachnida: Araneida). Mit Bemerkuegen zur Spinnenfauna der Ostalpen. Veroffentlichungen des Museum Ferdinandeum (Innsbruck) 73 (1993):69-119. Thaler, K. 1995. Beitrage zur Spinnenfauna von Nordtirol — 5. Linyphiidae 1: Linyphiinae (sensu Wiehle) (Arachnida: Araneida). Berichte des na- turwissenschaftlich-medizinischen Vereins in Innsbruck 82:153-190. Thaler, K. 1997a. Beitrage zur Spinnenfauna von Nordtirol — 3: “Lycosaeformia” (Agelenidae, Hahniidae, Argyronetidae, Pisauridae, Oxyopi- dae, Lycosidae) und Gnaphosidae (Arachnida: Araneida). Veroffentlichungen des Museum Fer~ dinandeum (Innsbruck) 75/76(1 995/96):97-146. Thaler, K. 1997b. Beitrage zur Spinnenfauna von Nordtirol — 4. Dionycha (Anyphaenidae, Clu™ bionidae, Heteropodidae, Liocranidae, Philod- romidae, Salticidae, Thomisidae, Zoridae). Ver- offentlichungen des Museum. Ferdinandeum (Innsbruck) 77:233-285. Thaler, K. 1998. Die Spinnen von Nordtirol (Arach- nida, Araneae): Faunistische Synopsis. Veroffen- tlichungee des Museum Ferdinandeum (Inns- bruck) 78:37-58. Thaler, K. 1999. Beitrage zur Spinnenfauna von Nordtirol — 6. Linyphiidae 2: Erigoninae (sensu Wiehle) (Arachnida: Araneae). Veroffentlichun- gen des Museum Ferdinandeum (Innsbruck) 79: 215-264. Thaler, K. 2003. The diversity of high altitude arachnids (Araneae, Opiliones, Pseudoscorpi- ones) in the Alps. Pp 281-296. In L. Nagy, G. Grabherr, C. Korner & D.B.A. Thompson (eds.), Alpine Biodiversity in Europe. Ecological Stud- ies 167. Springer, Berlin, Heidelberg. Thaler, K. (ed.) 2004. Diversitat und Biologic von Webspinnen, Skorpionen und anderen Spinnen- tieren. Denisia 12:1-586. Thaler, K. 2005. Fragmenta Faunistica Tirolensia — 17 (Arachnida: Araneae; Insecta: Psocoptera, Strepsiptera, Megaloptera, Neuroptera, Raphi- dioptera, Mecoptera, Siphonaptera, Diptera: My- cetophiloidea). Veroffentlichungen des Tiroler Landesmuseum Ferdinandeum 84:161-180. Thaler, K. & B. Knofiach. 2002. Zur Faunistik der Spinnen (Araneae) von Osterreich: Atypidae, Haplogynae, Eresidae, Zodariidae, Mimetidae. Linzer biologische Beitrage 34:413-444. Thaler, K. & B. Knoflach. 2003. Zur Faunistik der Spinnen (Araneae) von Osterreich: Orbiculariae p.p. (Araneidae, Tetragnathidae, Theridiosomati- dae, Uloboridae). Linzer biologische Beitrage 35:613-655. Thaler, K. & B. Knoflach. 2004. Zur Faunistik der Spinnen (Araneae) von Osterreich: Gnaphosidae, Thomisidae (Dionycha pro parte). Linzer biolo- gische Beitrage 36:417-484. INSTRUCTIONS TO AUTHORS (revised July 2006) General: Manuscripts are accepted in English only Authors whose primary language is not English may consult the editors for assistance in obtaining help with manuscript preparation. All manuscripts should be prepared in general accordance with the current edition of the Council of Biological Editors Style Manual unless instructed otherwise below. Authors are advised to consult a recent issue of the Journal of Aracknology for additional points of style. Manuscripts longer than three printed journal pages should be prepared as Feature Articles, shorter papers as Short Communications. One invited Review Article per year will be solicited by the editors and published in the third issue at the discretion of the editors. 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I'Ai'i'i ;Ju.! •'..fiJJ' ’oilar- • Wi i45i#(N„,= _- /• ' t. . , ■• T.. ■fSiastef .a IJ_ jn n i.--, }• sct/Trti'f-i b.r uii J-i • :tl"C Ki'] tn ;£;?Tmi» ' ’• am '*• -11 ^ . w. ii^-ii •* ■ 'u.v|f-Cii‘i ■ < *'<» -".f «t '' Mainosa, a new genus for the Australian ‘shuttlecock wolf spider’ (Araneae, Lycosidae) by Volker W. Framenau 206 Ecology of Thestylus aurantiurus of the Parque Estadual da Serra da Cantareira, Sao Paulo, Brazil (Scorpiones, Bothriuridae) by Humberto Y. Yamaguti & Ricardo Pinto-da-Rocha 214 Observations on Loxosceles reclusa (Araneae, Sicariidae) feeding on short-horned grasshoppers by Jennifer Parks, William V. Stoecker & Charles Kristensen 221 The systematic position of the Amazonian species of Albiorix (Pseudoscorpiones, Ideoroncidae) by Mark S. Harvey & Volker Mahnert 227 Short Communications Additional notes on the post-birth development of the scorpion Vaejovis coahuilae Williams (Vaejovidae) by W. David Sissom, Kari J. McWest & Anne L. Wheeler 231 Variations in web construction in Leucauge ventusa (Araneae, Tetragnathidae) by Yann Henaut, Jose Alvaro Garcia-Ballinas & Claude Alauzet 234 Nest site fidelity of Paraphidippus aurantia (Salticidae) by Kailen A. Mooney & JonR.Haloin 241 A new Mastophora from Argentina and the male of Mastophora vaquera (Araneae, Araneidae) by Herbert W. Levi 244 A replacement name for Iracema Perez-Miles 2000 (Araneae, Theraphosidae) by Fernando Perez-Miles 247 An extremely low genetic divergence across the range of Euscorpius italicus (Scorpiones, Euscorpiidae) by Victor Fet, Benjamin Gantenbein, Ay§egul Karataf & Ahmet Karata§ 248 Dispersal by Ummidia spiderlings (Araneae, Ctenizidae): ancient roots of aerial webs and orientation? by William G. Eberhard 254 Regurgitation among penultimate juveniles in the subsocial spider Anelosimus cf. studiosus (Theridiidae): are males favored? by Carmen Viera, Soledad Ghione & Fernando G. Costa 258 Activity of juvenile tarantulas in and around the maternal burrow by Cara Shillington & Brian McEwen 261 Types of shelter sites used by the giant whipscorpion Mastigoproctus giganteus (Arachnida, Uropygi) in a habitat characterized by hard adobe soils by Fred Punzo 266 First case of maternal care in the family Cranaidae (Opiliones, Laniatores) by Glauco Machado & Joseph Warfel 269 First unequivocal mermithid-linyphiid (Araneae) parasite-host association by David Penney & Susan P. Bennett 273 Three homonymous generic names in Araneae and Opiliones by Hiiseyin Ozdikmen & Adriano Brilhante Kury 279 Obituary Le temps marche si vite — In memory of Konrad Thaler by Christoph Muster & Jason A. Dunlop 281 USERNAME: akron05 PASSWORD: spider05 CONTENTS The Journal of Arachnology Volume 34 Featured Articles Number 1 The wolf spiders of artesian springs in arid South Australia, with a revalidation of Tetmlycosa (Araneae, Lycosidae) by Volker W. Framenau, Travis B. Gotch & Andrew D. Austin 1 The prey of a lithophilous crab s^idiQx Xysticus loeffleri (Araneae, Thomisidae) by Elchin Fizuli oglu Guseinov 37 First species of Hesperopilio (Opiliones, Caddoidea, Caddidae) from South America by Jeffrey W. Shultz & Tomas Cekalovic 46 Role of the anterior lateral eyes of the wolf spider Lycosa tarantula (Araneae, Lycosidae) during path integration by Joaquin Ortega-Escobar 51 An examination of agonistic interactions in the whip spider Phrynus mar^inemaculatus (Arachnida, Amblypygi) by Kasey D. Fowler-Finn & Eileen A. Hebets 62 Four new crab spiders from Taiwan (Araneae, Thomisidae) by Jun-Xia Zhang, Ming-Sheng Zhu & I-Min Tso 77 A review of the linyphiid spider genus Solenysa (Araneae, Linyphiidae) by Lihong Tu & Shuqiang Li 87 Spider size and guarding of offspring affect Paraphidippus aurantius (Araneae, Salticidae) response to predation threat by Kailen A. Mooney & Jon R. Haloin 98 Spider diversity in coffee plantations with different management in southeast Mexico by Miguel Angel Pinkus Rendon, Guillermo Ibarra-Niinez, Victor Parra-Tabla, Jose Alvaro Garcia-Ballinas & Yann Henaut 104 Systematics of the Afro-Macaronesian spider genus Sancus (Araneae, Tetragnathidae) by Matijaz Kuntner & Fernando Alvarez-Padilla 113 Three new species of Pholcus (Araneae, Pholcidae) from the Canary Islands with notes on the genus Pholcus in the archipelago by Dimitar Dimitrov & Carles Ribera 126 A new species of Cupiennius (Araneae, Ctenidae) coexisting with Cupiennius salei in a Mexican mangrove forest by Francisco J. Medina Soriano 135 Have you seen my mate? Description of unknown sexes of some North American species of Linyphiidae and Theridiidae (Araneae) by Nadine Dupere, Pierre Paquin & Donald J. Buckle 142 Capture efficiency and preservation attributes of different fluids in pitfall traps by Martin H. Schmidt, Yann Clough, Wenke Schulz, Anne Westphalen & Teja Tscharntke 159 Description and ecology of a new solifuge from Brazilian Amazonia (Arachnida, Solifugae, Mummuciidae) by Lincoln S. Rocha & Martinho C. Carvalho 163 Two new purse- web spiders of the genus Atypus (Araneae, Atypidae) from Korea by Seung-Tae Kim, Hun-Sung Kim, Myung-Pyo Jung, Joon-Ho Lee & Joon Namkung 170 Copulatory behavior and web of Indicoblemma lannaianum from Thailand (Arachnida, Araneae, Tetrablemmidae) by Matthias Burger, Alain Jacob & Christian Kropf 176 Prey choice by Nesticodes rufipes (Araneae, Theridiidae) on Musca domestica (Diptera, Muscidae) and Dermestes ater (Coleoptera, Dermestidae) by Marcelo N. Rossi & Wesley A.C. Godoy 186 A review of pholcid spiders from Tibet, China (Araneae, Pholcidae) by Feng Zhang, Ming-Sheng Zhu & Da-Xiang Song 194 3 9088 01277 3164 Contents continued on inside back cover