# Haplogynae

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{{Short description|Infraorder of spiders}}
[[File:Pholcus.phalangioides.6908.jpg|thumb|''[Pholcus phalangioides](/source/Pholcus_phalangioides)'']]
The '''Haplogynae''' or '''haplogynes''' are one of the two main groups into which [araneomorph](/source/Araneomorphae) [spider](/source/spider)s have traditionally been divided, the other being the [Entelegynae](/source/Entelegynae). Morphological phylogenetic studies suggested that the Haplogynae formed a [clade](/source/clade); more recent [molecular phylogenetic studies](/source/Molecular_phylogenetics) refute this, although many of the [ecribellate](/source/Cribellum) haplogynes do appear to form a clade, [Synspermiata](/source/Synspermiata).

Unlike the [Entelegynae](/source/Entelegynae), haplogynes lack hardened (sclerotized) female genitalia ([epigyne](/source/epigyne)s).

Most of the species within this group have six eyes, as opposed to most other spiders. Spiders in the genus ''[Tetrablemma](/source/Tetrablemma)'' ([Tetrablemmidae](/source/Tetrablemmidae)) have only four eyes, as do some members of the family [Caponiidae](/source/Caponiidae); caponiids may even have only two eyes. However, spiders in the family [Plectreuridae](/source/Plectreuridae) have the normal eight eyes.<ref>{{cite book | vauthors=((Gertsch,  Willis John)) | date= 1958 | title=The spider family Plectreuridae | publisher=American Museum of Natural History | volume=1920 | url=https://www.biodiversitylibrary.org/page/63417590}}</ref>

==Phylogeny==
{{Main|Spider taxonomy}}
The Haplogynae are one of the two major groups into which araneomorph spiders were traditionally divided, the other being the Entelegynae. In 2005, [Coddington](/source/Jonathan_A._Coddington) summarized the relationships of these groups as suggested by morphological [phylogenetic studies](/source/Phylogenetics):<ref name=Codd05/>

{{clade |style=line-height:90%;
|label1=[Araneomorphae](/source/Araneomorphae)
|1={{clade
   |label1=Paleocribellatae
   |1=[Hypochilidae](/source/Hypochilidae)
   |label2=Neocribellatae
   |2={{clade
      |1=[Austrochiloidea](/source/Austrochiloidea)
      |label2=Araneoclada
      |2={{clade
         |1='''Haplogynae'''
         |2=[Entelegynae](/source/Entelegynae)
         }}
      }}
   }}
}}

Subsequent molecular phylogenetic studies have confirmed the [monophyly](/source/monophyly) of some of the groups suggested by morphological studies, while rejecting many others.<ref name=AgnaCoddKunt13/><ref name=GarrRodrAgnaCodd15/><ref name=WheeCoddCrowDimi17/>

A study published in 2015 suggested that two families formerly placed in the Haplogynae do not belong there. [Filistatidae](/source/Filistatidae) groups with [Hypochilidae](/source/Hypochilidae) at the base of the Haplogynae; [Leptonetidae](/source/Leptonetidae) is basal to the Entelegynae.<ref name=GarrRodrAgnaCodd15/> The similarity of some morphological features of Leptonetidae to those of entelegynes had already been noted.<ref name=LedfGris10/>

{{barlabel
|size=5
|at1=3 |label1='''Haplogynae''' sensu Coddington (2005) |bar1=purple
|cladogram=
{{clade |style=line-height:90%
|label1=Araneomorphae
|1={{clade
   |1={{cladex
      |1={{cladex
         |1=Hypochilidae
         |2=Filistatidae |barbegin2=purple
         }}
      |2=remaining Haplogynae |bar2=purple
      }}
   |2={{cladex
      |1=Leptonetidae |barend1=purple
      |2=Entelegynae
      }}
   }}
}}
}}

In 2016, a large [molecular phylogenetic](/source/Molecular_phylogenetics) study was published online that included 932 spider species, representing all but one of the then known families. It "refutes important higher-level groups",<ref name=WheeCoddCrowDimi17/> 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 [Synspermiata](/source/Synspermiata) comprises all the ecribellate haplogynes and is consistently recovered, but with low support.<ref name=WheeCoddCrowDimi17/>

{{clade |style=line-height:100%
|label1=[Araneomorphae](/source/Araneomorphae)
|1={{clade
   |1={{clade
      |1={{clade
         |1=[Hypochilidae](/source/Hypochilidae)
         |2=<span style="background-color:yellow>[Filistatidae](/source/Filistatidae)</span>
         }}
       |2=<span style="background-color:yellow>[Synspermiata](/source/Synspermiata) (ecribellate haplogynes)</span>
      }}
   |2={{clade
      |1={{clade
         |1={{clade
            |1=''[Hickmania](/source/Hickmania)'' ([Austrochilidae](/source/Austrochilidae))
            |2=<span style="background-color:yellow>''[Archoleptoneta](/source/Archoleptoneta)'' ([Leptonetidae](/source/Leptonetidae))</span>
            }}
         |2=[Gradungulidae](/source/Gradungulidae)
         }}
      |2={{clade
         |1={{clade
            |1=<span style="background-color:yellow>''[Leptoneta](/source/Leptoneta)'' (Leptonetidae)</span>
            |2=''[Austrochilus](/source/Austrochilus)'' + ''[Thaida](/source/Thaida)'' (Austrochilidae)
            }}
         |2={{clade
            |state1=double
            |1=<span style="background-color:#CFF>[Palpimanoidea](/source/Palpimanoidea)</span> (paraphyletic in molecular analyses)
            |2={{clade
               |1=<span style="background-color:yellow>''Neoleptoneta'' + ''Calileptoneta'' (Leptonetidae)</span>
               |2=<span style="background-color:#CFF>[Entelegynae](/source/Entelegynae) ''sensu'' Wheeler et al. (2017)</span>
               }}
            }}
         }}
      }}
   }}
}}

==Families==
As shown above, [Filistatidae](/source/Filistatidae) and [Leptonetidae](/source/Leptonetidae) are placed outside the traditional haplogynes in the analysis by Wheeler et al. (2017). Traditional haplogyne families they place in Synspermiata are:<ref name=WheeCoddCrowDimi17/>
{{div col|colwidth=15em}}
*[Caponiidae](/source/Caponiidae)
*[Diguetidae](/source/Diguetidae)
*[Drymusidae](/source/Drymusidae)
*[Dysderidae](/source/Dysderidae)
*[Ochyroceratidae](/source/Ochyroceratidae)
*[Oonopidae](/source/Oonopidae)
*[Orsolobidae](/source/Orsolobidae)
*[Pacullidae](/source/Pacullidae)
*[Periegopidae](/source/Periegopidae)
*[Pholcidae](/source/Pholcidae)
*[Plectreuridae](/source/Plectreuridae)
*[Scytodidae](/source/Scytodidae)
*[Segestriidae](/source/Segestriidae)
*[Sicariidae](/source/Sicariidae)
*[Tetrablemmidae](/source/Tetrablemmidae)
{{div col end}}
[Telemidae](/source/Telemidae), traditionally placed in Haplogynae, was not included in the analysis by Wheeler et al.<ref name=WheeCoddCrowDimi17/> However, it is placed in Synspermiata in other studies.<ref name=MichRami14/> The recently discovered haplogyne family [Trogloraptoridae](/source/Trogloraptoridae)<ref name=ZK2012>{{Cite journal |last1=Griswold |first1=C. |last2 = Audisio |first2=T. |last3=Ledford |first3=J. |title=An extraordinary new family of spiders from caves in the Pacific Northwest (Araneae, Trogloraptoridae, new family) |doi=10.3897/zookeys.215.3547 |journal=[ZooKeys](/source/ZooKeys) |pages=77&ndash;102| year=2012 |issue=215 |pmid=22936872| pmc=3428790 |doi-access=free }}</ref> was also placed in Synspermiata.<ref name=WheeCoddCrowDimi17/>

Several extinct families have also been placed into the haplogynes.
*†[Pholcochyroceridae](/source/Pholcochyroceridae)

==References==
<references>
<ref name=AgnaCoddKunt13>{{Cite book |last1=Agnarsson |first1=Ingi |last2=Coddington |first2=Jonathan A. |last3=Kuntner |first3=Matjaž |year=2013 |editor-last=Penney |editor-first=David |contribution=Systematics : Progress in the study of spider diversity and evolution |title=Spider research in the 21st century: trends & perspectives |location=Manchester, UK |publisher=Siri Scientific Press |isbn=978-0-9574530-1-2 |name-list-style=amp }} pp. 82–83.</ref>

<ref name=Codd05>{{Cite book |last=Coddington |first=Jonathan A. |year=2005 |editor-last=Ubick |editor-first=D. |editor2-last=Paquin |editor2-first=P. |editor3-last=Cushing |editor3-first=P.E. |editor4-last=Roth |editor4-first=V. |contribution=Phylogeny and classification of spiders |title=Spiders of North America: an identification manual |pages=18–24 |publisher=American Arachnological Society |access-date=2015-09-24 |contribution-url=https://repository.si.edu/bitstream/handle/10088/4365/CoddingtonSNAPhylogeny05.pdf  |name-list-style=amp }}</ref>

<ref name=GarrRodrAgnaCodd15>{{Cite journal |last1=Garrison |first1=Nicole L. |last2=Rodriguez |first2=Juanita |last3=Agnarsson |first3=Ingi |last4=Coddington |first4=Jonathan A. |last5=Griswold |first5=Charles E. |last6=Hamilton |first6=Christopher A. |last7=Hedin |first7=Marshal |last8=Kocot |first8=Kevin M. |last9=Ledford |first9=Joel M. |last10=Bond |first10=Jason E. |date=2015 |title=Spider phylogenomics: untangling the Spider Tree of Life |journal=PeerJ |volume=3 |page=e1852 |doi=10.7717/peerj.1719 |name-list-style=amp |pmc=4768681 |pmid=26925338 |doi-access=free }}<!--|accessdate=2016-01-09 --></ref>

<ref name=LedfGris10>{{Citation |last1=Ledford |first1=Joel M. |last2=Griswold |first2=Charles E. |date=2010 |title=A study of the subfamily Archoleptonetinae (Araneae, Leptonetidae) with a review of the morphology and relationships for the Leptonetidae |journal=Zootaxa |volume=2391 |pages=1–32 |url=http://mapress.com/zootaxa/2010/f/z02391p032f.pdf |access-date=2016-01-09 |name-list-style=amp |doi=10.11646/zootaxa.2391.1.1 }}</ref>

<ref name=MichRami14>{{Citation |last1=Michalik |first1=Peter |last2=Ramírez |first2=Martín J. |date=2014 |title=Evolutionary morphology of the male reproductive system, spermatozoa and seminal fluid of spiders (Araneae, Arachnida)–Current knowledge and future directions |journal=Arthropod Structure & Development |volume=43 |issue=4 |pages=291–322 |doi=10.1016/j.asd.2014.05.005 |pmid=24907603 |name-list-style=amp |hdl=11336/19081 |hdl-access=free }}</ref>

<ref name=WheeCoddCrowDimi17>{{Citation |last1=Wheeler |first1=Ward C. |last2=Coddington |first2=Jonathan A. |last3=Crowley |first3=Louise M. |last4=Dimitrov |first4=Dimitar |last5=Goloboff |first5=Pablo A. |last6=Griswold |first6=Charles E. |last7=Hormiga |first7=Gustavo |last8=Prendini |first8=Lorenzo |last9=Ramírez |first9=Martín J. |last10=Sierwald |first10=Petra |last11=Almeida-Silva |first11=Lina |last12=Alvarez-Padilla |first12=Fernando |last13=Arnedo |first13=Miquel A. |last14=Benavides Silva |first14=Ligia R. |last15=Benjamin |first15=Suresh P. |last16=Bond |first16=Jason E. |last17=Grismado |first17=Cristian J. |last18=Hasan |first18=Emile |last19=Hedin |first19=Marshal |last20=Izquierdo |first20=Matías A. |last21=Labarque |first21=Facundo M. |last22=Ledford |first22=Joel |last23=Lopardo |first23=Lara |last24=Maddison |first24=Wayne P. |last25=Miller |first25=Jeremy A. |last26=Piacentini |first26=Luis N. |last27=Platnick |first27=Norman I. |last28=Polotow |first28=Daniele |last29=Silva-Dávila |first29=Diana |last30=Scharff |first30=Nikolaj |last31=Szűts |first31=Tamás |last32=Ubick |first32=Darrell |last33=Vink |first33=Cor J. |last34=Wood |first34=Hannah M. |last35=Zhang |first35=Junxia |date=2017 |orig-year=published online 2016 |title=The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling |journal=Cladistics |volume=33 |issue=6 |pages=574–616 |doi=10.1111/cla.12182 |s2cid=35535038 |name-list-style=amp |doi-access=free }}</ref>
</references>

{{Araneae}}
{{Taxonbar|from=Q136328}}

Category:Araneomorphae
Category:Callovian first appearances
Category:Extant Middle Jurassic first appearances
Category:Obsolete spider taxa

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Adapted from the Wikipedia article [Haplogynae](https://en.wikipedia.org/wiki/Haplogynae) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Haplogynae?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.