Himalmartensus, a new genus of the spider family Amaurobiidae from Nepal (Araneae).
Griswold (1990) addressed the higher level taxonomic question of the familial relationships of Phyxelidinae by emphasizing four character systems: the nature of the cribellum; the presence, form, number, and position of processes on the male palpal tibia; the pattern of tarsal and metatarsal trichobothria; and the form of the tracheal system. In examining those structures, we found that the striated texture of the small trichobothrial hood, plus the presence of PMS paracribellar spigots and PLS modified spigot (amaurobiid PLS spigot of Wang 2000), are unique and define Amaurobius (the type genus of the family) and related genera including, at least, Callobius Chamberlin 1947, Pimus Chamberlin 1947 and Taira Lehtinen 1967. This "true Amaurobiidae" group has been tested phylogenetically by Griswold et al. (2005). As a result, the family Amaurobiidae includes at least the above genera, and more are likely to be included with more detailed analysis that is beyond the scope of this study.
Wu et. al (2002) analyzed 12S rRNA gene sequences using 2 amaurobiid species, Coelotes plancyi Simon 1880 and Tamgrinia tibetana (Hu & Li 1987) and 2 agelenid species of the genus Agelena and concluded that Coelotes is more closely related to Tamgrinia than to Agelena. However, in another study (Bi et al. 2005) that used 18S and 28S rRNA in an attempt to resolve the phylogenetic position of Coelotinae, four taxa from two amaurobiid and two agelenid genera (e.g., Draconarius, Coelotes, Agelena, and Alloagelena) were used and results indicated that Coelotes + Draconarius are more closely related to Agelena + Alloagelena than to Tamgrinia, suggesting (Tamgrinia((Coelotes, Draconarius)(Agelena, Alloagelena))), which is in conflict with the findings of Wu et al. (2002). More recently, the analyses by Spagna & Gillespie (2008) using molecular data of non-orb-weaving spiders suggested a sister group relationship between Coelotinae and Ageleninae, which is consistent with the conclusion of Bi et al. (2005). Spagna & Gillespie (2008) found that Agelenidae (including Ageleninae and Coelotinae) could be the sister to Hahniidae + Cybaeidae related taxa. In addition to agelenids and coelotines, Spagna & Gillespie also selected four taxa for analyses from three "true amaurobiids" genera (e.g., Amaurobius, Callobius, and Pimus) and found none of them was closely related to Coelotinae. Unfortunately, other lineages of current Amaurobiidae species were not sampled by Spagna & Gillespie (2008) (e.g., Rubrius and Macrobunus related species from South America, the Holarctic Arctobius, and Tamgrinia from the Himalayan region) and their phylogenetic placements, either related to Agelenidae or to Amaurobiidae, still need further investigation. The synapomorphies of current Agelenidae (including Coelotinae) are still unknown, but some may be found in their spinneret structures. Both Ageleninae and Coelotinae are ecribellate spiders and build similar funnel-shaped webs (and also the cribellate amaurobiid genus Tamgrinia) (Wang 2002, 2003), although the webs of Coelotinae appear to be much smaller. It seems likely that species of the cribellate Tamgrinia are also related to agelenids and coelotines.
When we first examined the three amaurobiid species collected from Nepal and included in this paper, their generic placement was puzzling. After eight years of a failed search for male specimens in hopes of obtaining additional characters to support their generic status, we decided to publish the material on hand as a new genus, Himalmartensus, based only on females. Hopefully males will be found in the future based on our published female information. Even without male palpal characters, the differences between Himalmartensus new genus and other current amaurobiid and agelenid members are obvious. We defined this new genus as a member of the family Amaurobiidae because of its similarity to amaurobiid Rubrius and Macrobunus, despite the large geographic distance between them. Of course, the phylogenetic placement of Rubrius and Macrobunus related amaurobiids needs to be evaluated further. Of the similar genera (Table 1), the new genus Himalmartensus is similar to Rubrius and some Macrobunus species by having a single hairy colulus (Figs. 9, 27, 29), rather than a cribellum found in "true amaurobiids," Tamgrinia, Arctobius, and some Macrobunus, or two patches of setae as in the subfamily Coelotinae, Ageleninae, and the family Cybaeidae. Another similarity between Himalmartensus new genus and Rubrius is the smooth large hood of the trichobothrial base (Figs. 12, 26, 51), being either longitudinally or transversely striated in other studied taxa. But the four simple tracheal tubes in Himalmartensus new genus (Fig. 36) differ from the strongly branched tracheal tubes in Rubrius. While in Cybaeus jilinensis (Song et al. 1993), there are only two tracheal tubes, which are strongly branched (Fig. 61). In addition, the female epigynum of Himalmartensus new genus is modified with long and looping copulatory ducts. Similar coiled copulatory ducts are observed in coelotine lutulentus-group species and also in some Cybaeidae species, for example, Cybaeina minuta (Banks 1906), but the species of Himalmartensus new genus show no evidence of complex spermathecal pore structures as in Cybaeidae. According to Bennett (1992), the complex spermathecal pore structures are also absent in Coelotinae and other amaurobiids.
All measurements are in millimeters. Unless indicated otherwise, all scale bars are 0.2 mm length. Legs are not measured. Spinnerets, trichobothria, and tarsal organs are examined using SEM. Other photos are taken from the Olympus Stereo Scope eyepiece using a Nikon Coolpix 4500 camera. Prior to SEM examination, the specimens were either air-dried or critical point dried and coated. Tracheal tubes were examined using Griswold's (1990) method. The spigot names used in the text and figures follow Coddington (1989) and Griswold (1990). The distribution map was generated using GIS ArcView software and the .dbf files of the studied species are downloadable from http://www.amaurobiidae. com, which is published and maintained by Xin-Ping Wang. More photos of the type specimens included in this paper can be viewed from the website http://www.ChineseSpecies.com which was created and maintained by Shu-Qiang Li and XinPing Wang.
The types are deposited in the Senckenberg Museum, Frankfurt, Germany (SMF). Abbreviations: AC = aciniform spigots; ALE = anterior lateral eyes; ALS = anterior lateral spinneret; AME = anterior median eyes; CY = cylindrical spigots; mAP = minor ampullate spigots; MAP = major ampullate spigots; PI = piriform spigots; PLE = posterior lateral eyes; PLS = posterior lateral spinneret; PME = posterior median eyes; PMS = posterior median spinneret.
Family Amaurobiidae Thorell 1870
Himalmartensus Wang & Zhu new genus
Type species.--Himalmartensus martensi Wang & Zhu new species.
Other species.--Himalmartensus ausobskyi Wang & Zhu new species and H. nepalensis Wang & Zhu new species.
Etymology.--The genus is named in honor of Jochen Martens (Mainz, Germany) for his contribution of amaurobiid specimens that were collected from his Himalayan expeditions. These specimens included a new coelotine genus (Himalcoelotes Wang 2002) and 10 species of this genus (Wang 2002). Another 38 coelotines, including 36 new species, have been recognized from Martens material and will be published soon. The gender is masculine.
Diagnosis.--Himalmartensus new genus can be diagnosed from other amaurobiids, agelenids, and genera of other related families by at least: (1) AMS represented by colulus as in Rubrius and some Macrobunus species; (2) single chilum, as in Rubrius, and Amaurobius-related genera; (3) smooth trichobothrial base, as in Rubrius and agelenids, all others being either longitudinally or transversely striated, and (4) four simple tracheal tubes, but branched tubes in Cybaeus, Rubrius and Macrobunus. In addition, both the promargin and retromargin of Himalmartensus new genus have 5-8 teeth and the epigynum is modified with long and looping copulatory ducts.
Description.--Females: Medium size ecribellate spiders, with total length 8-10 mm. Carapace elongate, reddish brown, slightly narrowed in ocular area, sparsely covered with short, long black setae; few strong setae on clypeus, ocular area, and middle carapace; longitudinal fovea moderately depressed. Legs moderately long. Abdomen dark brown, with dark maculation, heavily covered with short setae (Figs. 3, 14, 15, 23, 33, 37, 45). Spinnerets short. From dorsal view, anterior eye row slightly procurved, posterior row procurved; eye sizes and arrangements: AME smallest, ALE and PLE largest and subequal, PME larger than AME, AME-AME separated by approximately AME diameter, AME-ALE widely separated by approximately 1-1.5 AME diameter, ALE-PLE separated by approximately their radius, PME-PLE and PME-PME distinctly separated by approximately 1.5-2 times PME diameter, AME-PME widely separated by at least 2 times AME diameter (Figs. 6, 16, 42). Clypeus height 1.5-2 times AME diameter, covered with long, strong setae; chilum undivided, hairless (Figs. 6, 16). Chelicerae with 6-7 promarginal teeth, the basal ones largest, and 5-8 retromarginal teeth, with basal large and distal small; condyle large; dorsal chelicerae covered with long setae, with strongly elevated base; chelicerae ventrally flat, with proximal short setae and inner long, fine setae; fangs moderately long; anterior face of chelicerae covered with dense, long, strong setae (Figs. 22, 44). Endites elongated, with anterior scopula and linear serrula. Labium longer than wide, slightly notched distally. Sternum shield-shaped, sparsely covered with long dark setae, heavily sclerotized (Figs. 7, 17, 43). Legs medium length, I, IV longest, almost subequal, leg III shortest; trochanters not notched; tibiae with about four rows of trichobothria; metatarsi and tarsi with one row of trichobothria; trichobothria with both large and small hoods smooth, not striated (Figs. 12, 26, 51). Tarsal organ with simple opening (Figs. 13, 28, 52). Tarsi with three claws, superior claws with 8-10 teeth; scopulae absent; leg spination often varies among individuals, typical leg spination pattern: femur: I p0-0-2, d1-1-0; II p0-0-1, d1-1 0; III p0-0-1, d1-1-0; IV d1-1-0; tibia: I p0-0-1, v2-2-2; II p0-0-1, v2-2-2; III p0-0-1, v1-2-2; IV d1-0-0; r1-1-0, v1-12; metatarsus: I v2-2-2; II v2-2-2; III p0-1-1, r0-1-1, v2-2-2; IV d0-0-1, v1-1-2. Tracheal tubes simple, limited to abdomen; spiracle situated close to spinnerets and connected to relatively narrow atrium from which two lateral and two median tubes arise (Fig. 36). Colulus present, covered with hairs (Figs. 9, 27); ALS short, apex with 2 major ampullate gland spigots (MAP) and 41-55 piriform gland spigots (PI); PMS small, with 2 minor ampullate gland spigots (mAP), 2 aciniform gland spigots (AC), and 1-3 cylindrical gland spigots (CY); PLS second segment short, with approximately 6-17 aciniform gland spigots, and 2 cylindrical gland spigots (Figs. 8-11, 29-32, 47-50). Epigynum simple, atrium small, situated close to epigastric furrow; copulatory ducts long, with 3-7 loops around spermathecae; spermathecae with bases relatively large, widely separated, anteriorly converging with spermathecal heads almost touching each other (Figs. 1, 2, 4, 5, 18-21, 24, 34, 35, 38-41, 46).
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Distribution.--Nepal (Fig. 53).
Himalmartensus ausobskyi Wang & Zhu new species Figs. 1-13, 53
Type specimens.--NEPAL: holotype female, Dholakha District, Jiri valley, elev. 2600-3000 m, oak forest, 86[degrees]14'E, 27[degrees]37'N, 16 January 1970, J. Martens (SMF, #44); 1 female paratype, Lalitpur District, Phulchoki Mt., foot-hills near Godavari, elev. 1770 m, 85[degrees]23'E, 27[degrees]36'N, 19 March 1980. J. Martens & A. Ausobsky (SMF, #105).
Etymology.--The specific name is after Albert Ausobsky, one of the collectors of the type specimens.
Diagnosis.--H ausobskyi resembles H nepalensis in having long spermathecal stalks (short in H martensi), with long copulatory ducts that loop around spermathecal stalks at least four times, and the narrow, less sclerotized posterior plate of atrium. The spermathecal stalks of this species converge gradually while extending anteriorly (Figs. 2, 5), rather than converging immediately as in H nepalensis.
Description.--Holotype female: Total length 11.3. Carapace 4.80 long, 3.60 wide. Abdomen 6.50 long, 4.90 wide (Fig. 3). Eye sizes and interdistances: AME and PME approximately same size, ALE largest, PLE slightly smaller than ALE (AME 0.16, ALE 0.26, PME 0.17, PLE 0.20); AME separated from each other by 2/3 of its diameter, from ALE by slightly more than AME diameter, from PME by approximately 1.5 times AME diameter; PME and PLE widely separated (AME-AME 0.10, AME-ALE 0.21, AME-PME 0.25, PME-PME 0.31, PME-PLE 0.34) (Fig. 6). Chelicerae with 6 promarginal, and 8 retromarginal teeth. Apex of ALS with 2 major ampullate gland spigots (MAP) and approximately 44 piriform gland spigots (PI); PMS with 2 minor ampullate gland spigots (mAP), 2 aciniform gland spigots (AC), 2 cylindrical gland spigots (CY); PLS with approximately 6 aciniform gland spigots, and 2 cylindrical gland spigots (Figs. 8-11). Epigynum with small atrium; atrium with narrow, less sclerotized posterior plate; copulatory ducts originate posteriorly between the spermathecal bases, extending anteriorly, each looping around the long and converging spermathecal stalks at least four times; spermathecae with large bases separated by approximately their width; spermathecal stalks extend anteriorly, converging gradually, with distal ends close together (Figs 1, 2, 4, 5).
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Distribution.--Nepal (Dholakha, Lalitpur) (Fig. 53).
Himalmartensus martensi Wang & Zhu new species Figs. 14-36, 53
Type specimens.--NEPAL: holotype female, Kathmandu District, Kathmandu valley, Balaju Park, elev. 1400 m, 85[degrees]17'E, 27[degrees]44'N, 1 May 1973, J. Martens (SMF, #36); 1 female paratype, Lamjung District, Marsyandi valley, between Tal and Dharapani, forest remnants in gorge, elev. 1580-1850 m, 84[degrees]21'E, 28[degrees]17'N, 12 April 1980. J. Martens & A. Ausobsky (SMF, #136); 1 female paratype, Ilan District, Mai Pokhari, elev. 2100-2200 m, Castganopsis forest remnants, 87[degrees]55' E, 26[degrees]58' N, 9-10 April 1988, J. Martens & W. Schawaller (SMF, #319, tracheal tubes examined); 1 female paratype, Myagdi District, southern Dhaulagiri range, Bobang S of Dhorpatan, elev. 2500 m, 26 April-1 May 1970, J. Martens (SMF, #3); 1 female paratype, Makawanpur District, Mahabarat Mts., Daman, elev. 2500-2900 m, 22-25 February 1970, J. Martgens (SMF, #33).
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Etymology.--This new species is named in honor of Jochen Martens who collected the specimens used in this and other studies.
Diagnosis.--Compared to the more than four copulatory duct loops in H ausobskyi and H nepalensis, this new species has only 3 loops around the relatively short spermathecal stalk (Figs. 19, 21, 35). In addition, the atrium of this new species has a broad, highly sclerotized plate (Figs. 18, 20, 24, 34).
Description.--Holotype female: Total length 11.3. Carapace 3.10 long, 2.30 wide. Abdomen 5.20 long, 3.40 wide (Figs. 14, 15). Eye sizes and interdistances: AME and PME subequal, ALE largest, PLE slightly smaller than ALE (AME 0.13, PME 0.14, ALE 0.18, PLE 0.16); AME separated from each other by 2/3 of its diameter, from ALE by about AME diameter, from PME by approximately 1.5 times AME diameter; posterior eyes widely separated (AME-AME 0.08, AMEALE 0.13, AME-PME 0.19, PME-PME 0.23, PME-PLE 0.22) (Fig. 16). Chelicerae with 6 promarginal, and 6 retromarginal teeth (Fig. 22). Apex of ALS with 2 major ampullate gland spigots (MAP) and approximately 55 piriform gland spigots (PI); PMS with 2 minor ampullate gland spigots (mAP), 2 aciniform gland spigots (AC), and 2 cylindrical gland spigots (CY); PLS with approximately 10 aciniform gland spigots, and 2 cylindrical gland spigots (Figs. 29-32). Epigynum with small atrium; atrium with broad, highly sclerotized posterior plate; copulatory ducts originate posteriorly between the spermathecal bases, extend anteriorly and loop around the converging spermathecal stalks 3 times; spermathecae with large bases that are separated by approximately their width; spermathecal stalks extend anteriorly and converge gradually, with distal ends close together (Figs. 1821).
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Distribution.--Nepal (Kathmandu, Lamjung, Ilan) (Fig. 53).
Himalmartensus nepalensis Wang & Zhu new species Figs. 37-53
Type specimens.--NEPAL: holotype female, Rasuwa District, Trisuli Valley, Gosainkund, mixed forest, elev. 24002600 m, 85[degrees]23'E, 28[degrees]8'N, 23 April 1973 (SMF, #38); 1 female paratype, Rasuwa District, Trisuli Valley, Gosainkund, moist forest in gorge, elev. 1000-2000 m, 85[degrees]19'E, 28[degrees]8'N, 23 June? 1973, J. Martens (SMF, #38A); 1 female paratype, Rasuwa District, Trisuli Valley, between Ramche and Dhunche, elev. 1800-2000 m, 85[degrees]14'E, 28[degrees]5'N, 22 April 1973, J. Martens (SMF, #37).
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Etymology.--The specific name refers to the type locality of the species, Nepal.
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Diagnosis.--H. nepalensis is similar to H. ausobskyi in having long spermathecal stalks and copulatory ducts which loop at least 5 times, and the narrow, less sclerotized posterior plate of atrium. The spermathecal stalks of this new species converge immediately and then extend anteriorly (Figs. 39, 41), rather than converge gradually as in H. ausobskyi.
Description.--Holotype female: Total length 11.3. Carapace 4.20 long, 3.40 wide. Abdomen 4.60 long, 3.20 wide (Fig. 37). Eye sizes and interdistances: AME smallest, PME slightly larger than AME, ALE and PLE about the same size (AME 0.11, ALE 0.20, PME 0.15, PLE 0.18); AME separated from each other by less than its diameter, from ALE and PME by about 1.5 times AME diameter; posterior eyes are widely separated (AME-AME 0.08, AME-ALE 0.17, AME-PME 0.17, PME-PME 0.23, PME-PLE 0.26) (Fig. 42). Chelicerae with 7 promarginal and 5 retromarginal teeth (Fig. 44). ALS with 2 major ampullate gland spigots (MAP) and approximately 41 piriform gland spigots (PI); PMS with 2 minor ampullate gland spigots (mAP), 2 aciniform gland spigots (AC), and 1 cylindrical gland spigot (CY); PLS with approximately 17 aciniform gland spigots and 2 cylindrical gland spigots (Figs. 47-50). Epigynum with small atrium having a narrow, weakly sclerotized posterior plate; copulatory ducts originate posteriorly between the spermathecal bases, extend anteriorly and loop around the long and converging spermathecal stalks at least 6 times; spermathecae with large bases separated by approximately their width; spermathecal stalks extend anteriorly and converge gradually, with anterior ends contiguous (Figs. 38-41).
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Distribution.--Nepal (Rasuwa) (Fig. 53).
This research is the result of the Himalaya Expeditions of J. Martens, no. 257. For no. 256, see: Acta Arachnologica 56:1519, 2007. J. Martens was sponsored by Deutscher Akademischer Austauschdienst, Deutsche Forschungsgemeinschaft and Feldbausch Foundation, Mainz University.
We thank J. Martens (Institute of Zoology, Mainz University, Germany) for providing the specimens used in this study and for helping create the distribution map. J. Martens, C.E. Griswold (California Academy of Sciences, San Francisco), R.G. Bennett (British Columbia Ministry of Forests, Victoria, Canada) and S.Q. Li (Institute of Zoology, Chinese Academy of Sciences, Beijing) kindly read the manuscript and gave valuable comments. Z.S. Zhang (Southwest University, Chongqing, China) helped with agelenid SEM photos and the illustration of Cybaeus tracheal tubes. We like to thank I. Agnarsson (University of Akron, Ohio) and two anonymous reviewers for their comments. The first author thanks the American Museum of Natural History (New York, N.I. Platnick), the California Academy of Sciences (San Francisco, C.E. Griswold) and the Florida State Collection of Arthropods (Gainesville, G.B. Edwards) for the use of their facilities.
Manuscript received 10 December 2007, revised 16 March 2008.
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Xin-Ping Wang and Ming-Sheng Zhu: College of Life Sciences, Hebei University, Baoding 071002, China. E-mail: email@example.com; firstname.lastname@example.org
Table 1.--Comparison of eight selected characters of twelve representative species from ten Amaurobiidae genera: Amaurobius, Callobius, Taira, Tamgrinia, Arctobius, Pimus, Macrobunus, Rubrius, Himalmartensus, and Coelotes, Agelenidae genus Agelena, and Cybaeidae genus Cybaeus. Characters Amaurobius Callobius Taira fenestralis bennetti sp. AMS cribellum cribellum cribellum Cribellum divided divided divided PMS present present present paracribellar spigots PLS modified present present present spigot (amaurobiid PLS spigot of Wang 2000) Chilum single single single Trichobothria transverse transverse transverse base, large hood striations Trichobothria striated striated striated base, small hood striations Tracheal four simple four simple four middle tubes tubes simple tubes (Griswold tubes et al. 2005) Characters Tamgrinia Arctobius Pimus laticeps ageleneides pitus AMS cribellum cribellum cribellum (Griswold et al. 2005) Cribellum divided divided -- PMS absent absent present paracribellar spigots PLS modified absent absent present spigot (amaurobiid PLS spigot of Wang 2000) Chilum paired paired ? Trichobothria transverse transverse transverse base, large hood striations Trichobothria smooth smooth striated base, small hood striations Tracheal four simple ? four simple middle tubes tubes tubes (Griswold et al. 2005: Pimus sp.) Characters Macrobunus Rubrius Himalmartensus multidentatus antarcticus martensi (Figs. 14-36) AMS colulus colulus colulus (Griswold et al. 2005) Cribellum -- -- -- PMS absent absent absent paracribellar spigots PLS modified absent absent absent spigot (amaurobiid PLS spigot of Wang 2000) Chilum absent single single Trichobothria longitudinal smooth or smooth base, large finely hood striated striations Trichobothria smooth smooth smooth base, small hood striations Tracheal four tubes, four tubes, four simple middle with median with tubes four tubes tubes slightly median branched tubes (Griswold et strongly al. 2005) branched Characters Coelotes Cybaeus Agelena atropos spp labyrinthica (Wang 2002) (Figs. 60, 61) (Fig. 58) AMS setae setae setae (Roth (Bennett & Brame 1988) 1972) Cribellum -- -- -- PMS absent absent absent paracribellar spigots PLS modified absent absent absent spigot (amaurobiid PLS spigot of Wang 2000) Chilum paired paired (Ihara paired & Nojima 2004) Trichobothria transverse smooth or smooth base, large finely hood striated striations longi- tudinally Trichobothria smooth smooth smooth base, small hood striations Tracheal four two strongly four simple middle simple branched tubes tubes tubes tubes
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|Author:||Wang, Xin-Ping; Zhu, Ming-Sheng|
|Publication:||The Journal of Arachnology|
|Date:||May 1, 2008|
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