Journal of Natural History A review of Coelotinae epigynal teeth

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A review of Coelotinae epigynal teeth morphology, with descriptions oftwo species from China (Araneae: Amaurobiidae)Xin-Ping Wang a;Peter Jäger b

a College of Life Sciences, Hebei University, Baoding, Hebei, China b Arachnology, SenckenbergResearch Institute, Frankfurt am Main, Germany

Online publication date: 23 April 2010

To cite this Article Wang, Xin-Ping andJäger, Peter(2010) 'A review of Coelotinae epigynal teeth morphology, withdescriptions of two species from China (Araneae: Amaurobiidae)', Journal of Natural History, 44: 19, 1165 — 1187To link to this Article: DOI: 10.1080/00222931003632708URL: http://dx.doi.org/10.1080/00222931003632708

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Journal of Natural HistoryVol. 44, Nos. 19–20, May 2010, 1165–1187

ISSN 0022-2933 print/ISSN 1464-5262 online© 2010 Taylor & FrancisDOI: 10.1080/00222931003632708http://www.informaworld.com

TNAH0022-29331464-5262Journal of Natural History, Vol. 0, No. 0, Feb 2010: pp. 0–0Journal of Natural HistoryA review of Coelotinae epigynal teeth morphology, with descriptions of two species from China (Araneae: Amaurobiidae)Journal of Natural HistoryX.-P. Wang and P. JägerXin-Ping Wanga* and Peter Jägerb

aCollege of Life Sciences, Hebei University, Baoding, Hebei, China; bArachnology, Senckenberg Research Institute, Frankfurt am Main, Germany

(Received 3 June 2009; final version received 18 January 2010)

The morphology of the epigynal teeth of the spider subfamily Coelotinae is studied.Of the 460 coelotine species, 413 are known from females (261 have both sexesdescribed and 152 have only females described) but six of them have incompleteepigynum data. Among the 407 species studied, six characters have been recog-nized that might be informative for the estimation of Coelotinae phylogeny, i.e.epigynal teeth: presence/absence, number, length, position on epigynum, separa-tion from each other, and width. Their congruence with the current coelotinegenera is discussed. Two coelotine species from Yunnan, China are described:Draconarius wrasei sp. nov. (female) and Draconarius immensus Xu and Li 2006(both sexes). Both species are similar to the incertus group species in having abifurcate conductor, simple median apophysis, prolaterally originated embolus inmale, and in having a similar epigynum, absence of epigynal teeth, and short,more or less anteriorly extended spermathecae.

Keywords: Araneae; Coelotinae; female genitalia; systematics; Yunnan Province

Introduction

Spiders of the subfamily Coelotinae F.O. Pickard-Cambridge, 1893 are widespreadfrom Europe and Asia to eastern North America (Wang 2002, 2009; Platnick 2009)and have been the subject of considerable interest in recent years, particularly thosefrom East and Southeast Asia. A total of 31 coelotine species have been discovered in2008, with the total number of species increased from 429 (Wang and Jäger 2008) tothe current 460 (Platnick 2009; Wang 2009), and most of them (62%) were assigned totwo of the 22 genera, Draconarius Ovtchinnikov, 1999 and Coelotes Blackwall, 1841(Table 1). However, Wang (2003) and Wang and Jäger (2007) indicated that manyspecies in these two genera are not congeneric with any of the existing coelotine gen-era and are only temporarily assigned to either Draconarius or Coelotes. Genericassignments of many species in Coelotinae, as well as the phylogenetic analysis oftheir species relationships, are difficult because of the high degree of morphologicalvariation and a lack of comparative study of their genitalia. Wang and Jäger (2008)dealt with the male palpal embolus morphology and provided the starting point forsuch studies in the future. In addition to the embolus in the male palp, the subfamilyCoelotinae is also characterized by epigynal teeth, which arise from the ventral sideof the epigynum. The possible informative characters related to the epigynal teethinclude their presence/absence, number, length, position on epigynum, separation

*Corresponding author. Email: [email protected]

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1166 X.-P. Wang and P. Jäger

from each other and width. In this study, the epigynal teeth of all coelotine speciesare compared. Two species of the genus Draconarius, one of them a new species, arealso described from Zhongdian County, Yunnan, China.

Materials and methods

Measurements are in millimetres. Scale lines in habitus photos represent a rulerdivided into 1-mm increments; scale bars on all other illustrations represent0.2 mm. Eye diameters are taken at the widest point. The lengths of body, prosomaand opisthosoma do not include the length of the chelicerae or spinnerets. Termi-nology used in the text and figures follows Wang (2002). Specimens described in thecurrent paper are deposited in the Senckenberg Museum, Frankfurt (SMF). Epigynalteeth morphology of most Coelotinae species is based upon specimens examined bythe authors; the rest of the species are judged from the published illustrations. Thedistribution map was generated using GIS ARCVIEW software and data of the stud-ied species are downloadable from Wang (2009). More type specimen photos of thespecies included in this paper can be viewed from Li and Wang (2009). Abbrevia-tions used in the text are: AME, anterior median eyes; ALE, anterior lateral eyes;PME, posterior median eyes; PLE, posterior lateral eyes; RTA, retrolateral tibialapophysis.

Table 1. Coelotinae genera and number of species (all datafrom Platnick 2009 and Wang 2009).

Genus No. of species Percentage

Alloclubionoides 24 5.22%Bifidocoelotes 2 0.43%Coelotes 141 30.65%Coras 15 3.26%Draconarius 145 31.52%Eurocoelotes 11 2.39%Femoracoelotes 2 0.43%Himalcoelotes 11 2.39%Iwogumoa 17 3.70%Leptocoelotes 2 0.43%Lineacoelotes 5 1.09%Longicoelotes 3 0.65%Notiocoelotes 5 1.09%Orumcekia (= Coronilla) 8 1.74%Pireneitega 20 4.35%Platocoelotes 14 3.04%Robusticoelotes 2 0.43%Spiricoelotes 3 0.65%Tegecoelotes 6 1.30%Tonsilla 8 1.74%Urocoras 5 1.09%Wadotes 11 2.39%

Total 460 100.00%


Journal of Natural History 1167

Coelotinae epigynal teeth morphology

The presence of epigynal teeth in female coelotine spiders is a synapomorphy of aclade that includes 15 genera (lost in Platocoelotes Wang, 2002 and SpiricoelotesWang, 2002), according to Wang (2002). We define the epigynal teeth as the structurethat arises from the ventral surface of the epigynum. Most of the early workers onlyprovided illustrations but lacked a detailed description of the epigynal teeth (andother genital structures). The epigynal teeth have been described as tubercles (forCoras by Muma 1946), scape (for Wadotes by Muma 1947 and Bennett 1987), spurs(Paik et al. 1969), projections (Paik 1971, Shimojana 2000), processes (Shimojana1982), pricks (Deltshev 1990), teeth or epigynal teeth (Blauwe 1973; Brignoli 1982;Wang et al. 1990; Peng and Wang 1997; Wang 2002). The term epigynal teeth hasbeen used in recent years consistently (Wang 2003; Dankittipakul and Wang 2004;Ovtchinnikov and Inayatullah 2005; Xu et al. 2006, 2008; Xu and Li 2007). The func-tions of the epigynal teeth may be correlated to functional aspects during copulation,e.g. anchoring of male structures. If involved in a functional unit it is suggested thatabsence, presence, size, distance, position etc. are most likely more similar in closelyrelated groups, as male and female structures are adapted as co-evolving traits andtherefore cannot be changed separately or dramatically, but gradually (Jäger 2006).Of the 460 coelotine species, 413 (90%) have females described (261 of them haveboth sexes described and 152 have only females described) but the following sixspecies have incomplete epigynum data: Coelotes cavicola (Komatsu, 1961), Coelotesobesus Simon, 1875, Coelotes simoni Strand, 1907, Coelotes simplex O. Pickard-Cambridge, 1885, Coelotes stylifer Caporiacco, 1935 and Coelotes tegenarioidesO. Pickard-Cambridge, 1885. As a result, 407 species are dealt with in this study. Ourstudy on the epigynal teeth found the following six characters, which could be thevaluable sources for phylogenetic signals for future Coelotinae matrices.

Character one: presence/absence of epigynal teeth

Presence of epigynal teeth is a synapomorphy of node 4 species in the Coelotinae clado-gram of Wang (2002). Of the 407 studied Coelotinae, 290 of them (71%) have epigynalteeth and 117 (29%) do not have epigynal teeth (Table 2). Absence of epigynal teeth isshared by all species of eight genera, i.e. Alloclubionoides Paik, 1992, FemoracoelotesWang, 2002, Longicoelotes Wang, 2002, Notiocoelotes Wang, Li and Xu, 2008, Plato-coelotes Wang, 2002, Orumcekia Koçak and Kemal, 2008 (= Coronilla Wang, 1994,according to Koçak and Kemal 2008), Robusticoelotes Wang, 2002 and SpiricoelotesWang, 2002.

Fifteen species, which have no epigynal teeth, are not congeneric with their cur-rent genus Coelotes, and their correct generic placement is still unknown. Three otherspecies with no epigynal teeth might also be misplaced in their current genera, i.e.Iwogumoa xieae Liu and Li, 2008, Leptocoelotes edentulus (Wang and Ono, 1998) andPireneitega bidens (Caporiacco, 1935). Iwogumoa species exhibit the epigynal teethexcept I. xieae Liu and Li, 2008, but the presence of a conductor dorsal apophysis inthe I. xieae male palp proves that it is not congeneric to Iwogumoa. Both Leptocoe-lotes species have a weakly sclerotized epigynum. The anteriorly situated epigynalhoods and the posteriorly originated copulatory ducts in L. edentulus differ from thelaterally situated epigynal hoods and the anteriorly originated copulatory ducts in the



Table 2. Character one: epigynal teeth, presence/absence. Character two: epigynal teeth,number (all data from Wang 2009).

Epigynal teeth, presence/absence, number

Genus No. of species

Remarks

Absent (n = 117) Alloclubionoides* 23Coelotes 15 All 15 species are not congeneric with Coelotes, they

are: C. amplilamnis Saito, 1936; C. galeiformis Wang et al., 1990; C. iheyaensis Shimojana, 2000; C. inabaensis Arita, 1974; C. introhamatus Xu and Li, 2006; C. luculli Brignoli, 1978; C. maculatus Zhang, Peng and Kim, 1997; C. ningmingensis Peng et al., 1998; C. noctulus Wang et al., 1990; C. samaksanensis Namkung, 2002; C. satoi Nishikawa, 2003; C. septus Wang et al., 1990; C. tiantongensis Zhang, Peng and Kim, 1997; C. titaniacus Brignoli, 1977; C. uenoi Yamaguchi and Yaginuma, 1971

Coronilla* 7Draconarius 41Femoracoelotes* 2Iwogumoa 1

I. xieae Liu and Li, 2008 is not congeneric with Iwogumoa

Leptocoelotes 1 Le. edentulus (Wang and Ono, 1998) is not congeneric with Leptocoelotes

Longicoelotes* 3Notiocoelotes* 4Pireneitega 1 P. bidens (Caporiacco, 1935) is not congeneric

with PireneitegaPlatocoelotes* 13Robusticoelotes* 2Spiricoelotes* 3

One (n = 11) Bifidocoelotes* 2Coelotes 1 C. wangi Chen and Zhao, 1997 is not congeneric

with CoelotesWadotes* 8

Two (n = 279) Coelotes 109Coras* 15Draconarius 77Eurocoelotes* 10Himalcoelotes* 11Iwogumoa* 15Leptocoelotes 1Lineacoelotes* 5Pireneitega 19Tegecoelotes* 6Tonsilla* 7Urocoras* 5

Total 407

Note: The asterisk indicates that all species of the genus have this character state. For a list of spe-cies included in each category, see Wang 2009.



type species L. pseudoluniformis (Zhang, Peng and Kim, 1997). Male L. edentulus isunknown. Judging from the female epigynum and vulva, L. edentulus is not conge-neric with Leptocoelotes. Species of Pireneitega have moderately long epigynal teeth.Not only lacking the epigynal teeth, P. bidens also shows a different atrium and adifferent vulva and as a consequence is not congeneric with Pireneitega.

Character two: number of epigynal teeth

The presence of a single epigynal tooth is a synapomorphy for Bifidocoelotes Wang,2002 from Hong Kong and Taiwan, and Wadotes Chamberlin, 1925 from NorthAmerica (Table 2). The epigynal tooth of Bifidocoelotes is relatively slender,distinctly bifurcate, arises at level of anterior atrial margin (Figure 1A), while thetooth of Wadotes is broad, may be slightly bifurcate in some species (Bennett 1987),arises at a level anterior of the atrium, and is distinctly separated from the atrium slit(Figure 1B).

Coelotes wangi Chen and Zhao, 1997, with only the female described, has a singleepigynal tooth (Figure 1C) and is misplaced in its current genus. It has a small AME(half the size of ALE), two retromarginal cheliceral teeth, and a single broad, shortepigynal tooth arising anteriorly and distinctly separated from the small, mediallysituated atrium (Chen and Zhao 1997; Wang and Jäger 2007). Its correct genericplacement is unknown.

Character three: length of epigynal teeth

We define the length of epigynal teeth as long, which is more than three times longerthan wide (Figure 1A,B,D–F), or short, which is as long as wide, or slightly longerthan wide (Table 3). An intermediate condition between long and short did exist(called “more or less long” here) and could fall into the ambiguity code meaning thatthis observation is uncertain (Figure 1G,H). In most Coelotinae, the epigynal teethare short. Approximately 24 species have the epigynal teeth that are defined as “moreor less long” and 55 species as long. Species of nine genera are consistent in length ofepigynal teeth, i.e. long in Bifidocoelotes, Himalcoelotes, Urocoras, Wadotes, short inCoras, Iwogumoa, Leptocoelotes, Tegecoelotes, and with intermediate length inLineacoelotes. Among the 13 Coelotes that have long epigynal teeth, six of them fromEastern Europe and Middle Asia belong to the Coelotes charitonovi species group(Wang and Zhu 2009) but others from East Asia are apparently not congeneric withCoelotes. The four Draconarius from China and Thailand with long epigynal teethcould also belong to other genera, they are: D. huizhunesis (Wang and Xu, 1988),D. strophadatus (Zhu and Wang, 1991), D. acidentatus (Peng and Yin, 1998),D. elatus (Dankittipakul and Wang, 2004). Eurocoelotes, Pireneitega and Tonsillashow high levels of variation in length of epigynal teeth.

Character four: position of epigynal teeth

Position of epigynal teeth could be a highly homoplasious character and is difficult tomeasure due to the presence of an intermediate condition. This character defines theepigynal teeth by their longitudinal position relative to the position of the atrium, butthe position and size of the atrium itself might vary in many coelotine species (Table 4).



The epigynal teeth may arise anteriorly (at a level anterior to atrium, with their basesdistinctly separated from anterior atrial margin) (Figures 1B–D, 2A–C), medially (atabout the same level as anterior atrial margin) (Figures 1A,E,G–I, 2D,F), or posteri-orly (at a level posterior to anterior atrial margin) (Figures 1F, 2E,G–I). Approxi-mately 27 species have the teeth arising at a level slightly anterior of the atrium andnine others slightly posterior of the anterior atrial margin. They could fall into theambiguity code. Species of six genera are consistent in their position of epigynal

Figure 1. Epigynal tooth morphology. (A) Bifidocoelotes primus (Fox, 1937): single, long,strongly bifurcate, arises at level of anterior atrial margin. (B) Wadotes dixiensis Chamberlin,1925: single, more or less broad, arises at level anteriorly of atrium. (C) Coelotes wangi Chenand Zhao, 1997: single, more or less broad, arises at level anteriorly of atrium. (D) Coelotesbrachiatus Wang et al., 1990: long, arise at level anteriorly of atrium, separated by less than anatrial width. (E) Pireneitega spinivulva (Simon, 1880): long, arise at level of anterior atrial mar-gin, separated by an atrial width. (F) Eurocoelotes jurinitschi (Drensky, 1915): long, arise atlevel posteriorly of anterior atrial margin, separated by more than an atrial width. (G) Tonsillatruculenta Wang and Yin, 1992: more or less long, contiguous, arise at level of anterior atrialmargin. (H) Lineacoelotes longicephalus Xu, Li and Wang, 2008: more or less long, arise atlevel of anterior atrial margin, separated by less than an atrial width. (I) Tegecoelotes ignotus(Bösenberg and Strand, 1906): short, broad, arise at level of anterior atrial margin, separatedby less than an atrial width.



Table 3. Character three: epigynal teeth, length (all data from Wang 2009).

Epigynal teeth, length


Species list

Long (n = 55) Bifidocoelotes* 2 AllCoelotes 13 C. caudatus de Blauwe, 1971; C. arganoi Brignoli,

1978; C. coenobita Brignoli, 1978; C. vignai Brignoli, 1978; C. musashiensis Nishikawa, 1989; C. brachiatus Wang et al., 1990; C. nenilini Ovtchinnikov, 1999; C. turkestanicus Ovtchinnikov, 1999; C. insulanus Shimojana, 2000; C. kumensis Shimojana, 2000; C. chenzhou Zhang and Yin, 2001; C. pedodentalis Zhang, Zhu, Sun and Song, 2006; C. hiradoensis Okumura and Ono, 2006

Draconarius 4 D. huizhunesis (Wang and Xu, 1988); D. strophadatus (Zhu and Wang, 1991); D. acidentatus (Peng and Yin, 1998); D. elatus Dankittipakul and Wang, 2004

Eurocoelotes 1 E. jurinitschi (Drensky, 1915)Himalcoelotes* 11 AllPireneitega 10 P. involuta (Wang et al., 1990); P. major

(Kroneberg, 1875); P. luctuosus (L. Koch, 1878); P. spinivulva (Simon, 1880); P. taishanensis (Wang et al., 1990); P. luniformis (Zhu and Wang, 1994); P. triglochinata (Zhu and Wang, 1994); P. neglecta (Hu, 2001); P. xinping (Zhang, Zhu and Song, 2002); P. liansui (Bao and Yin, 2004)

Tonsilla 1 T. imitata Wang and Yin, 1992Urocoras* 5 AllWadotes* 8 All

More or less long (n = 24)

Coelotes 6 C. exitialis Koch, 1878; C. modestus Simon, 1880; C. mastrucatus Wang et al., 1990; C. saccatus Peng and Yin, 1998; C. kumejimanus Shimojana, 2000; C. tokaraensis Shimojana, 2000

Draconarius 2 D. yichengensis Wang, 2003; D. tongi Xu and Li, 2007

Eurocoelotes 3 E. falciger (Kulczynski, 1897); E. karlinskii (Kulczynski, 1906); E. deltshevi (Dinitrov, 1996)

Iwogumoa 1 I. pengi (Ovtchinnikov, 1999)Lineacoelotes* 4 AllPireneitega 5 P. armeniaca (Brignoli, 1978); P. cottarellii

(Brignoli, 1978); P. fedotovi (Charitonov, 1946); P. spasskyi (Charitonov, 1946)

Tonsilla 3 T. defossa Xu and Li, 2006; T. eburniformis Wang and Yin, 1992; T. truculenta (Wang and Yin, 1992)

(Continued)



teeth, i.e. arising anteriorly at a level anterior to atrium in Wadotes, medially at aboutthe same level as anterior atrial margin in Bifidocoelotes, Coras, Leptocoelotes,Tegecoelotes, or posteriorly at a level posterior to anterior atrial margin in Eurocoe-lotes. The epigynal teeth in Iwogumoa and Himalcoelotes in general arise anteriorlybut could arise slightly anteriorly or even posteriorly in one Iwogumoa species. Ton-silla, Urocoras, Lineacoelotes and Pireneitega usually have the epigynal teeth arisingmedially at the same level as the anterior atrial margin but could be anteriorly or pos-teriorly. Coelotes has medially or posteriorly arising epigynal teeth, and those specieswith anteriorly arising teeth seem not to be congeneric with Coelotes. Most of theDraconarius species have anteriorly or medially, only occasionally posteriorly, arisingepigynal teeth. One species, Draconarius postremus Wang and Jäger, 2007 from Laos,has the epigynal teeth arising from the posterior margin of the epigynum (Figure 2I)(Wang and Jäger 2007). The position of epigynal teeth could be informative in defin-ing the species group within a genus.

Character five: separation of epigynal teeth

We define the separation of epigynal teeth by comparing their position transverselyrelative to the width of atrium (Table 5). The epigynal teeth could be contiguous(Figures 1G, 2A), separated by less than an atrial width (Figures 1D,H,I, 2B,F,H),separated by approximately an atrial width (Figures 1E, 2G), or separated by morethan an atrial width (Figures 1F, 2C–E,I). This character shows a high level of varia-tion among species of each genus. Two genera have the epigynal teeth separated by

Table 3. (Continued).

Epigynal teeth, length


Species list

Short (n = 211) Coelotes 91 Entries rest of genusCoras* 15 AllDraconarius 71 Entries rest of genusEurocoelotes 6 E. anoplus (Kulczynski, 1897); E. brevispinus

(Deltshev and Dimitrov, 1996); E. gasperinii (Simon, 1891); E. inermis (L. Koch, 1855); E. kulczynskii (Drensky, 1915); E. microlepidus (de Blauwe, 1973)

Iwogumoa* 14 AllLeptocoelotes* 1 Le. speudoluniformis (Zhang, Peng and Kim, 1997)Pireneitega 5 P. garibaldii (Kritscher, 1969); P. pyrenaea

(Simon, 1870); P. segestriformis (Dufour, 1820); P. taiwanensis (Wang and Ono, 1998); P. tianchiensis (Wang et al., 1990)

Tegecoelotes* 6 AllTonsilla 3 T. lyrata (Wang et al., 1990); T. lautispina (Wang

et al., 1990); T. variegata (Wang et al., 1990)

Total 291

Note: The asterisk indicates that all species of the genus have this character state.



Table 4. Character four: epigynal teeth, position (all data from Wang 2009).

Epigynal teeth, position


Species list

Anteriorly (n = 74) At level anteriorly of atrium (n = 74)

Coelotes 20 See Wang (2009)

Draconarius 23 See Wang (2009)Himalcoelotes 9 H. brignolii Wang, 2002;

H. bursarius Wang, 2002; H. diatropos Wang, 2002; H. gyirongensis (Hu and Li, 1987); H. martensi Wang, 2002; H. pirum Wang, 2002; H. sherpa (Brignoli, 1976); H. subsherpa Wang, 2002; H. xizangensis (Hu, 1992)

Iwogumoa 13 Entries rest of genusTonsilla 1 T. tautispina (Wang et al.,

1990)Urocoras 1 U. longispinus

(Kulczynski, 1897)Wadotes* 8 All

Medially (n = 172) At level slightly anteriorly of atrium (n = 27)

Coelotes 6 C. amamiensis Shimojana, 1989; C. antri (Komatsu, 1961); C. caudatus de Blauwe, 1971; C. gotoensis Okumura, 2007; C. multannulatus Zhang, Zhu and Song, 2006; C. yanlingensis Zhang, Yin and Kim, 2000

Draconarius 16 See Wang (2009)Himalcoelotes 2 H. aequoreus Wang, 2002;

H. syntomos Wang, 2002

Iwogumoa 1 I. pengi (Ovtchinnikov, 1999)

Lineacoelotes* 1 L. strenuus Xu, Li and Wang, 2008

Tonsilla 1 T. lyrata (Wang et al., 1990)

At level of anterior atrial margin (n = 136)

Bifidocoelotes* 2 All

(Continued)






Species list

Coelotes 54 See Wang (2009) for a list of species

Coras* 15 AllDraconarius 33 See Wang (2009) for a list

of speciesLeptocoelotes* 1 Le. pseudoluniformis

(Zhang, Peng and Kim, 1997)

Lineacoelotes 4 L. bicultratus (Chen, Zhao and Wang, 1991); L. funiushanensis (Hu, Wang and Wang, 1991); L. longicephalus Wang, Xu and Li, 2008; L. nitidus (Li and Zhang, 2002)

Pireneitega 12 P. involuta (Wang et al., 1990); P. liansui (Bao and Yin, 2004); P. luctuosus (L. Koch, 1878); P. luniformis (Zhu and Wang, 1994); P. major (Kroneberg, 1875); P. neglecta (Hu, 2001); P. spasskyi (Charitonov, 1946); P. spinivulva (Simon, 1880); P. taishanensis (Wang et al., 1990); P. tianchiensis (Wang et al., 1990); P. triglochinata (Zhu and Wang, 1994); P. xinping (Zhang, Zhu and Song, 2002)

Tegecoelotes* 6 AllTonsilla 5 T. defossa Xu and Li,

2006; T. eburniformis Wang and Yin, 1992; T. imitata Wang and Yin, 1992; T. truculenta Wang and Yin, 1992; T. variegata Wang et al., 1990






Species list

Urocoras 4 U. matesianus (de Blauwe, 1973); U. munieri (Simon, 1880); U. nicomedis (Brignoli, 1978); U. phthisicus (Brignoli, 1978)

At level slightly posteriorly of anterior atrial margin (n = 9)

Coelotes 6 C. kirgisicus Ovtchinnikov, 2001; C. motobuensis Shimojana, 2001; C. pseudoterrestris Schenkel, 1963; C. sordidus Ovtchinnikov, 2001; C. striatilamnis ketmenensis Ovtchinnikov, 2001; C. terrestris (Wider, 1834)

Pireneitega 3 P. armeniaca (Brignoli, 1978); P. cottarellii (Brignoli, 1978); P. taiwanensis (Wang and Ono, 1998)

Posteriorly (n = 44)

At level posteriorly of anterior atrial margin (n = 43)

Coelotes 24 See Wang (2009)Draconarius 4 D. paraterebratus

Wang, 2003; D. pervicax (Hu and Li, 1987); D. qingzhangensis (Hu, 2001); D. sublutulentus Xu and Li, 2008

Eurocoelotes* 10 AllIwogumoa 1 I. dicranatus (Wang et al.,

1990)Pireneitega 4 P. fedotovi (Charitonov,

1946); P. garibaldii (Kritscher, 1969); P. pyrenaea (Simon, 1870); P. segestriformis (Dufour, 1820)

At level close to epigastric furrow (n = 1)

Draconarius 1 D. postremus Wang and Jäger, 2007

Total 291

Note: The asterisk means that all species of the genus have this character state.



less than an atrial width, i.e. Lineacoelotes and Leptocoelotes, whereas in Pireneitegathey are separated by about an atrial width. Tonsilla species in general have the con-tiguous or slightly separated teeth except T. defossa Xu and Li, 2006, which has theteeth distinctly separated by less than an atrial width. The generic placement ofT. defossa is doubtful because it also lacks a large atrium. The epigynal teeth of

Figure 2. Epigynal tooth morphology. (A) Tonsilla tautispina (Wang et al., 1990): short,arise at level anteriorly of atrium, contiguous. (B) Iwogumoa ensifer (Wang and Ono, 1998):short, arise at level anteriorly of atrium, separated by less than an atrial width. (C) Draco-narius episomos Wang, 2003: short, arise at level slightly anteriorly of atrium, separated bymore than an atrial width. (D) Coelotes alpinus Polenec, 1972: short, arise at level of anterioratrial margin, separated by more than an atrial width. (E) Coelotes pseudoterrestris Schenkel,1963: short, arise at level slightly posteriorly of anterior atrial margin, separated by morethan an atrial width. (F) Draconarius adligansus (Peng and Yin, 1998): short, arise at level ofanterior atrial margin, separated by less than an atrial width. (G) Coelotes guttatus Wang etal., 1990: short, arise at level posteriorly of anterior atrial margin, separated by about anatrial width. (H) Coelotes rugosus (Wang, Peng and Kim, 1996): short, arise at level posteri-orly of anterior atrial margin, separated by about an atrial width. (I) Draconarius postremusWang and Jäger, 2008: short, arise at level of epigastric furrow, separated by more than anatrial width.



Table 5. Character five: epigynal teeth, separation (all data from Wang 2009).

Epigynal teeth, separation


Species list

Contiguous (n = 22) Coelotes 14 See Wang (2009)Draconarius 2 D. acidentatus (Peng and Yin 1998);

D. digitusiformis (Wang et al. 1990)Tonsilla 6 T. eburniformis Wang and Yin, 1992;

T. imitata Wang and Yin, 1992; T. lyrata (Wang et al., 1990); T. tautispina (Wang et al., 1990); T. truculenta Wang and Yin, 1992; T. variegata (Wang et al., 1990)

Separated by less than atrial width (n = 51)

CoelotesDraconariusHimalcoelotesIwogumoa

201219

See Wang (2009)See Wang (2009)H. aequoreus Wang, 2002I. acco (Nishikawa, 1987); I. ensifer (Wang

and Ono, 1998); I. illustrata (Wang et al., 1990); I. interuna (Nishikawa, 1977); I. montivaga (Wang and Ono, 1998); I. nagasakiensis Okumura, 2007; I. pengi (Ovtchinnikov, 1999); I. taoyuandong (Bao and Yin,2004); I. yushanensis (Wang and Ono, 1998)

Leptocoelotes* 1 Le. pseudoluniformis (Zhang, Peng and Kim, 1997)

Lineacoelotes* 5 All Lineacoelotes speciesTonsilla 1 T. defossa Xu and Li, 2006Tegecoelotes 3 Te. corasides (Bösenberg and Strand, 1906);

Te. ignotus (Bösenberg and Strand, 1906); Te. secundus (Paik, 1971)

Separated by atrial width (n = 123)

CoelotesCorasDraconariusEurocoelotes

3314334

See Wang (2009)Entries rest of genusSee Wang (2009)E. anoplus (Kulczynski, 1897);

E. deltshevi (Dimitrov, 1996); E. falciger (Kulczynski, 1897); E. microlepidus (de Blauwe, 1973)

Himalcoelotes 10 Entries rest of genusIwogumoa 4 I. dicranata (Wang et al., 1990); I. insidiosa (L.

Koch, 1878); I. songminjae (Paik and Yaginuma, 1969); I. tengchihensis (Wang and Ono, 1998)

Pireneitega* 19 AllTegecoelotes 2 Te. dysodentatus Zhang and Zhu, 2004;

Te. mizuyamae Ono, 2008

(Continued)



Himalcoelotes, Coras and Urocoras are separated by about an atrial width, butoccasionally by either less than or more than an atrial width. In Eurocoelotes, theepigynal teeth can be separated by an atrial width or slightly more than an atrialwidth (Wang et al. 2010). Tegecoelotes species also show mixed character states,with one species separated by more than an atrial width, two by about an atrialwidth, and three others by less than an atrial width. In general, Iwogumoa hasshort, closely set epigynal teeth, which could be less than, more than, or about anatrial width because of the variation in atrium size. The genus Coelotes usuallyexhibit epigynal teeth separated either by an atrial width or by more than anatrial width. The contiguous epigynal teeth are found in six species of the Coe-lotes charitonovi group (Wang and Zhu 2009) but the other two species from thesame group are distinctly separated by less than an atrial width. Eight other Coe-lotes from China and Japan also share the closely set teeth but they could be mis-placed in Coelotes. Of the 20 Coelotes with the epigynal teeth separated by lessthan an atrial width, two of them belong to the Coelotes charitonovi group andothers might be misplaced. Draconarius might have the epigynal teeth that areseparated by more than, less than, or by about an atrial width, but the genericplacement of many currently involved species need to be further justified. Thetwo species that have the contiguous teeth are obviously not congeneric withDraconarius.

Similar to the position of epigynal teeth, the separation of epigynal teeth isrelative to the width of the atrium, which could vary itself in size and position.Another possible reference for epigynal tooth position might be their relativeposition to the epigynal hood. Epigynal teeth could be situated: at a level anterior


Epigynal teeth, separation


Species list

Urocoras 4 U. longispinus (Kulczynski, 1897); U. munieri (Simon, 1880); U. nicomedis (Brignoli, 1978); U. phthisicus (Brignoli, 1978)

Separated by more than atrial width (n = 83)

CoelotesCorasDraconariusEurocoelotes

421

306

See Wang (2009)C. juvenilis (Keyserling, 1881)See Wang (2009)E. brevispinus (Deltshev and Dimitrov, 1996);

E. gasperinii (Simon, 1891); E. inermis (L. Koch, 1855); E. jurinitschi (Drensky, 1915); E. karlinskii (Kulczynski, 1906); E. kulczynskii (Drensky, 1915)

Iwogumoa 2 I. xinhuiensis (Chen, 1984); I. yaeyamensis (Shimojana, 1982)

Urocoras 1 U. matesianus (de Blauwe, 1973)Tegecoelotes 1 Te. michikoae (Nishikawa, 1977)

Total 280

Note: The asterisk indicates that all species of the genus have this character state.



of the epigynal hoods, at the level of the epigynal hoods, or posterior of the epig-ynal hoods. Epigynal teeth could be separated by: the distance of epigynal hoods,less than half, about half or more than half the distance of epigynal hoods.Although the position of the epigynal hoods shows a relatively high level of con-sistency, the position and size of atrium could be closely related to the epigynalteeth in their functions.

Character six: width of epigynal teeth

The epigynal teeth can be broad (wider than long, or subequal in length and width)(Figure 1I) or slender (either small or at least twice as long as wide) (Table 6). Theteeth of Wadotes and Coelotes wangi Chen and Zhao, 1997 show an intermediatecondition between broad and slender (Figure 1B,C). The broad teeth are present inLeptocoelotes and Tegecoelotes, whereas the slender teeth are unique in all other gen-era except one Coelotes species and four Draconarius species. The generic status ofthe four Draconarius with broad epigynal teeth also needs to be tested in any futureanalysis.

Table 6. Character six: epigynal teeth, width (all data from Wang 2009).

Epigynal teeth, width


Species list

Broad (n = 12) Coelotes 2 C. uozumii Nishikawa, 2002Draconarius 4 D. latidens Wang and Jäger, 2008;

D. stemmleri (Brignoli, 1978); D. trifasciatus (Wang and Zhu, 1991); D. yadongensis (Hu, 1987)

Leptocoelotes* 1 Le. pseudoluniformis (Zhang, Peng and Kim, 1997)

Tegecoelotes* 6 All

More or less broad (n = 9)

Wadotes*Coelotes

81

AllC. wangi Chen and Zhao, 1997

Slender (n = 269) Bifidocoelotes* 2 AllCoelotes 108 Entries rest of genusCoras* 15 AllDraconarius 73 Entries rest of genusEurocoelotes* 10 AllHimalcoelotes* 11 AllIwogumoa* 15 AllLineacoelotes* 5 AllPireneitega* 19 AllTonsilla* 7 AllUrocoras* 5 All

Total 291

Note: The asterisk means that all species of the genus have this character state.



Taxonomy

Family AMAUROBIIDAE Thorell, 1870Subfamily COELOTINAE F.O. Pickard-Cambridge, 1893

Genus Draconarius Ovtchinnikov, 1999Draconarius wrasei sp. nov.

(Figures 3–5, 9)

Type material

Holotype. Female, China: Yunnan: Zhongdian County, 36 km east-southeast Zhong-dian, 3500–3550 m, 27°40.9′ N, 100°01.05′ E, overgrown rock hillside with old mixedforest, bamboo, dead wood, vinegar trap, 23–24 August 2003, D.W. Wrase, depos-ited in SMF.

Etymology

The specific name is a patronym in honour of D.W. Wrase, the German carabid spe-cialist who collected the specimen; noun in genitive case.

Diagnosis

The female of this new species has an epigynum similar to D. incertus Wang, 2003and related species, but can be easily recognized by the posteriorly arising spermat-hecal heads (Figures 3A,B, 4A,B).

Description

Female (holotype). Medium-sized Coelotinae, total length 6.09 (Figure 5A–D). Dor-sal shield of prosoma 3.09 long, 2.12 wide; opisthosoma 3.00 long, 2.00 wide. AMEsmallest, half the size of other eyes, which being subequal, or with PME slightlysmaller (AME 0.06, ALE 0.13, PME 0.12, PLE 0.13); AME separated from eachother by slightly more than their size, from ALE by slightly less than AME diameter;PME separated from each other by slightly less than their diameter, from PLE by

Figure 3. Draconarius wrasei, sp. nov., female holotype, epigynum: (A) ventral, (B) dorsal.



PME diameter (AME–AME 0.08, AME–ALE 0.05, PME–PME 0.11, PME–PLE0.13, AME–PME 0.09) (Figure 5D). Labium slightly wider than long (length/width = 0.90) (Figure 5E). Chelicera with three promarginal and two retromarginalteeth. Epigynal teeth absent; atrium large, slightly wider than long, anterior atrialmargin recurved medially to a distinct septum, lateral atrial margins indistinct, ridgesin atrium distinct, limited to posterior half of atrium; copulatory ducts small, origi-nating posteriorly; spermathecae with large, distinctly separated bases, which extendanterior of spermathecal heads by a length at least as long as spermathecal bases;spermathecal heads arising posteriorly close to bases (Figures 3A,B, 4A,B).

Male. Unknown.

Relationships

Draconarius wrasei sp. nov. is a member of the incertus group because it lacks theepigynal teeth and has a similar epigynum, but the posteriorly arising spermathecal

Figure 4. Draconarius wrasei, sp. nov., female holotype, epigynum: (A) ventral, (B) dorsal(part of the spermathecal bases are covered by tissues in B).

Figure 5. Draconarius wrasei, sp. nov., female holotype: (A–C) habitus, dorsal, lateral andventral view; (D) eyes, view between front and dorsal; (F) labium, ventral view.



heads and the long, distinctly anterior extending spermathecae distinguish it fromother members of this group.

Distribution

China (Yunnan) (Figure 9).

Draconarius immensus Xu and Li, 2006(Figures 6–9)

Material examined

Five males, one female, China: Yunnan, Zhongdian County, 36 km east-southeastZhongdian, 3500–3550 m, 27°40.9′ N, 100°01.05′ E, overgrown rock hillside with oldmixed forest, bamboo, dead wood, vinegar trap, 23–24 August 2003, D.W. Wrase(SMF).

Remarks

The male specimens examined are the same as D. immensus but the female differsslightly in the shape and separation of spermathecae. This might be because of theindividual variation in vulva structure, which is common in the incertus speciesgroup. The female is similar to D. papillatus Xu and Li 2006, which is also found inZhongdian County, Yunnan, in addition to its distribution in Sichuan and Tibet (Xuand Li 2006). Draconarius papillatus might be a junior synonym of D. immensus butthis needs to be confirmed by collecting and examining male D. papillatus from theholotype locality (Sumdo, 29.1° N, 100.1° E, Xiangcheng County, Sichuan) (Xu andLi 2006). The eyes and body of the specimens examined in this study are also meas-ured for the purpose of future comparison regarding the species status ofD. immensus and D. papillatus.

Diagnosis

This species is similar to D. incertus in having a similar epigynum, a bifurcate conduc-tor, and a simple median apophysis, but can be distinguished by the spermathecaeextending anteriorly beyond their heads and the spermathecal heads arising mediallybetween spermathecae in the female, and by the short median apophysis, the embolusoriginating between prolateral and proximal at about the 6 to 7 o’clock position andrunning more than one-quarter of an oval (Figures 6, 7).

Description

Female. Large sized Coelotinae, total length 11.50 (Figure 8A–C). Dorsal shield ofprosoma 5.50 long, 3.16 wide; opisthosoma 6.00 long, 4.20 wide. AME smallest,slightly more than half the size of ALE; ALE largest; posterior eyes subequal,three-quarters the size of ALE (AME 0.13, ALE 0.21, PME 0.15, PLE 0.16); AME



separated from each other by their diameter, from ALE by slightly less than AMEdiameter; PME separated from each other by more than their diameter, widely sep-arated from PLE by slightly more than 1.5 times PME diameter (AME–AME 0.13,AME–ALE 0.10, PME–PME 0.19, PME–PLE 0.25, AME–PME 0.18) (Figure 8D).Labium slightly longer than wide (length/width = 1.06) (Figure 8E). Promargin ofchelicera with three teeth, retromargin with two teeth. Epigynal teeth absent;atrium large, wider than long, anterior atrial margin recurved medially to a distinctseptum, lateral atrial margins indistinct, ridges in atrium distinct and extending toanterior half of atrium; copulatory ducts small, originating posteriorly betweenspermathecae; spermathecae broad, distinctly separated by about half of theirwidth, slightly extending anterior of their heads; spermathecal heads small, arisingmedially between spermathecae (Figures 6A,B, 7A,B).

Male. Large Coelotinae, total length 10.15 (Figures 8F–H). Dorsal shield of pro-soma 5.15 long, 3.00 wide; opisthosoma 5.00 long, 3.00 wide. AME smallest,ALE largest; posterior eyes subequal in size, slightly smaller than ALE (AME0.14, ALE 0.18, PME 0.16, PLE 0.17); AME separated from each other byslightly more than half of their diameter, from ALE by one-third of AME diame-ter; posterior eyes almost equally separated by slightly less than their diameter(AME–AME 0.08, AME–ALE 0.05, PME–PME 0.14, PME–PLE 0.15, AME–PME 0.13) (Figure 8I). Labium slightly longer than wide (length/width = 1.16)(Figure 8J). Promargin of chelicera with three teeth, retromargin with two teeth.Palpal patellar apophysis short, curved dorsally; RTA more than half of tibial

Figure 6. Draconarius immensus Xu and Li 2006: (A,B) female pigynum, ventral and dorsalviews; (C,D) male palp, ventral and retrolateral views.



length, with distinctly protruding distal end; lateral tibial apophysis distinct;cymbial furrow less than half of tibial length; conductor short, with a broad base,a slender, bifurcate apex, a small basal lamella, and a slender dorsal apophysis;median apophysis short, not spoon-shaped; embolus short, filiform, originatingbetween prolateral and proximal at approximately 6 to 7 o’clock position,extending posteriorly to distal part of tibia, anteriorly not coiling beyond distalpart of bulb (Figures 6C,D, 7C–E).

Relationships

Draconarius immensus could be the sister species to D. incertus.

Distribution

China (Yunnan, Sichuan) (Figure 9).

Figure 7. Draconarius immensus Xu and Li 2006 : (A,B) female epigynum, ventral and dorsalviews; (C–E) male palp, prolateral, ventral and retrolateral views.



Figure 8. Draconarius immensus Xu and Li 2006, female (A–E) and male (F–J): (A–C, F–H)habitus, dorsal, lateral and ventral views; (D,I) eyes, view between front and dorsal; (E,J)labium, ventral view.

Figure 9. Distribution records of Draconarius wrasei sp. nov. (black circle only) and Draco-narius immensus Xu and Li 2006 (black square and circle).



Acknowledgements

We thank David W. Wrase (Berlin) for collecting the specimens and Charles E. Griswold(California Academy of Sciences, San Francisco), Norman I. Platnick (American Museum ofNatural History, New York), Hirotsugu Ono (National Museum of Nature and Science,Tokyo) for discussion of coelotine epigynal tooth morphology. Coelotinae colleagues Shu-Qiang Li, Jie Liu (Institute of Zoology, Beijing), Xiang Xu (Hunan Norman University,Changsha), Zhi-Sheng Zhang (Northwest University, Chongqin) and Ming-Sheng Zhu (HebeiUniversity, Baoding) exchanged ideas on Coelotinae genital morphology, read the manuscriptand provided critical comments.

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Documents

Journal of Natural History A review of Coelotinae epigynal teeth