2005. The Journal of Arachnology 33:415±425

LARVAL CHAETOTAXY IN WOLF (ARANEAE, LYCOSIDAE): SYSTEMATIC INSIGHTS AT THE SUBFAMILY LEVEL

Beata Tomasiewicz: Zoological Institute, Department of and Evolutionary , Przybyszewskego 63/77, 51±148 Wrocøaw, Poland. E-mail: [email protected] Volker W. Framenau: Department of Terrestrial Invertebrates, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia.

ABSTRACT. Studies into the systematics of wolf spiders have mainly employed morphological char- acters of adult spiders, in particular features of the male and female genitalia, and more recently mito- chondrial DNA sequence data. However, there is still no established phylogenetic framework for the Lycosidae, even at the subfamily level. This study uses a novel morphological character set, the chaetotaxy of lycosid larvae (presence and arrangement of setae and slit organs), to infer systematic information on seven species of wolf spiders that are currently listed in three subfamilies: Lycosinae [ pulver- ulenta (Clerck 1757), Hogna antelucana (Montgomery 1904), Rabidosa rabida (Walckenaer 1837), Tro- chosa ruricola (DeGeer 1778)], Piratinae [Hygrolycosa rubrofasciata (Ohlert 1865), Pirata hygrophilus (Clerck 1757)], and Sosippinae (Sosippus californicus Simon 1898). Cheliceral and tarsal (legs I and II) chaetotaxic patterns of the ®rst postembryo showed equivalent chaetotaxic complexes amongst all species but revealed considerable differences between representatives of the three subfamilies. Sosippus califor- nicus showed the most complex pattern and P. piraticus the most reduced arrangement. In addition, it casts doubt on the previous listings of H. rubrofasciata in either the Lycosinae or Piratinae, as its chae- totaxic setae arrangement was more similar to S. californicus than to any other species investigated here. Keywords: Larval stages, chaetotaxic complex, Lycosinae, Piratinae, Sosippinae

Chaetotaxy, the presence and arrangement and larval chaetotaxy has been ar- of setae and other sensory structures on the gued to represent an excellent character set for integument of arthropods, has been widely systematic studies (Pomorski 1996). Larval used for systematic and taxonomic studies in morphology is of particular interest in holo- a variety of groups, including insects (e.g. metabolic insects such as beetles (Kilian Alarie & Watts 2004) and such as 1998; Borowiec & SÂwieÎtojanÂska 2003) and mites (Tuzovsky 1987; Grif®th et al. 1990) butter¯ies (Kitching 1984, 1985) as different and pseudoscorpions (e.g. Chamberlin 1931; expressions of the same genotype can com- Harvey 1992). In contrast, investigations into plement the morphological characters of the chaetotaxy of spiders are comparatively adults (Alarie & Watts 2004; Grebennikov rare and have focused mainly on trichoboth- 2004; Ashe 2005). However, larval chaetotaxy rial patterns. These have been argued to be a has also been useful in phylogenetic and tax- suitable feature in higher level systematics onomic studies of arthropods with gradual (Lehtinen 1980; Scioscia 1992). They may larval development such as springtails, mites also serve as an important tool in identi®ca- and pseudoscorpions (Nayrolles & Betsch tion at the species level. For example, the po- 1993; Pomorski 1996; Grif®th et al. 1990; sition of the metatarsal trichobothrium has Harvey 1992). Early stages of development been used as an essential character in the iden- may last for only a short period of time, ti®cation of central European Micryphantinae which in many cases eliminates the devel- Bertkau 1872 (Wiehle 1960; Heimer & Nen- opment of distinct adaptive traits. In addition, twig 1990). the morphology of juveniles is less variable There is a considerable difference between and complex than that of adults (Pomorski the chaetotaxic pattern of immature and adult 1996).

415 416 THE JOURNAL OF ARACHNOLOGY

Figures 1, 2.ÐLarval setae of wolf spiders. 1. Seta form [position: apical/etc leg/chelicera etc] of [species]; 2. Serrated seta from [position: apical/etc., leg/chelicera etc.] of [species]. Scale bar: 10 ␮m (Fig. 1), 5 ␮m (Fig. 2).

Studies on chaetotaxic structures in imma- de®ned morphological characters that could ture spiders are rare and initially focused on classify and separate particular genera and trichobothrial patterns (Emerit 1964). A recent subfamilies. However, there appears to be a study of the linyphiid Bathyphanthes consensus that web-building wolf spiders, eumenis (L. Koch 1879) included all sensory such as the genera Sosippus Simon 1888 structures of the protonymph and showed that (sheet-web) and Pirata Sundevall 1833 (tube- the arrangement of sensory organs such as se- shaped retreat) represent more ancient evolu- tae, trichobothria and slit organs was constant tionary lines in comparison to genera within in all examined specimens and may have the the Lycosinae Simon 1898 ( C.L. potential to serve in the identi®cation of spi- Koch 1847, Alopecosa Simon 1885, Rabidosa ders at the generic and species level (Rybak Roewer 1960 and Hogna Simon 1885) that are & Pomorski 2003). The nomenclature of the considered representatives of more recent evo- chaetotaxic patterns developed for B. eumenis lutionary lineages (Dondale 1986; Zehethofer was subsequently used in a detailed compar- & Sturmbauer 1998; Vink et al. 2002). ative study including the Trochosa The genus Hygrolycosa Dahl 1908 was for- ruricola (DeGeer 1778) (Lycosidae) (Rybak merly placed in the Lycosinae along with, & Tomasiewicz 2005). Although this study showed considerable differences in chaetotax- amongst others, Alopecosa, Hogna and Tro- ic pattern between both species, some body chosa (Dondale 1986). However, more re- parts showed very similar setae distribution, cently, it was listed in a separate subfamily, which suggested homology for a large number Piratinae Zyuzin 1993, based on the shape and of chaetotaxic complexes. location of the embolus and the functional Despite recent investigations into the sys- conductor in the male pedipalp (Zyuzin 1993). tematics of wolf spiders, there is still no ac- Current molecular evidence suggests that Hy- cepted phylogenetic framework for the Lycos- grolycosa is a sister taxon to Aulonia albi- idae, even at the subfamily level (e.g. Dondale mana (Walckenaer 1805) in a clade that also 1986; Zyuzin 1993; Vink et al. 2002). This includes Pirata, Venonia Thorell 1894 (Ven- problem can be attributed to a lack of well- oniinae Lehtinen & Hippa 1979) and Xeroly- TOMASIEWICZ & FRAMENAUÐLARVAL CHAETOTAXY IN WOLF SPIDERS 417

Table 1.ÐNomenclature of chaetotaxic complexes on the larval integuments of A. pulverulenta, H. antelucana, H. rubrofasciata, P. hygrophilus, R. rabida, S. californicus and T. ruricola.

Abbreviation Chaetotaxic complex Illustrations Chelicerae

ChDA dorsal apical complex Figs. 1±4 ChDM dorsal median complex Figs. 1±4 ChVAM ventral apico-median complex Figs. 5±8 ChVM ventral median complex Figs. 7±8 Tarsi I and II

TDA dorsal apical complex Fig. 9 TDAI,TDAII. . . ®rst, second, . . . dorsal apical complex Figs. 10±11 TDM dorsal median complex Fig. 9 TDMI,TDMII. . . ®rst, second, . . . dorsal median complex Figs. 10±13 TDP dorsal proximal complex Figs. 10±13 TVAI,TVAII. . . ®rst, second, . . . ventral apical complex Figs. 12±14 TVM ventral median complex Fig. 12 TVMI,TVMII. . . ®rst, second, . . . ventral medial complex Figs. 13±14 TVP ventral proximal complex Figs. 13±14

cosa Dahl 1908 (Evippinae Zyuzin 1985) (N. iation in regard to the number of structures Murphy et al. in press). within chaetotaxic complexes, which allowed The main objective of this study was to analysis of data without statistical consider- evaluate the signi®cance of larval chaetotaxic ation of variation. patterns for systematic analyses in wolf spi- Specimens were transferred to 5% KOH ders. More speci®cally, we used the ambigu- and cleared in distilled water. Subsequently, ous subfamily placement of H. rubrofasciata they were placed in chloramphenol and to assess its previous listings in either the Ly- mounted in Swan medium (20 g distilled wa- cosinae or Piratinae by including representa- ter, 60 g chloral hydrate, 15 g gum arabic, 3 tives of these subfamilies in our comparative g glucose, 2 g glacial acetic acid). All slides analysis. were examined under a phase contrast micro- METHODS scope (Nikon Eclipse E 600) with a drawing attachment. Scanning electron microscope We analyzed the larval stages of seven spe- (SEM) photographs were taken with a JEOL cies of wolf spiders currently listed in three JSM-5800 LV at 15kV after spray-coating the different subfamilies: Lycosinae [Alopecosa specimen with gold. Voucher specimens of the pulverulenta (Clerck 1757), Hogna antelu- cana (Montgomery 1904), Rabidosa rabida species examined were lodged at the Museum (Walckenaer 1837), and of Natural History, Wrocøaw (A. pulverulenta, (DeGeer 1778)], Piratinae [Hygrolycosa rub- P. hygrophilus, H. rubrofasciata) and the Cal- rofasciata and Pirata hygrophilus (Clerck ifornia Academy of Sciences, San Francisco 1757)], and Sosippinae (Sosippus californicus (H. antelucana, R. rabida, S. californicus). Simon 1898). We obtained immature stages Larval stages.ÐWe investigated the ®rst through laboratory colonies (T. ruricola, A. immature stage that possesses chaetotaxic pulverulenta, H. rubrofasciata and P. hygro- structures on the integument, i.e. the ®rst pos- philus) or loan and donation of material from tembryo. These young spiders develop inside overseas collections (H. antelucana, R. rabida the egg-sac followed by the protonymph, and S. californicus). which abandons the egg-sac (Vachon 1957). Overall, we studied 64 specimens of T. rur- Vachon (1957) proposed the term `larva' for icola, 10 specimens each of H. rubrofasciata the ®rst postembryo, which corresponds to and P. hygrophilus and 5 specimens each of `stage D' (Holm 1940), `preÂjuvenile (Ji 1)' A. pulverulenta, H. antelucana, R. rabida, and (Canard 1987), `larva ``setose stage''' (Hallas S. californicus. There was no intraspeci®c var- 1988), and `IV instar' (Galiano 1991). Con- 418 THE JOURNAL OF ARACHNOLOGY sequently, all references to `larvae' or `larval' allowed a comparison between species and in this study refer to the ®rst postembryo. genera. These structures were most complex Chaetotaxic structures.ÐAlthough nu- in S. californicus (Figs. 6, 10, 13, 16) and H. merous chaetotaxic structures such as spines, rubrofasciata (Figs. 5, 9, 12, 15) and most trichobothria, proprioreceptors in the form of reduced in P. piraticus (Figs. 3, 7, 11, 14). hair plates, and chemoreceptors in the form of Chelicerae dorsal.ÐAll species possessed tarsal organs, and taste hairs exist in spiders the apical complex ChDA. The number of setae (Foelix 1996; Rybak & Pomorski 2003), this within this complex differed between P. hy- study deals with setae and slit organs because grophilus (four setae; Fig. 3), a group com- only these structures were observed on the lar- prising T. ruricola, A. pulverulenta, R. rabida, val integument. In adult spiders, setae are tri- and H. antelucana (seven setae; Fig. 4) and a ply innervated hair-like structures that serve group with H. rubrofasciata and S. californi- purely mechanical tasks (tactile receptors). cus (10 setae; Figs. 5, 6). Hygrolycosa rub- They consist of a long exocuticular shaft of rofasciata and S. californicus had an addition- variable shape (including serrated and plu- al median complex ChDM that consisted of mose), which is suspended in a slipper-shaped three setae, which were long in S. californicus socket in which it can move (Rybak & Po- and very short in H. rubrofasciata. All species morski 2003). In contrast, spines are rigid had one slit sensilla in the median section of structures that are regarded as hemolymph the chelicerae and two slit sensillae apically pressure receptors (Foelix & Chu-Wang (Figs. 3±6). 1973). In immature spiders, it is dif®cult to Chelicerae ventral.ÐAll species showed distinguish between spines and setae as the an apico-median complex ChVAM that consist- socket and the setae are generally not fully ed of one or two setae in P. hygrophilus (Fig. developed (Figs. 1, 2), although different 7), and four setae in all other species (Figs. types of setae may exist (Bond 1994). Con- 8±10). Hygrolycosa rubrofasciata (Fig. 9) and sequently, within the scope of our study, we S. californicus (Fig. 10) possessed a further do not differentiate between setae and spines. structure ChVM, consisting of a single seta in Slit organs occur both in adult and larval spi- H. rubrofasciata and two setae in S. califor- ders. They sense mechanical stress in the exo- nicus. The latter species showed an additional skeleton caused by vibrations, gravity or the apical seta ChVA that did not exist in any of spider's own movement and occur singly (`slit the other lycosids. All species possessed two sensillae') or in groups where slits run parallel slit sensillae apically (Figs. 7±10). to each other (`lyriform organs') (Foelix Tarsi of legs I and II dorsal.ÐAll species

1996). In this study, the chaetotaxic structures examined showed two similar complexes, TDA on larval chelicerae and tarsi were grouped and TDM in T. ruricola, H. antelucana, R. ra- into distinct complexes. The nomenclature of bida, A. pulverulenta, and P. hygrophilus (Fig. these complexes follows Rybak & Pomorski 11), corresponding to TDAIII and TDMIV in H. (2003) and Tomasiewicz & Rybak (2005) (see rubrofasciata (Fig. 12) and TDAII and TDMIV in also Table 1). S. californicus) (Fig. 13). Hygrolycosa rub- rofasciata (Fig. 12) and S. californicus (Fig. RESULTS 13) showed seven more complexes in which There were considerable differences in the the apical ones had a larger number of setae number of chaetotaxic structures on the larval in S. californicus. All lycosids showed slit bodies of the investigated species, which al- sensillae located laterally in the median part lowed separating them into two main groups of the tarsi (Figs. 11±13). (Tables 2 & 3). While A. pulverulenta, H. an- Tarsi of legs I and II ventral.ÐAll spe- telucana, P. hygrophilus, R. rabida and T. cies showed three identical complexes, TVAI, ruricola possessed chaetotaxic structures only TVAII, and TVM in T. ruricola, H. antelucana, on the chelicerae, labium, maxillae, legs and A. pulverulenta and P. hygrophilus (Fig. 14), pedipalps, H. rubrofasciata and S. californi- corresponding to TVAI,TVAIII, and TVMI in H. cus exhibited chaetotaxy on all body parts in- rubrofasciata and S. californicus (Figs. 15± cluding sternum, carapace, abdomen and spin- 16). The complex TVMI consisted of three setae nerets. Chelicerae and the tarsi of legs I and in H. rubrofasciata (Fig. 15) (as the equiva-

II showed distinct chaetotaxic patterns, which lent complex TVM in the other above-men- TOMASIEWICZ & FRAMENAUÐLARVAL CHAETOTAXY IN WOLF SPIDERS 419

Figures 3±6.ÐChaetotaxic pattern on dorsal side of the chelicerae in wolf spider larvae: 3. Pirata hygrophilus;4.Alopecosa pulverulenta, Hogna antelucana, Trochosa ruricola, Rabidosa rabida;5.Hy- grolycosa rubrofasciata;6.Sosippus californicus. Scale bar: 0.1 mm. Multiple scale bars in Fig. 4 re¯ect the comparative scale of the species in the given sequence. 420 THE JOURNAL OF ARACHNOLOGY

Figures 7±10.ÐChaetotaxic pattern on ventral side of the chelicerae in wolf spider larvae: 7. Pirata hygrophilus;8.Alopecosa pulverulenta, Hogna antelucana, Trochosa ruricola, Rabidosa rabida;9.Hy- grolycosa rubrofasciata; 10. Sosippus californicus. Scale bar: 0.1 mm. Multiple scale bars in Fig. 8 re¯ect the comparative scale of the species in the given sequence. TOMASIEWICZ & FRAMENAUÐLARVAL CHAETOTAXY IN WOLF SPIDERS 421 tioned lycosids), but included four setae in S. by the overall distribution of chaetotaxic com- californicus (Fig. 16). Sosippus californicus plexes on the bodies of the spiders (Table 3), and H. rubrofasciata showed six additional does not re¯ect current phylogenetic hypoth- complexes (TVAII,TVC,TVMII,TVMIII,TVMIV,TVP eses for wolf spiders. Morphological (Dondale in H. rubrofasciata and TVAII,TVMII,TVMIII, 1986) and molecular (Zehethofer & Sturm- TVMIV,TVMV,TVP in S. californicus (Figs. 15, bauer 1998; Vink et al. 2002; Murphy et al. 16). Although these two species showed the in press) phylogenies consider the Lycosinae most similar chaetotaxic patterns, there are as the most derived lineage of wolf spiders, complexes (TVMIV,TVC in H. rubrofasciata and whereas the Piratinae and Sosippinae are TVMIV,TVMV in S. californicus) (Figs. 15, 16) thought to represent more basal evolutionary among which it is dif®cult to establish ho- lines. mology. Both species showed slit sensillae sit- Although we included a wide range of taxa uated medially near the apical part of the tarsi, from different currently recognized subfam- which were absent in all other species (Figs. ilies our study is ambiguous in regards to the 14±16). plesiomorphic condition for larval chaetotax- ic structures. Both Pirata and the sheet-web DISCUSSION building Sosippus are thought to represent Our analysis of chaetotaxic patterns in wolf basal lineages in the evolution of wolf spi- spiders showed distinct and regular complexes ders but they differ considerably in their lar- for all species examined. These complexes ap- val chaetotaxy. Preliminary studies on the pear to be similar to the arrangement in other chaetotaxy of Pisaura mirabilis (Clerck spider families such as the Linyphiidae (Ry- 1757) representing the Pisauridae, a putative bak & Pomorski 2003), which suggests that sister taxon of the Lycosidae (Dondale 1986; larval chaetotaxy may serve as a very useful Griswold 1993), show considerably reduced character set in systematic studies if homolo- chaetotaxic patterns (Tomasiewicz unpub. gies can be established on a higher taxonomic data) supporting P. hygrophilus to display the level. However, there was no difference of plesiomorphic state. In this case, and in com- chaetotaxic patterns among any of the species bination with current phylogenetic hypothe- currently included in the subfamily Lycosinae. ses (Murphy et al. in press), an increase in In contrast to other arthropods, in particular chaetotaxic structures has evolved twice insects (e.g. Deruaz et al. 1991; Alarie & within our sampled taxa, in Sosippus and Hy- Watts 2004), larval chaetotaxy does not seem grolycosa. to be suitable for the identi®cation of taxa be- The chaetotaxic pattern of H. rubrofascia- low subfamily level in wolf spiders. ta differs considerably from all other lycos- There were signi®cant differences in the ine and piratinae species examined, the two number of complexes of cheliceral and tarsal subfamilies where it was previously listed setae and the number and size within these (Dondale 1986; Zyuzin 1993) and our study complexes. Sosippus californicus showed the suggests an alternative placement within the most complex pattern along with H. rubrofas- Sosippinae. However, current molecular data ciata that differed only in the absence of two place H. rubrofasciata in a basal lineage setae on the ventral side of the chelicerae, the within in the Lycosidae, close to the Venon- absence of the complex equivalent to TVMIV in iinae, Piratinae and Evippinae (Murphy et al. S. californicus, and a reduction in the number in press), providing support for Zyuzin's of setae in the apical and proximal complexes (1993) placement of the genus and at the of the tarsi. On the other hand, all four species same time rejecting chaetotaxic patterns as of Lycosinae and P. piraticus showed very informative for the subfamilial placement of similar setal arrangements (Table 2). Here, Hygrolycosa. chaetotaxy was heavily reduced in comparison This preliminary study shows that larval to Sosippus and Hygrolycosa, in particular in chaetotaxy may provide some additional mor- regard to the tarsal setae. Pirata piraticus had phological evidence that bears phylogenetic the lowest number of setae as complex CHDA information in spiders although some discrep- and CHVM had two setae less each than the ancies with common tenets of current phylo- equivalent complexes in the Lycosinae. This genetic hypotheses in wolf spiders exist. It is separation into two major groups, supported not possible to distinguish species or even 422 THE JOURNAL OF ARACHNOLOGY

Figures 11±13.ÐChaetotaxic pattern on dorsal side of the tarsi of legs I and II in wolf spider larvae: 11. Pirata hygrophilus, Alopecosa pulverulenta, Hogna antelucana, Trochosa ruricola, Rabidosa rabida; 12. Hygrolycosa rubrofasciata; 13. Sosippus californicus. Scale bar: 0.1 mm. Multiple scale bars in Fig. 11 re¯ect the comparative scale of the species in the given sequence. Figures 14±16.ÐChaetotaxic pattern on ventral side of the tarsi of legs I and II in wolf spider larvae: 14. Pirata hygrophilus, Alopecosa pulverulenta, Hogna antelucana, Trochosa ruricola, Rabidosa rabida; 15. Hygrolycosa rubrofasciata; 16. Sosippus californicus. Scale bar: 0.1 mm. Multiple scale bars in Fig. 14 re¯ect the comparative scale of the species in the given sequence. genera on the basis of this feature, but the and the shape of the setae, similar to a study analysis of chaetotaxic patterns may help to in astigmatid mites (Grif®th et al. 1990), may establish relationships among subfamilies or prove helpful in establishing a detailed and above. A more detailed analysis not only into more informative character set based on larval the presence and absence but also the position chaetotaxy. Presence or absence of setae con- TOMASIEWICZ & FRAMENAUÐLARVAL CHAETOTAXY IN WOLF SPIDERS 423

Table 2.ÐNumber of setae per chaetotaxic complex on the chelicerae and tarsi of leg II and III of A. pulverulenta, H. antelucana, H. rubrofasciata, P. hygrophilus, R. rabida, S. californicus and T. ruricola.

A. pulverulenta H. antelucana P. hygrophilus R. rabida, T. ruricola H. rubrofasciata S. californicus

ChDA 7 7 10 10 ChDM ÐÐ 3 3 ChVAM 24 44 ChVM ÐÐ 1 2 TDA 33ÐÐ TDAI ÐÐ 5 5 TDAII ÐÐ 2 3 TDAIII ÐÐ 3 2 TDM 22ÐÐ TDMI ÐÐ 4 4 TDMII ÐÐ 3 3 TDMIV ÐÐ 2 2 TDMV ÐÐ 2 2 TDP ÐÐ 2 2 TVAI 22 22 TVAII ÐÐ 4 4 TVAIII ÐÐ 3 3 TVM 33ÐÐ TVMI ÐÐ 3 4 TVMII ÐÐ 4 5 TVMIII ÐÐ 3 2 TVMIV ÐÐ 4 3 TVMV ÐÐ Ð 3 TVC ÐÐ 2Ð TVP ÐÐ 4 4 tain only a limited amount of information homological chelal trichobothria in pseudo- since a reduction of structures may have easily scorpions (Harvey 1992) and the setal ar- occurred in multiple evolutionary lines. Dis- rangement in astigmatid mites (Grif®th et al. tinguishable morphological categories of setae 1990). exist in wolf spider larvae, for example Currently, it remains dif®cult to acquire lar- smooth and serrated forms (Figs. 1, 2). Future val material for morphological studies since research could explore an expanded character larvae and juveniles are often discarded dur- set and subsequently code it as morphological ing the collection of spiders, and, if collected, matrix for a phylogenetic analysis similar to the material may not represent a suitable de- some studies of insects (e.g., Alarie & Watts velopmental stage for comparative studies. 2004; Ashe 2005). This could then be incor- However, if larval chaetotaxy can be estab- porated in an exhaustive morphological and lished as an important morphological tool in molecular dataset for higher phylogenetic phylogenetic studies of spiders, the collection analysis in spiders. and preservation of spider larvae may receive The analysis of larval chaetotaxy may bear much stronger support. considerable importance in interpreting struc- tures of mature spiders, in particular during ACKNOWLEDGMENTS character polarization as part of a phyloge- We are grateful to Darrell Ubick (California netic analysis (ontogenetic criterion, see Hen- Academy of Science) for the loan and dona- nig 1966; Nelson 1978; Mabee 2000). For ex- tion of specimens of H. antelucana, R. rabida ample, the study of setal arrangement during and S. californicus for this study. Melissa postembryonic development has been helpful Thomas and Mark Harvey provided helpful in determining the phylogenetic migration of comments to improve this manuscript. TB was 424 THE JOURNAL OF ARACHNOLOGY

Table 3.ÐPresence of chaetotaxic structures on the larval bodies of A. pulverulenta, H. antelucana, H. rubrofasciata, P. hygrophilus, R. rabida, S. californicus and T. ruricola. sl ± slit sensillae, ly ± lyriform organs.

A. pulverulenta H. antelucana, P. hygrophilus H. rubrofasciata R. rabida, T. ruricola S. californicus Chelicerae setae, sl setae, sl Labium setae setae Maxillae setae setae Sternum Ð setae, sl Carapace Ð setae Pedipalps setae, ly, sl setae, ly, sl Legs setae, ly, sl setae, ly, sl Abdomen Ð setae Spinnerets Ð setae funded through the Zoological Institute of the Emerit, M. 1964. La trichothriotaxie et ses varia- Wrocøaw University and VWF through the tions au cour du deÂveloppement post-embryon- Australian Biological Resources Studies naire chez l'araigneÂe Gasteracantha versicolor (ABRS) to Mark Harvey (Western Australian (Walck.) (Argiopidae). Comptes Rendus de Museum) and Andy Austin (The University of l'Academie des Sciences Paris 258:4853±4854. Adelaide). Foelix, R.F. 1996. Biology of Spiders. Oxford Uni- versity Press, New York. LITERATURE CITED Foelix, R.F. & I.-W. Chu-Wang. 1973. The mor- Alarie, Y. & H.S. Watts. 2004. Larvae of the genus phology of the spider sensilla. I. Mechanorecep- Antiporus (Coleoptera: Dytiscidae) and phylo- tors. Tissue Cell 5:451±478. genetic implications. Invertebrate Systematics Galiano, M.E. 1991. Postembryonic development in 18:523±546. ten species of neotropical Salticidae (Araneae). Ashe, J.S. 2005. Phylogeny of the tachyporine Bulletin of British Arachnological Society 8: group subfamilies and basal lineages of aleo- 209±218. charines (Coleoptera: Staphylinidae) based on Grebennikov, V.V. 2004. Review of larval mor- larval and adult characteristics. Systematic En- phology of the beetle suborder Archostemata (In- tomology 30:3±37. secta: Coleoptera) with emphasis on ®rst-instar Bond, J.E. 1994. Seta-spigot homology and silk chaetotaxy. European Journal of Entomology production in ®rst instar Antrodiaetus unicolour 101:273±292. spiderlings (Araneae: Antrodiaetidae). Journal of Grif®th, D.A., W.T. Atyeo, R.A. Norton & C.A. Arachnology 22:19±22. Lynch. 1990. The idiosomal chaetotaxy of astig- Â Borowiec, L. & J. SwieÎtojanÂska. 2003. The ®rst in- matid mites. Journal of Zoology, London 220:1± star larva of Cassida nebulosa L. (Coleoptera: 32. Chrysomelidae: Cassidinae)- a model descrip- Griswold, C.D. 1993. Investigations into the phy- tion. Annales Zoologici 53:189±200. logeny of the lycosoid spiders and their kin Canard, A. 1987. Analyse nouvelle du deÂveloppe- (Arachnida: Araneae: Lycosoidea). Smithsonian ment postembryonnaire des araigneÂes. Revue Ar- Contributions to Zoology 539:1±39. achnologique 7:91±128. Hallas, S.E.A. 1988. Hatching and early postem- Chamberlin, J.C. 1931. The order Chelo- nethida. Stanford University Publications Uni- bryonic development in the Salticidae. Bulletin versity Series, Biological Sciences 7:1±284. of British Arachnological Society 7:231±236. Deruaz, D., J. Deruaz & J. Pichot. 1991. Corre- Harvey, M.S. 1992. The phylogeny and classi®ca- spondence analysis of larval chaetotaxy in the tion of Pseudoscorpionida (: Arach- ``Anopheles maculipennis complex'' (Diptera, nida). Invertebrate Taxonomy 6:1373±1435. Culicidae). Annales de Parasitologie Humaine et Heimer, S. & W. Nentwig. 1991. Spinnen Mitteleu- Comparee 66:166±172. ropas. Ein Bestimmungsbuch. Paul Parey, Berlin Dondale, C.D. 1986. The subfamilies of wolf spi- and Hamburg. ders (Araneae: Lycosidae). Actas X Congreso In- Hennig, W. 1966. Phylogenetic Systematics. Uni- ternacional de AracnologõÂa, Jaca, EspanÄa 1:327± versity of Illinois Press, Urbana. 332. Holm, AÊ . 1940. Studien uÈber die Entwicklung und TOMASIEWICZ & FRAMENAUÐLARVAL CHAETOTAXY IN WOLF SPIDERS 425

Entwicklungsbiologie der Spinnen. Zoologiska 1879) (Araneae: Linyphiidae). Genus 14:585± Bidrag fran Uppsala 19:1±289. 602. Kilian, A. 1998. Morphology and phylogeny of the Rybak, J. & B. Tomasiewicz. 2005. The larval larval stages of the tribe Agathidiini (Coleoptera: chaetotaxy of Bathyphantes eumenis (L. Koch Leiodidae: Leiodinae). Annales Zoologici 48: 1879) (Araneae: Linyphiidae) and Trochosa rur- 125±220. icola (De Geer 1778) (Araneae: Lycosidae)Ða Kitching, I.J. 1984. The use of larval chaetotaxy in model description. Genus 16:129±143. butter¯y systematics, with special reference to Scioscia, C.L. 1992. EvolucioÂn de la tricobotriotax- the Danaini (Lepidoptera: Nymphalidae). Sys- ia durante el desarrollo de tres Salticidae neo- tematic Entomology 9:49±61. tropicales (Araneae). EOS 68:99±206. Kitching, I.J. 1985. Early stages and the classi®ca- Tuzovsky, P.V. 1987. Morphology and Postembry- tion of the milkweed butter¯ies (Lepidoptera: onic Development of Water Mites. Nauka Pub- Danainae). Zoological Journal of the Linnean lications, Moscow. Society 85:1±97. Vachon, M. 1957. Contribution a l'eÂtude du deÂvel- Lethinen, P. 1980. Trichobothrial patterns in high oppement postembryonnaire des araigneÂes. 1ere level taxonomy of spiders. Proceedings of the note: geÂneÂraliteÂs et nomenclature des stades. Bul- VIIIth International Arachnological Congress, letin de la SocieÂte Zoologique de France 82:337± Vienna. Pp. 493±498. 354. Mabee, P.M. 2000. The usefulness of ontogeny in Vink, C.J., A.D. Mitchell & A.M. Paterson. 2002. interpreting morphological characters. Pp. 84± A preliminary molecular analysis of phylogenet- 114. In Phylogenetic Analysis of Morphological ic relationships of Australasian wolf spider gen- Data. (J.J. Wiens, ed.). Smithsonian Institution era (Araneae, Lycosidae). Journal of Arachnol- Press, Washington and London. ogy 30:227±237. Murphy, N.P., V.W. Framenau, S.C. Donnellan, Wiehle, H. 1960. Spinnentiere oder Arachnoidea M.S. Harvey, Y.-C. Park & A.D. Austin. In (Araneae). XI. MicryphantidaeÐZwergspinnen. press. Phylogenetic reconstruction of the wolf Pp. 1±620. In Die Tierwelt Deutschlands und der spiders (Araneae, Lycosidae) using sequences angrenzenden Meeresteile nach ihren Merkmalen from 12S rRNA, 28S rRNA and NADH1 genes: und nach ihrer Lebensweise. 47. Teil (Dahl, F., implications for classi®cation, biogeography and M. Dahl & H. Bischoff, eds.). Gustav Fischer the evolution of web-building behavior. Molec- Verlag, Jena ular Phylogenetics and Evolution. Zehethofer, K. & C. Sturmbauer. 1998. Phylogenetic Nayrolles, P. & J.M. Betsch. 1993. Pour une theÂorie relationships of Central European wolf spiders de la description cheÂtotaxique chez les collem- (Araneae: Lycosidae) inferred from 12S ribosomal boles. Annales de la SocieÂte Entomologique de DNA sequences. Molecular Phylogenetics and France (N.S.) 29:5±15. Evolution 10:391±398. Nelson, G.J. 1978. Ontogeny, phylogeny, palaeon- Zyuzin, A.A. 1993. Studies on wolf spiders (Ara- tology, and the biogenetic law. Systematic Zo- neae: Lycosidae). I. A new genus and species ology 27:324±345. from Kazakhstan, with comments on the Lycos- Pomorski, R.J. 1996. The ®rst instar larvae of On- inae. Memoirs of the Queensland Museum 33: ychiurinaeÐa systematic study (Collembola: 693±700. Onychiuridae). Genus 7:1±102. Rybak, J. & R.J. Pomorski. 2003. Morphology of Manuscript received 4 January 2005, revised 17 protonymph of Bathyphantes eumenis (L. Koch, June 2005.