Atlas of phylogenetic data for entelegyne spiders (Araneae: Araneomorphae: Entelegynae) with comments on their phylogeny

2 Introduction 2 Taxon choice 3 Conventions 5 Materials and methods 5 Specimen preparation 6 Acknowledgments 7 Results 8 The Entelegyne exemplars 8 Agelenidae 8 Amaurobiidae 9 Amphinectidae 12 Araneoidea 14 Archaeidae 14 Austrochilidae 15 Ctenidae 17 Deinopidae 18 Desidae 19 Dictynidae 21 Eresidae 24 Filistatidae 27 Gradungulidae, Huttoniidae 28 Hypochilidae 29 Mimetidae 30 Neolanidae 31 Nicodamidae 32 Oecobiidae 33 Pararchaeidae 34 Phyxelididae 35 Psechridae 37 Segestriidae 39 Stiphidiidae 40 Tengellidae 41 2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, Supplement II Titanoecidae 42 Uloboridae 43 Zorocratidae 44 Zoropsidae 46 Characters 47 Discussion 70 Character systems and homoplasy levels 71 Groups 71 Conclusions 76 Literature cited 76 Appendicies 83 Appendix 1. Taxa examined to provide exemplar data 85 Appendix 2. Data matrix 94 Appendix 3. Comments on data matrix 97 Illustrations 101 Index: Systematic and Geographic 319 We present a phylogenetic analysis of higher groups of entelegyne spiders, with representatives of all entelegyne families containing cribellate members and of Palpimanoidea. We examined 55 exemplar species of eresoids (Oecobiidae, Eresidae), Orbiculariae (Deinopidae, Uloboridae, Araneidae), Palpimanoidea (Archaeidae, Huttoniidae, Mimetidae, Pararchaeidae), titanoecoids (Phyxelididae, Titanoecidae), lycosoids and related groups (Ctenidae, Psechridae, Tengellidae, Zorocratidae, Zoropsidae), and several other entelegyne families of contentious relationships (Agelenidae, Amphinectidae, Desidae, Neolanidae, Stiphidiidae, Amaurobiidae, Dictynidae, Nicodamidae), plus representatives of the relatively basal araneomorph groups Palaeocribellatae (Hypochilidae), Austrochiloidea (Austrochilidae and Gradungulidae), and Haplogynae (Filistatidae, Segestriidae). We also included the enigmatic cribellate genera Aebutina (Dictynidae? ) and Poaka (Psechridae? Amaurobiidae? ). This selection of taxa covers much of the morphological diversity found through major groups of araneomorph spiders. The 154 characters in our dataset include all the classical character sources in higher level spider taxonomy. We present a large collection of labeled images to document character definitions and observations. The phylogenetic trees obtained with our dataset differ according to parameters of analysis (equal or implied weights), and diverge from previous results. We obtain a paraphyletic Araneoclada, excluding members of Haplogynae. At least some taxa previously assigned to Palpimanoidea appear to be nested within orb weavers. The outgroups to Orbiculariae remain an open question, and the monophyly of Nicodamidae, Amaurobiidae and Zorocratidae are questioned. We corroborated Austrochiloidea, Eresoidea, and the Divided Cribellum, Oval Calamistrum and RTA Clades. The Entelegynae comprise the largest group of spiders with more than 38,000 described species (Platnick 2004). Ideas about entelegyne spider evolution were long dominated by the fauna of the northern hemisphere, especially Europe. This impoverished fauna includes distinct and distantly related taxa, most of which are ecribellate or colulate. Spiders with cribella, the remarkable spinning plate that works in conjunction with a comb on the fourth leg to make a "hackled band" capture thread, form a small subset of this northern fauna. For nearly a century the higher spiders were divided into two groups: those with cribella, and those without. All this changed in the late 20 century with global taxonomic studies of higher spiders (Lehtinen 1967), intense studies of the spiders of the southern hemisphere (Forster 1970; Forster and Wilton 1973), and application of Pfennig's cladistic principles to spider taxonomy (e.g., Platnick and Gertsch 1976; Platnick 1977). GRISWOLD, RAMIREZ, CODDINGTON, & PLATNICK: ENTELEGYNE SPIDER DATA 3 The cribellum was reinterpreted as a primitive feature common to all higher spiders, albeit, for most, in greatly modified form. The cribellum is clearly more ancient than even the entelegyne condition, i.e., that of having separate copulatory and fertilization ducts. Entelegynes have repeatedly been the subject of quantitative phylogenetic analysis. The earliest studies focused on clades of special interest, e.g., orb weavers (Coddington 1990a, 1990b; Griswold et al. 1998, Fig. 211), haplogynes (Platnick et al. 1991, Fig. 209), lycosoids (Griswold 1993, Fig. 213). The first attempt at a comprehensive entelegyne phylogeny was by Griswold et al. (1999), who chose and coded cribellate members from families across the araneomorph spectrum. Their approach was guided by the words of Lehtinen (1967:202) who declared, "because of the central position of the Cribellate groups in Araneomorphae, a detailed revision of them is a short cut to a rough classification of the whole suborder. " They chose exemplars from all cribellate families, reasoning that taxa retaining this plesiomorphic feature are more likely to straddle the basal nodes of the phylogeny of higher groups than are their relatives that have lost the cribellum: therefore they are most likely to reflect phylogenetic groundplans. Although phylogenetically ancient, the cribellum is a complex feature unlikely to have evolved more than once. Most major araneomorph clades have cribellate members (exceptions are Palpimanoidea and Dionycha). A phylogeny of these basal taxa should mirror the relationships of the large clades they exemplify. The provisional phylogeny of Griswold et al. (1999) tested many suprafamilial hypotheses of the last 30 years and was the first attempt to relate them using quantitative phylogenetic techniques. The cladogram confirmed some accepted groupings, refuted others, and several novel phylogenetic and nomenclatural changes were proposed (Fig. 212). Confirmed were the monophyly of Neocribellatae, Araneoclada, Entelegynae, and Orbiculariae. The Lycosoidea, Amaurobiidae and some included subfamilies, Dictynoidea, and Amaurobioidea (sensu Forster and Wilton 1973) appeared polyphyletic. Phyxelididae Lehtinen was raised to family level and Zorocratidae Dahl was revalidated. A group including all other entelegynes other than Eresoidea was weakly supported as the sister group of Orbiculariae and several new, informative, informal clades were proposed or redefined: the "Canoe Tapetum Clade," "Divided Cribellum Clade," the "Titanoecoids," the "RTA Clade," the "Fused Paracribellar Clade," the "Stiphidioids" and the "Agelenoids. " This slender paper (Griswold et al. 1999), constrained by publication in a congress volume, offered only the briefest outline of the data. Griswold and Wang (2001) presented a fuller account of the results, presenting character state trees for each of the 137 characters and figures depicting many of the character states. In this study we will present and illustrate in detail the morphology and other characteristics of the exemplar taxa, explain the character coding, and discuss some implications of our trees for spider evolution. We hope that this paper, especially the new data presented herein, will provide a springboard to further, more detailed and more comprehensive analyses of araneomorph phylogeny. This paper is dedicated to the memory of Ray Forster.