Atlas of phylogenetic data for entelegyne spiders (Araneae: Araneomorphae: Entelegynae) with comments on their phylogeny
Atlas of phylogenetic data for entelegyne spiders (Araneae: Araneomorphae: Entelegynae) with comments on their phylogeny
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发表时间：
2005
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通讯作者：
C. Griswold;M. Ramírez;J. Coddington;N. Platnick
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作者：
C. Griswold;M. Ramírez;J. Coddington;N. Platnick
2 简介 2 分类群选择 3 惯例 5 材料和方法 5 标本制备 6 致谢 7 结果 8 Entelegyne 范例 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 加州科学院论文集《科学》第 56 卷，补编 II 泰坦科 42 Uloboridae 43 Zorocratidae 44 Zoropsidae 46 性状 47 讨论 70 性状系统和同质性水平 71 类群 71 结论 76 引用文献 76 附录 83 附录 1. 分类单元检查以提供示例数据 85 附录 2. 数据矩阵 94 附录 3. 对数据矩阵的评论 97 插图 101 索引：系统和分类地理 319 我们提出一个 对更高等的 Entelegyne 蜘蛛类群进行系统发育分析，其中包含所有包含丝网成员和 Palpimanoidea 的 Entelegyne 科的代表。我们检查了 55 个 eresoids 范例物种（Oecobiidae、Eresidae）、Orbillaryiae（Deinopidae、Uloboridae、Araneidae）、Palpimanoideae（Archaeidae、Huttoniidae、Mimetidae、Pararchaeidae）、titanoecoids（Phyxelididae、Titanoecidae）、lycosoids 和相关类群 （Ctenidae、Psechridae、Tengellidae、Zorocratidae、Zoropsidae）和其他几个有争议关系的 entelegyne 科（Agelenidae、Amphinectidae、Desidae、Neolanidae、Stiphidiidae、Amaurobiidae、Dictynidae、Nicodamidae），以及相对基础的蜘蛛形群体 Palaeocribellatae 的代表 （Hypochilidae）、Austrochiloidea（Austrochilidae 和 Gradungulidae）和 Haplogynae（Filistatidae、Segestriidae）。我们还包括了神秘的卷草属 Aebutina (Dictynidae?) 和 Poaka (Psechridae? Amaurobiidae?)。这种分类单元的选择涵盖了通过主要蜘蛛类群发现的大部分形态多样性。我们数据集中的 154 个字符包括高级蜘蛛分类学中的所有经典字符源。我们提供了大量标记图像来记录字符定义和观察结果。使用我们的数据集获得的系统发育树根据分析参数（相等或隐含权重）而有所不同，并且与之前的结果有所不同。我们获得了并系的Araneoclada，不包括Haplogynae的成员。至少一些先前归属于帕尔皮马诺总科的类群似乎嵌套在球织虫体内。 Orbulariae 的外群仍然是一个悬而未决的问题，Nicodamidae、Amaurobiidae 和 Zorocratidae 的单系也受到质疑。我们证实了 Austrochiloidea、Eresoidea 和 Divided Cribellum、Oval Calamistrum 和 RTA 分支。 Entelegynae 是最大的蜘蛛类群，描述的物种超过 38,000 个（Platnick 2004）。关于恩特莱吉尼蜘蛛进化的想法长期以来一直由北半球动物群主导，尤其是欧洲。这种贫乏的动物群包括独特且亲缘关系较远的类群，其中大多数是横纹或排列的。带有“cribella”的蜘蛛是这种北方动物群的一小部分，这种蜘蛛是一种引人注目的旋转板，与第四条腿上的梳子一起工作，形成“锯齿带”捕捉线。近一个世纪以来，高等蜘蛛被分为两类：有底纹的蜘蛛和没有底纹的蜘蛛。 20世纪末，随着高等蜘蛛的全球分类学研究（Lehtinen 1967）、对南半球蜘蛛的深入研究（Forster 1970；Forster和Wilton 1973）以及Pfennig的分支原理在蜘蛛分类学中的应用（例如Platnick和Gertsch 1976；Platnick 1977），所有这一切都发生了变化。 GRISWOLD、RAMIREZ、CODDINGTON 和 PLATNICK：ENTELEGYNE 蜘蛛数据 3 筛网被重新解释为所有高等蜘蛛所共有的原始特征，尽管对于大多数蜘蛛来说，其形式已经过很大修改。筛网显然比 entelegyne 条件（即具有独立的交配管和受精管）更古老。 Entelegynes 多次成为定量系统发育分析的主题。最早的研究集中在特别感兴趣的分支，例如，球织虫（Coddington 1990a，1990b；Griswold 等人，1998，图 211）、单合体（Platnick 等人，1991，图 209）、类狼人（Griswold 1993，图 213）。 Griswold 等人首次尝试全面的 entelegyne 系统发育。 (1999)，他从蜘蛛形谱系的家族中选择并编码了鸣叫成员。他们的方法以 Lehtinen (1967:202) 的话为指导，Lehtinen 宣称，“由于 Cribellate 类群在 Araneomorphae 中的中心地位，对它们的详细修改是对整个亚目进行粗略分类的捷径。”他们从所有小笠科中选择了样本，理由是保留这种拟形特征的类群比失去小笠科的近亲更有可能跨越较高类群系统发育的基本节点：因此它们最有可能反映系统发育的基本规划。尽管在系统发育上很古老，但小筛是一个复杂的特征，不太可能进化多次。大多数主要的蜘蛛形分支都有条状成员（掌毛总科和酒神甲除外）。这些基础类群的系统发育应该反映它们所代表的大进化枝的关系。 Griswold 等人的临时系统发育。 (1999) 测试了过去 30 年的许多超家族假说，并首次尝试使用定量系统发育技术将它们联系起来。该分支图证实了一些已被接受的分组，反驳了其他分组，并提出了一些新的系统发育和命名法变化（图212）。确认了Neocribellatae、Araneoclada、Entelegynae 和Orbillaryiae 的单系。石狼总科、Amaurobiidae 和一些包括的亚科、Dictynoidea 和 Amaurobioidea (sensu Forster and Wilton 1973) 似乎是多系的。 Phyxelididae Lehtinen 被提升到科级别，Zorocratidae Dahl 被重新验证。包括除 Eresoidea 之外的所有其他 Entelegynes 的群体作为 Orbulariae 的姐妹群体受到了微弱的支持，并且提出或重新定义了几个新的、信息丰富的、非正式的分支：“Canoe Tapetum Clade”、“Divided Cribellum Clade”、“Titanoecoids”、“RTA Clade”、“Fused Paracribellar Clade”、“Stiphidioids”和 “阿杰雷诺德。”这篇薄薄的论文（Griswold 等人，1999）由于在国会卷中发表的限制，只提供了最简短的数据概要。 Griswold 和 Wang (2001) 对结果进行了更全面的说明，为 137 个字符和描绘了许多字符状态的图形中的每一个都提供了字符状态树。在这项研究中，我们将详细介绍和说明示例类群的形态和其他特征，解释字符编码，并讨论我们的树对蜘蛛进化的一些影响。我们希望本文，特别是本文中提出的新数据，将为进一步、更详细和更全面的蜘蛛形系统发育分析提供一个跳板。谨以此文纪念雷·福斯特。
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.