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Haplogynae
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<h2 id="phylogeny">Phylogeny</h2> <p class="note">Main article: <a href="/facts/Spider_taxonomy/7dwD8Qvk">Spider taxonomy</a></p> <p>The Haplogynae are one of the two major groups into which araneomorph spiders were traditionally divided, the other being the Entelegynae. In 2005, <a href="/facts/Jonathan_A._Coddington/30aCaRIW">Coddington</a> summarized the relationships of these groups as suggested by morphological <a href="/facts/Phylogenetics/bw4rsovt">phylogenetic studies</a>:<a class="footnote-ref" id="fnref:2" href="#fn:2"><sup>2</sup></a> </p> <table><tbody><tr><td><a href="/facts/Araneomorphae/tN5eRbnK">Araneomorphae</a></td><td rowspan="2"><table><tbody><tr><td>Paleocribellatae</td><td rowspan="2"><p><a href="/facts/Hypochilidae/EMuLw7SG">Hypochilidae</a></p></td></tr><tr><td></td></tr><tr><td>Neocribellatae</td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><a href="/facts/Austrochiloidea/nCS9dFpa">Austrochiloidea</a></p></td></tr><tr><td></td></tr><tr><td>Araneoclada</td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>Haplogynae</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Entelegynae/22sKpTIm">Entelegynae</a></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table> <p>Subsequent molecular phylogenetic studies have confirmed the <a href="/facts/Monophyly/pD8Yrrn6">monophyly</a> of some of the groups suggested by morphological studies, while rejecting many others.<a class="footnote-ref" id="fnref:3" href="#fn:3"><sup>3</sup></a><a class="footnote-ref" id="fnref:4" href="#fn:4"><sup>4</sup></a><a class="footnote-ref" id="fnref:5" href="#fn:5"><sup>5</sup></a> </p><p>A study published in 2015 suggested that two families formerly placed in the Haplogynae do not belong there. <a href="/facts/Filistatidae/fK7DdJoA">Filistatidae</a> groups with <a href="/facts/Hypochilidae/EMuLw7SG">Hypochilidae</a> at the base of the Haplogynae; <a href="/facts/Leptonetidae/nqucjfBA">Leptonetidae</a> is basal to the Entelegynae.<a class="footnote-ref" id="fnref:6" href="#fn:6"><sup>6</sup></a> The similarity of some morphological features of Leptonetidae to those of entelegynes had already been noted.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> </p> <table><tbody><tr><td><table><tbody><tr><td>Araneomorphae</td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>Hypochilidae</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p>Filistatidae</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p>remaining Haplogynae</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>Leptonetidae</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p>Entelegynae</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td><td>Haplogynae sensu Coddington (2005)</td></tr></tbody></table> <p>In 2016, a large <a href="/facts/Molecular_phylogenetics/FysplHP5">molecular phylogenetic</a> study was published online that included 932 spider species, representing all but one of the then known families. It "refutes important higher-level groups",<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a> including Paleocribellatae, Neocribellatae, Araneoclada and Haplogynae. In the preferred cladogram, the "Haplogynae" are divided among a number of clades basal to the Entelegynae, forming at most a grade. "Haplogynae" in the sense of Coddington (2005) are shaded yellow in the cladogram below; Entelegynae in the same sense are shaded blue. The clade <a href="/facts/Synspermiata/M0PxnkAh">Synspermiata</a> comprises all the ecribellate haplogynes and is consistently recovered, but with low support.<a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a> </p> <table><tbody><tr><td><a href="/facts/Araneomorphae/tN5eRbnK">Araneomorphae</a></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><a href="/facts/Hypochilidae/EMuLw7SG">Hypochilidae</a></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Filistatidae/fK7DdJoA">Filistatidae</a></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Synspermiata/M0PxnkAh">Synspermiata</a> (ecribellate haplogynes)</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Hickmania/uls9SMHw">Hickmania</a></i> (<a href="/facts/Austrochilidae/bASmT8zK">Austrochilidae</a>)</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i><a href="/facts/Archoleptoneta/Xk8SUpts">Archoleptoneta</a></i> (<a href="/facts/Leptonetidae/nqucjfBA">Leptonetidae</a>)</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Gradungulidae/z7UjBx8J">Gradungulidae</a></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Leptoneta/RbrtPuhq">Leptoneta</a></i> (Leptonetidae)</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i><a href="/facts/Austrochilus/mKpCyCkr">Austrochilus</a></i> + <i><a href="/facts/Thaida/6Ppnlqc7">Thaida</a></i> (Austrochilidae)</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><a href="/facts/Palpimanoidea/Wab7jfer">Palpimanoidea</a> (paraphyletic in molecular analyses)</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>Neoleptoneta</i> + <i>Calileptoneta</i> (Leptonetidae)</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Entelegynae/22sKpTIm">Entelegynae</a> <i>sensu</i> Wheeler et al. (2017)</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table> <h2 id="families">Families</h2> <p>As shown above, <a href="/facts/Filistatidae/fK7DdJoA">Filistatidae</a> and <a href="/facts/Leptonetidae/nqucjfBA">Leptonetidae</a> are placed outside the traditional haplogynes in the analysis by Wheeler et al. (2017). Traditional haplogyne families they place in Synspermiata are:<a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a> </p> <ul><li><a href="/facts/Caponiidae/SJAdTjhc">Caponiidae</a></li> <li><a href="/facts/Diguetidae/mfPk8DJ8">Diguetidae</a></li> <li><a href="/facts/Drymusidae/p2rsyvmq">Drymusidae</a></li> <li><a href="/facts/Dysderidae/U7VChKwm">Dysderidae</a></li> <li><a href="/facts/Ochyroceratidae/rfIgGkdk">Ochyroceratidae</a></li> <li><a href="/facts/Oonopidae/wrD6N5bd">Oonopidae</a></li> <li><a href="/facts/Orsolobidae/f5pRnXTs">Orsolobidae</a></li> <li><a href="/facts/Pacullidae/6jDXh2TJ">Pacullidae</a></li> <li><a href="/facts/Periegopidae/GeJVbjwh">Periegopidae</a></li> <li><a href="/facts/Pholcidae/azdPyEsr">Pholcidae</a></li> <li><a href="/facts/Plectreuridae/xcE4qlS1">Plectreuridae</a></li> <li><a href="/facts/Scytodidae/w8v30t8C">Scytodidae</a></li> <li><a href="/facts/Segestriidae/czdkNL2T">Segestriidae</a></li> <li><a href="/facts/Sicariidae/D1dfF8a0">Sicariidae</a></li> <li><a href="/facts/Tetrablemmidae/2kbrsVEv">Tetrablemmidae</a></li></ul> <p><a href="/facts/Telemidae/HTaEPlEm">Telemidae</a>, traditionally placed in Haplogynae, was not included in the analysis by Wheeler et al.<a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a> However, it is placed in Synspermiata in other studies.<a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a> The recently discovered haplogyne family <a href="/facts/Trogloraptoridae/KFwJIQ2z">Trogloraptoridae</a><a class="footnote-ref" id="fnref:13" href="#fn:13"><sup>13</sup></a> was also placed in Synspermiata.<a class="footnote-ref" id="fnref:14" href="#fn:14"><sup>14</sup></a> </p><p>Several extinct families have also been placed into the haplogynes. </p> <ul><li>†Pholcochyroceridae</li></ul> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Gertsch, Willis John (1958). The spider family Plectreuridae. Vol. 1920. American Museum of Natural History. <a href="https://www.biodiversitylibrary.org/page/63417590" target="_blank">https://www.biodiversitylibrary.org/page/63417590</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Coddington, Jonathan A. (2005). "Phylogeny and classification of spiders" (PDF). In Ubick, D.; Paquin, P.; Cushing, P.E. & Roth, V. (eds.). Spiders of North America: an identification manual. American Arachnological Society. pp. 18–24. Retrieved 2015-09-24. <a href="https://repository.si.edu/bitstream/handle/10088/4365/CoddingtonSNAPhylogeny05.pdf" target="_blank">https://repository.si.edu/bitstream/handle/10088/4365/CoddingtonSNAPhylogeny05.pdf</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>Agnarsson, Ingi; Coddington, Jonathan A. & Kuntner, Matjaž (2013). "Systematics : Progress in the study of spider diversity and evolution". In Penney, David (ed.). Spider research in the 21st century: trends & perspectives. Manchester, UK: Siri Scientific Press. ISBN 978-0-9574530-1-2. pp. 82–83. <a href="978-0-9574530-1-2" target="_blank">978-0-9574530-1-2</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>Garrison, Nicole L.; Rodriguez, Juanita; Agnarsson, Ingi; Coddington, Jonathan A.; Griswold, Charles E.; Hamilton, Christopher A.; Hedin, Marshal; Kocot, Kevin M.; Ledford, Joel M. & Bond, Jason E. (2015). "Spider phylogenomics: untangling the Spider Tree of Life". PeerJ. 3: e1852. doi:10.7717/peerj.1719. PMC 4768681. PMID 26925338. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4768681" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4768681</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>Wheeler, Ward C.; Coddington, Jonathan A.; Crowley, Louise M.; Dimitrov, Dimitar; Goloboff, Pablo A.; Griswold, Charles E.; Hormiga, Gustavo; Prendini, Lorenzo; Ramírez, Martín J.; Sierwald, Petra; Almeida-Silva, Lina; Alvarez-Padilla, Fernando; Arnedo, Miquel A.; Benavides Silva, Ligia R.; Benjamin, Suresh P.; Bond, Jason E.; Grismado, Cristian J.; Hasan, Emile; Hedin, Marshal; Izquierdo, Matías A.; Labarque, Facundo M.; Ledford, Joel; Lopardo, Lara; Maddison, Wayne P.; Miller, Jeremy A.; Piacentini, Luis N.; Platnick, Norman I.; Polotow, Daniele; Silva-Dávila, Diana; Scharff, Nikolaj; Szűts, Tamás; Ubick, Darrell; Vink, Cor J.; Wood, Hannah M. & Zhang, Junxia (2017) [published online 2016], "The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling", Cladistics, 33 (6): 574–616, doi:10.1111/cla.12182, S2CID 35535038 <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></p></li> <li id="fn:6"><p>Garrison, Nicole L.; Rodriguez, Juanita; Agnarsson, Ingi; Coddington, Jonathan A.; Griswold, Charles E.; Hamilton, Christopher A.; Hedin, Marshal; Kocot, Kevin M.; Ledford, Joel M. & Bond, Jason E. (2015). "Spider phylogenomics: untangling the Spider Tree of Life". PeerJ. 3: e1852. doi:10.7717/peerj.1719. PMC 4768681. PMID 26925338. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4768681" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4768681</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></p></li> <li id="fn:7"><p>Ledford, Joel M. & Griswold, Charles E. (2010), "A study of the subfamily Archoleptonetinae (Araneae, Leptonetidae) with a review of the morphology and relationships for the Leptonetidae" (PDF), Zootaxa, 2391: 1–32, doi:10.11646/zootaxa.2391.1.1, retrieved 2016-01-09 <a href="http://mapress.com/zootaxa/2010/f/z02391p032f.pdf" target="_blank">http://mapress.com/zootaxa/2010/f/z02391p032f.pdf</a> <a href="#fnref:7" class="footnote-back-ref">↩</a></p></li> <li id="fn:8"><p>Wheeler, Ward C.; Coddington, Jonathan A.; Crowley, Louise M.; Dimitrov, Dimitar; Goloboff, Pablo A.; Griswold, Charles E.; Hormiga, Gustavo; Prendini, Lorenzo; Ramírez, Martín J.; Sierwald, Petra; Almeida-Silva, Lina; Alvarez-Padilla, Fernando; Arnedo, Miquel A.; Benavides Silva, Ligia R.; Benjamin, Suresh P.; Bond, Jason E.; Grismado, Cristian J.; Hasan, Emile; Hedin, Marshal; Izquierdo, Matías A.; Labarque, Facundo M.; Ledford, Joel; Lopardo, Lara; Maddison, Wayne P.; Miller, Jeremy A.; Piacentini, Luis N.; Platnick, Norman I.; Polotow, Daniele; Silva-Dávila, Diana; Scharff, Nikolaj; Szűts, Tamás; Ubick, Darrell; Vink, Cor J.; Wood, Hannah M. & Zhang, Junxia (2017) [published online 2016], "The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling", Cladistics, 33 (6): 574–616, doi:10.1111/cla.12182, S2CID 35535038 <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></p></li> <li id="fn:9"><p>Wheeler, Ward C.; Coddington, Jonathan A.; Crowley, Louise M.; Dimitrov, Dimitar; Goloboff, Pablo A.; Griswold, Charles E.; Hormiga, Gustavo; Prendini, Lorenzo; Ramírez, Martín J.; Sierwald, Petra; Almeida-Silva, Lina; Alvarez-Padilla, Fernando; Arnedo, Miquel A.; Benavides Silva, Ligia R.; Benjamin, Suresh P.; Bond, Jason E.; Grismado, Cristian J.; Hasan, Emile; Hedin, Marshal; Izquierdo, Matías A.; Labarque, Facundo M.; Ledford, Joel; Lopardo, Lara; Maddison, Wayne P.; Miller, Jeremy A.; Piacentini, Luis N.; Platnick, Norman I.; Polotow, Daniele; Silva-Dávila, Diana; Scharff, Nikolaj; Szűts, Tamás; Ubick, Darrell; Vink, Cor J.; Wood, Hannah M. & Zhang, Junxia (2017) [published online 2016], "The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling", Cladistics, 33 (6): 574–616, doi:10.1111/cla.12182, S2CID 35535038 <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>Wheeler, Ward C.; Coddington, Jonathan A.; Crowley, Louise M.; Dimitrov, Dimitar; Goloboff, Pablo A.; Griswold, Charles E.; Hormiga, Gustavo; Prendini, Lorenzo; Ramírez, Martín J.; Sierwald, Petra; Almeida-Silva, Lina; Alvarez-Padilla, Fernando; Arnedo, Miquel A.; Benavides Silva, Ligia R.; Benjamin, Suresh P.; Bond, Jason E.; Grismado, Cristian J.; Hasan, Emile; Hedin, Marshal; Izquierdo, Matías A.; Labarque, Facundo M.; Ledford, Joel; Lopardo, Lara; Maddison, Wayne P.; Miller, Jeremy A.; Piacentini, Luis N.; Platnick, Norman I.; Polotow, Daniele; Silva-Dávila, Diana; Scharff, Nikolaj; Szűts, Tamás; Ubick, Darrell; Vink, Cor J.; Wood, Hannah M. & Zhang, Junxia (2017) [published online 2016], "The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling", Cladistics, 33 (6): 574–616, doi:10.1111/cla.12182, S2CID 35535038 <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>Wheeler, Ward C.; Coddington, Jonathan A.; Crowley, Louise M.; Dimitrov, Dimitar; Goloboff, Pablo A.; Griswold, Charles E.; Hormiga, Gustavo; Prendini, Lorenzo; Ramírez, Martín J.; Sierwald, Petra; Almeida-Silva, Lina; Alvarez-Padilla, Fernando; Arnedo, Miquel A.; Benavides Silva, Ligia R.; Benjamin, Suresh P.; Bond, Jason E.; Grismado, Cristian J.; Hasan, Emile; Hedin, Marshal; Izquierdo, Matías A.; Labarque, Facundo M.; Ledford, Joel; Lopardo, Lara; Maddison, Wayne P.; Miller, Jeremy A.; Piacentini, Luis N.; Platnick, Norman I.; Polotow, Daniele; Silva-Dávila, Diana; Scharff, Nikolaj; Szűts, Tamás; Ubick, Darrell; Vink, Cor J.; Wood, Hannah M. & Zhang, Junxia (2017) [published online 2016], "The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling", Cladistics, 33 (6): 574–616, doi:10.1111/cla.12182, S2CID 35535038 <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:11" class="footnote-back-ref">↩</a></p></li> <li id="fn:12"><p>Michalik, Peter & Ramírez, Martín J. (2014), "Evolutionary morphology of the male reproductive system, spermatozoa and seminal fluid of spiders (Araneae, Arachnida)–Current knowledge and future directions", Arthropod Structure & Development, 43 (4): 291–322, doi:10.1016/j.asd.2014.05.005, hdl:11336/19081, PMID 24907603 <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:12" class="footnote-back-ref">↩</a></p></li> <li id="fn:13"><p>Griswold, C.; Audisio, T.; Ledford, J. (2012). "An extraordinary new family of spiders from caves in the Pacific Northwest (Araneae, Trogloraptoridae, new family)". ZooKeys (215): 77–102. doi:10.3897/zookeys.215.3547. PMC 3428790. PMID 22936872. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3428790" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3428790</a> <a href="#fnref:13" class="footnote-back-ref">↩</a></p></li> <li id="fn:14"><p>Wheeler, Ward C.; Coddington, Jonathan A.; Crowley, Louise M.; Dimitrov, Dimitar; Goloboff, Pablo A.; Griswold, Charles E.; Hormiga, Gustavo; Prendini, Lorenzo; Ramírez, Martín J.; Sierwald, Petra; Almeida-Silva, Lina; Alvarez-Padilla, Fernando; Arnedo, Miquel A.; Benavides Silva, Ligia R.; Benjamin, Suresh P.; Bond, Jason E.; Grismado, Cristian J.; Hasan, Emile; Hedin, Marshal; Izquierdo, Matías A.; Labarque, Facundo M.; Ledford, Joel; Lopardo, Lara; Maddison, Wayne P.; Miller, Jeremy A.; Piacentini, Luis N.; Platnick, Norman I.; Polotow, Daniele; Silva-Dávila, Diana; Scharff, Nikolaj; Szűts, Tamás; Ubick, Darrell; Vink, Cor J.; Wood, Hannah M. & Zhang, Junxia (2017) [published online 2016], "The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling", Cladistics, 33 (6): 574–616, doi:10.1111/cla.12182, S2CID 35535038 <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:14" class="footnote-back-ref">↩</a></p></li> </ol>