We present a phylogenetic analysis of spiders using a dataset of 932 spider species, representing 115 families (only the family Synaphridae is unrepresented), 700 known genera, and additional representatives of 26 unidentified or undescribed genera. Eleven genera of the orders Amblypygi, Palpigradi, Schizomida and Uropygi are included as outgroups. The dataset includes six markers from the mitochondrial (12S, 16S, COI) and nuclear (histone H3, 18S, 28S) genomes, and was analysed by multiple methods, including constrained analyses using a highly supported backbone tree from transcriptomic data. We recover most of the higher‐level structure of the spider tree with good support, including Mesothelae, Opisthothelae, Mygalomorphae and Araneomorphae. Several of our analyses recover Hypochilidae and Filistatidae as sister groups, as suggested by previous transcriptomic analyses. The Synspermiata are robustly supported, and the families Trogloraptoridae and Caponiidae are found as sister to the Dysderoidea. Our results support the Lost Tracheae clade, including Pholcidae, Tetrablemmidae, Diguetidae, Plectreuridae and the family Pacullidae (restored status) separate from Tetrablemmidae. The Scytodoidea include Ochyroceratidae along with Sicariidae, Scytodidae, Drymusidae and Periegopidae; our results are inconclusive about the separation of these last two families. We did not recover monophyletic Austrochiloidea and Leptonetidae, but our data suggest that both groups are more closely related to the Cylindrical Gland Spigot clade rather than to Synspermiata. Palpimanoidea is not recovered by our analyses, but also not strongly contradicted. We find support for Entelegynae and Oecobioidea (Oecobiidae plus Hersiliidae), and ambiguous placement of cribellate orb‐weavers, compatible with their non‐monophyly. Nicodamoidea (Nicodamidae plus Megadictynidae) and Araneoidea composition and relationships are consistent with recent analyses. We did not obtain resolution for the titanoecoids (Titanoecidae and Phyxelididae), but the Retrolateral Tibial Apophysis clade is well supported. Penestomidae, and probably Homalonychidae, are part of Zodarioidea, although the latter family was set apart by recent transcriptomic analyses. Our data support a large group that we call the marronoid clade (including the families Amaurobiidae, Desidae, Dictynidae, Hahniidae, Stiphidiidae, Agelenidae and Toxopidae). The circumscription of most marronoid families is redefined here. Amaurobiidae include the Amaurobiinae and provisionally Macrobuninae. We transfer Malenellinae (Malenella, from Anyphaenidae), Chummidae (Chumma) (new syn.) and Tasmarubriinae (Tasmarubrius, Tasmabrochus and Teeatta, from Amphinectidae) to Macrobuninae. Cybaeidae are redefined to include Calymmaria, Cryphoeca, Ethobuella and Willisius (transferred from Hahniidae), and Blabomma and Yorima (transferred from Dictynidae). Cycloctenidae are redefined to include Orepukia (transferred from Agelenidae) and Pakeha and Paravoca (transferred from Amaurobiidae). Desidae are rede...
Understanding the web construction behaviour of theridiid (comb-footed) spiders is fundamental to formulating specific evolutionary hypotheses and predictions regarding the reduction of orb-webs. We describe for the first time in detail the web construction behaviour of Achaearanea tepidariorum , Latrodectus geometricus , Theridion sisyphium and T. varians as well as webs of a range of other theridiids. In our survey we distinguish four major web types. Among webs with gumfooted lines, we distinguish between webs with a central retreat ( Achaearanea -type) and those with a peripheral retreat ( Latrodectus -type). Among webs without gumfooted lines, we distinguish between those which contain viscid silk ( Theridion -type) and those with a sheet-like structure, which do not ( Coleosoma -type). Theridiid gumfoot-webs consist of frame lines that anchor them to surroundings and support threads which possess viscid silk. Building of gumfooted lines constitutes a unique stereotyped behaviour and is most probably homologous for Nesticidae and Theridiidae. Webs remained in place for extended periods and were expanded and repaired, but no regular pattern of replacement was observed. We suggest that the cost of producing and maintaining viscid silk might have led to web reduction, at least in theridiids.ADDITIONAL KEYWORDS: behavioural patternscharacter evolutioncapture threadresource allocation viscid silkweb constructionweb reduction. Figure 8. Schematic representation of theridiid webs (not to scale): A, Achaearanea-type web with a central retreat. B, Latrodectus-web with a peripheral retreat. C, Theridion-type web with viscid elements. D, Coleosoma-type without viscid elements but with a sheet and KN structure.Benjamin SP, Düggelin M, Zschokke S. 2002. Fine structure of sheet-webs of Linyphia triangularis (Clerck) and Microlinyphia pusilla (Sundevall), with remarks on the presence of viscid silk. Acta Zoologica 83: 49-59. Benjamin SP, Zschokke S. 2002. A computerised method to observe spider web building behaviour in a semi-natural light environment. In: Toft S, Scharff N, eds. European Arachnology 2000. Aarhus, Denmark: University of Aarhus Press, 177-122. Breed AL, Levine VD, Peakall DB, Witt PN. 1964. The fate of the intact orb web of the spider Araneus diadematus Cl. Behaviour 23: 43-60. Bristowe WS. 1958. The world of spiders. London: Collins. Carico JE. 1986. Web removal patterns in orb-weaving spiders. In: Shear WA, ed. Spiders -webs, behavior, and evolution. Stanford: Stanford University Press, 306-318. Coddington JA. 1986. Orb webs in non orb weaving ogrefaced spiders (Araneae: Dinopidae): a question of genealogy. Cladistics 2: 53-67. Comstock JH. 1940. The spider book. Ithaca, NY: Comstock Pub. Assoc. Eberhard WG. 1972. Observations on the biology of Achaearanea tesselata (Araneae: Theridiidae). Psyche: A Journal of Entomology, Boston 78: 209-212. Eberhard WG. 1979. Argyrodes attenuatus (Theridiidae): a web that is not a snare. Psyche: A Journal of Entomology, Boston 86: 407-413. Eberhard WG. 1981. The single ...
Images are paramount in documentation of morphological data. Production and reproduction costs have traditionally limited how many illustrations taxonomy could afford to publish, and much comparative knowledge continues to be lost as generations turn over. Now digital images are cheaply produced and easily disseminated electronically but pose problems in maintenance, curation, sharing, and use, particularly in long-term data sets involving multiple collaborators and institutions. We propose an efficient linkage of images to phylogenetic data sets via an ontology of morphological terms; an underlying, fine-grained database of specimens, images, and associated metadata; fixation of the meaning of morphological terms (homolog names) by ostensive references to particular taxa; and formalization of images as standard views. The ontology provides the intellectual structure and fundamental design of the relationships and enables intelligent queries to populate phylogenetic data sets with images. The database itself documents primary morphological observations, their vouchers, and associated metadata, rather than the conventional data set cell, and thereby facilitates data maintenance despite character redefinition or specimen reidentification. It minimizes reexamination of specimens, loss of information or data quality, and echoes the data models of web-based repositories for images, specimens, and taxonomic names. Confusion and ambiguity in the meanings of technical morphological terms are reduced by ostensive definitions pointing to features in particular taxa, which may serve as reference for globally unique identifiers of characters. Finally, the concept of standard views (an image illustrating one or more homologs in a specific sex and life stage, in a specific orientation, using a specific device and preparation technique) enables efficient, dynamic linkage of images to the data set and automatic population of matrix cells with images independently of scoring decisions.
The subfamily Ballinae is revised. To test its monophyly, 41 morphological characters, including the first phylogenetic use of scale morphology in Salticidae, were scored for 16 taxa (1 outgroup and 15 ingroup). Parsimony analysis of these data supports monophyly based on five unambiguous synapomorphies. The paper provides new diagnoses, descriptions of new genera, species, and a key to the genera. At present, Ballinae comprises 13 nominal genera, three of them new: Afromarengo, Ballus, Colaxes, Cynapes, Indomarengo, Leikung, Marengo, Philates and Sadies. Copocrossa, Mantisatta, Pachyballus and Padilla are tentatively included in the subfamily. Nine new species are described and illustrated: Colaxes horton, C. wanlessi, Philates szutsi, P. thaleri, P. zschokkei, Indomarengo chandra, I. sarawakensis, Leikung kinabaluensis and Marengo deelemanae. Colaxes nitidiventris and Ballus segmentatus are redescribed. The Seychelles species Baviola spatulata is considered a junior synonym of B. braueri. The phylogenetic analysis suggests two independent origins of tentative Batesian mimicry of ants within Ballinae.
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