Entelegynae

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Entelegynae
Temporal range: Jurassic–present
Araneus diadematus qtl1.jpg
Underside of Araneus diadematus showing epigynum
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Araneae
Infraorder: Araneomorphae
Clade: Entelegynae
Simon, 1893
Families

Unnamed clade

Unnamed clade

Incertae sedis:

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]

Contents

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]

Spider fertilization systems
Spider fertilization systems - haplogyne.svg
Non-entelegyne
Spider fertilization systems - entelegyne.svg
Entelegyne
Schematic diagrams showing sperm entering and being stored in the spermathecae; eggs leaving the ovaries and being fertilized; and finally a fertilized egg leaving the female's body

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]

No representative of the family Synaphridae was included in the 2017 analysis, [1] but previous analyses placed it in the Entelegynae. [5]

Related Research Articles

Araneomorphae Infraorder of arachnids

The Araneomorphae are an infraorder of spiders. They are distinguishable by chelicerae (fangs) that point diagonally forward and cross in a pinching action, in contrast to the Mygalomorphae, where they point straight down. Araneomorphs comprise the vast majority of living spiders.

Crevice weaver Family of spiders

Crevice weaver spiders (Filistatidae) comprise cribellate spiders with features that have been regarded as "primitive" for araneomorph spiders. They are weavers of funnel or tube webs. The family contains 18 genera and more than 120 described species worldwide.

Spider taxonomy

Spider taxonomy is that part of taxonomy that is concerned with the science of naming, defining and classifying all spiders, members of the Araneae order of the arthropod class Arachnida with about 46,000 described species. However, there are likely many species that have escaped the human eye to this day, and many specimens stored in collections waiting to be described and classified. It is estimated that only one third to one half of the total number of existing species have been described.

Cribellum

Cribellum literally means "little sieve", and in biology the term generally applies to anatomical structures in the form of tiny perforated plates.

Nephilinae Spider family

Nephilinae is a spider subfamily of the family Araneidae with seven genera. The various genera in Nephilinae were formerly grouped in the family Nephilidae, and before that in the Tetragnathidae and in the Araneidae. All nephiline genera partially renew their webs. Spiders in the subfamily Nephilinae are commonly referred to as golden orb-weavers.

Palpimanoidea Superfamily of spiders

The Palpimanoidea or palpimanoids, also known as assassin spiders, are a group of araneomorph spiders, originally treated as a superfamily. As with many such groups, its circumscription has varied. As of September 2018, the following five families were included:

Leptonetoidea Superfamily of arachnids

The Leptonetoidea are a superfamily of haplogyne araneomorph spiders with three families. Phylogenetic studies have provided weak support for the relationship among the families. The placement of one of the families within the Haplogynae has been questioned.

Dysderoidea Superfamily of spiders

The Dysderoidea are a clade or superfamily of araneomorph spiders. The monophyly of the group, initially consisting of the four families Dysderidae, Oonopidae, Orsolobidae and Segestriidae, has consistently been recovered in phylogenetic studies. In 2014, a new family, Trogloraptoridae, was created for a recently discovered species Trogloraptor marchingtoni. It was suggested that Trogloraptoridae may be the most basal member of the Dysderoidea clade. However, a later study found that Trogloraptoridae was placed outside the Dysderoidea and concluded that it was not part of this clade.

Eresoidea

The Eresoidea or eresoids are a group of araneomorph spiders that have been treated as a superfamily. As usually circumscribed, the group contains three families: Eresidae, Hersiliidae and Oecobiidae. Studies and reviews based on morphology suggested the monophyly of the group; more recent gene-based studies have found the Eresidae and Oecobiidae to fall into different clades, placing doubt on the acceptability of the taxon. Some researchers have grouped Hersiliidae and Oecobiidae into the separate superfamily Oecobioidea, a conclusion supported in a 2017 study, which does not support Eresoidea.

Agelenoidea

The Agelenoidea or agelenoids are a superfamily or informal group of entelegyne araneomorph spiders. Phylogenetic studies since 2000 have not consistently recovered such a group, with more recent studies rejecting it.

The Dictynoidea or dictynoids are a group of araneomorph spiders that have been treated as a superfamily. The composition of the group has varied. Phylogenetic studies in the 21st century have failed to confirm the monophyly of the dictynoids as originally defined.

Austrochiloidea Superfamily of spiders

The Austrochiloidea or austrochiloids are a group of araneomorph spiders, treated as a superfamily. The taxon contains two families of eight-eyed spiders:

Haplogynae

The Haplogynae or haplogynes are one of the two main groups into which araneomorph spiders have traditionally been divided, the other being the Entelegynae. Morphological phylogenetic studies suggested that the Haplogynae formed a clade; more recent molecular phylogenetic studies refute this, although many of the ecribellate haplogynes do appear to form a clade, Synspermiata.

Spider anatomy

The anatomy of spiders includes many characteristics shared with other arachnids. These characteristics include bodies divided into two tagmata, eight jointed legs, no wings or antennae, the presence of chelicerae and pedipalps, simple eyes, and an exoskeleton, which is periodically shed.

This glossary describes the terms used in formal descriptions of spiders; where applicable these terms are used in describing other arachnids.

Palpal bulb

The two palpal bulbs – also known as palpal organs and genital bulbs – are the copulatory organs of a male spider. They are borne on the last segment of the pedipalps, giving the spider an appearance often described as like wearing boxing gloves. The palpal bulb does not actually produce sperm, being used only to transfer it to the female. Palpal bulbs are only fully developed in adult male spiders and are not completely visible until after the final moult. In the majority of species of spider, the bulbs have complex shapes and are important in identification.

Orbiculariae

Orbiculariae is a potential clade of araneomorph spiders, uniting two groups that make orb webs. Phylogenetic analyses based on morphological characters have generally recovered this clade; analyses based on DNA have regularly concluded that the group is not monophyletic. The issue relates to the origin of orb webs: whether they evolved early in the evolutionary history of entelegyne spiders, with many groups subsequently losing the ability to make orb webs, or whether they evolved later, with fewer groups having lost this ability. As of September 2018, the weight of the evidence strongly favours the non-monophyly of "Orbiculariae" and hence the early evolution of orb webs, followed by multiple changes and losses.

Synspermiata Clade of spiders

Synspermiata is a clade of araneomorph spiders, comprising most of the former "haplogynes". They are united by having simpler genitalia than other araneomorph spiders, lacking a cribellum, and sharing an evolutionary history of synspermia – a particular way in which spermatozoa are grouped together when transferred to the female.

<i>Leucauge mariana</i> Species of spider

Leucauge mariana is a long-jawed orb weaver spider, native to Central America and South America. Its web building and sexual behavior have been studied extensively. Males perform several kinds of courtship behavior to induce females to copulate and to use their sperm.

Larinia jeskovi is a species of the family of orb weaver spiders and a part of the genus Larinia. It is distributed throughout the Americas, Africa, Australia, Europe, and Asia and commonly found in wet climes such as marshes, bogs, and rainforests. Larinia jeskovi have yellow bodies with stripes and range from 5.13-8.70 millimeters in body length. They build their webs on plants with a small height above small bodies of waters or wetlands. After sunset and before sunrise are the typical times they hunt and build their web. Males usually occupy a female's web instead of making their own. The mating behavior is noteworthy as male spiders often mutilate external female genitalia to reduce sperm competition while female spiders resort to sexual cannibalism to counter such mechanisms. The males also follow an elaborate courtship ritual to attract the female. The bite of Larinia jeskovi is not known to be of harm to humans.

References

  1. 1 2 3 4 5 6 7 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
  2. 1 2 3 Foelix, Rainer F. (2011), Biology of Spiders (3rd p/b ed.), Oxford University Press, ISBN   978-0-19-973482-5
  3. 1 2 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
  4. 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
  5. 1 2 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.