Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Chelicerata
Classis: Arachnida
Ordo: Araneae
Subordo: Opisthothelae
Infraordo: Araneomorphae
Taxon: Neocribellatae
Series: Entelegynae
Superfamiliae: Dionycha (sectio) – Agelenoidea – Amaurobioidea – Araneoidea – Archaeoidea – Dictynoidea – Eresoidea – Lycosoidea – Mimetoidea – Nicodamoidea – Oecobioidea – Palpimanoidea – Tengelloidea – Titanoecoidea – Uloboroidea – Zodaroidea – †Archaeometoidea – †Pyritaraneoidea
References
Griswold, C.E., Coddington, J.A., Platnick, N.I. & Forster, R.R. 1999. Towards a phylogeny of entelegyne spiders (Araneae, Araneomorphae, Entelegynae). Journal of Arachnology 27: 53–63. PDF.
Griswold, C.E., Ramírez, M.J., Coddington, J.A. & Platnick, N.I. 2005. Atlas of phylogenetic data for entelegyne spiders (Araneae: Araneomorphae: Entelegynae) with comments on their phylogeny. Proceedings of the California Academy of Sciences 56(Suppl. II): 1–324. PDF.
Miller, J.A., Carmichael, A., Ramírez, M.J., Spagna, J.C., Haddad, C.R., Řezáč, M., Johannesen, J., Král, J., Wang, X.-P. & Griswold, C.E. 2010. Phylogeny of entelegyne spiders: affinities of the family Penestomidae (new rank), generic phylogeny of Eresidae, and asymmetric rates of change in spinning organ evolution (Araneae, Araneoidea, Entelegynae). Molecular phylogenetics and evolution 55: 786–804. DOI: 10.1016/j.ympev.2010.02.021
The Entelegynae or entelegynes are a subgroup of araneomorph spiders, the largest of the two main groups into which the araneomorphs were traditionally divided. Females have a genital plate (epigynum) and a "flow through" fertilization system; males have complex palpal bulbs. Molecular phylogenetic studies have supported the monophyly of Entelegynae (whereas the other traditional subgroup, the Haplogynae, has been shown not to be monophyletic).[1]
The clade contains both cribellate and ecribellate spiders.
Characterization
The Entelegynae are characterized primarily by the nature of the female genital system. The ancestral (plesiomorphic) system is found in non-entelegyne spiders, where there is a single external genital opening in the female's abdomen. One or more males inject sperm from their palpal bulbs via this opening; the sperm is usually stored in special spermathecae (absent in some spiders, e.g. Pholcus). When eggs are released from the ovaries, sperm is also released, and the fertilized eggs pass out of the female's body by the same opening. Sperm that enters first is likely to be the last to fertilize eggs.[2][1][3]
In entelegyne spiders, there are three external openings in the female's body. Sperm is injected via one or other of the two separate copulatory openings and enters the spermathecae. Egg release and fertilization occurs in the same way as in non-entelegyne spiders. In this "flow through" system, sperm that enters first can be the first to fertilize eggs.[2][1][3]
The copulatory openings are usually surrounded by a hardened (sclerotized) area called the epigynum. In some entelegyne families, such as araneids, the epigynum includes a projection that covers or partially covers the copulatory openings. This plays a role in aligning the male's palpal bulb during mating.[2] Male entelegyne spiders generally have more complex palpal bulbs than other groups of spiders and the bulbs are expanded and moved by haemolytic pressure alone, as there are no muscles attached to them.[4]
Phylogeny
In 2016, a large molecular phylogenetic study was published online that included 932 spider species, representing all but one of the then known families. It supported the monophyly of Entelegynae, but presented a somewhat complex picture of its position within Araneomorphae. Araneomorph spiders were divided into two clades: one comprising the families Filistatidae and Hypochilidae plus the clade Synspermiata, the other comprising three non-monophyletic families (Austrochilidae, Gradungulidae and Leptonetidae) plus Palpimanoidea, the four being basal to Entelegynae.[1]
Araneomorphae
Filistatidae + Hypochilidae
Synspermiata (ecribellate haplogynes)
Austrochilidae + Gradungulidae + Leptonetidae (part) (paraphyletic)
Palpimanoidea (paraphyletic in molecular analyses)
Leptonetidae (part)
Entelegynae
Most members of the former Haplogynae are placed in the Synspermiata. Filistatidae is placed outside the Synspermiata; Leptonetidae, which was found not to be monophyletic, is placed basal to the Entelegynae.[1]
Families
The cladogram in Wheeler et al. (2017) includes the following families in the Entelegynae. The main difference from previous circumscriptions, such as Coddington in 2005,[5] is that the Palpimanoidea are excluded. A few groups within otherwise entelegyne families have reverted to a haplogyne state: the genera Comaroma, Tangaroa and Waitkera, and some members of the family Tetragnathidae.[1]
Agelenidae
Amaurobiidae
Ammoxenidae
Anapidae
Anyphaenidae
Araneidae
Arkyidae
Cheiracanthiidae (syn. Eutichuridae)
Cithaeronidae
Clubionidae
Corinnidae
Ctenidae
Cyatholipidae
Cybaeidae
Cycloctenidae
Deinopidae
Desidae
Dictynidae
Eresidae
Gallieniellidae
Gnaphosidae
Hahniidae
Hersiliidae
Homalonychidae
Lamponidae
Linyphiidae
Liocranidae
Lycosidae
Malkaridae
Megadictynidae
Mimetidae
Miturgidae
Mysmenidae
Nesticidae
Nicodamidae
Oecobiidae
Oxyopidae
Penestomidae
Philodromidae
Phrurolithidae
Physoglenidae
Phyxelididae
Pimoidae
Pisauridae
Psechridae
Salticidae
Selenopidae
Senoculidae
Sparassidae
Stiphidiidae
Symphytognathidae
Synotaxidae
Tetragnathidae
Theridiidae
Theridiosomatidae
Thomisidae
Titanoecidae
Toxopidae
Trachelidae
Trechaleidae
Trochanteriidae
Udubidae
Uloboridae
Viridasiidae
Xenoctenidae
Zodariidae
Zoropsidae
No representative of the family Synaphridae was included in the 2017 analysis,[1] but previous analyses placed it in the Entelegynae.[5]
References
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
Foelix, Rainer F. (2011), Biology of Spiders (3rd p/b ed.), Oxford University Press, ISBN 978-0-19-973482-5
Uhl, Gabriele; Nessler, Stefan H. & Schneider, Jutta M. (2010), "Securing paternity in spiders? A review on occurrence and effects of mating plugs and male genital mutilation", Genetica, 138 (1): 75–104, doi:10.1007/s10709-009-9388-5
Huber, Bernhard A. (2004), "Evolutionary transformation from muscular to hydraulic movements in spider (Arachnida, Araneae) genitalia: A study based on histological serial sections", Journal of Morphology, 261 (3): 364–376, doi:10.1002/jmor.10255
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. p. 20.
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