Lycosoidea

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Lycosoidea
Lycosidae female carrying young.jpg
Wolf spider carrying young
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Araneae
Infraorder: Araneomorphae
Clade: Entelegynae
Superfamily: Lycosoidea
Families

See text.

Lycosoidea is a clade or superfamily of araneomorph spiders. The traditional circumscription was based on a feature of the eyes. The tapetum is a reflective layer at the back of the eye, thought to increase sensitivity in low light levels. Lycosoids were then defined by having a "grate-shaped" tapetum. Research from the late 1990s onwards suggests that this feature has evolved more than once, possibly as many as five times, [1] so that the original Lycosoidea is paraphyletic. [2] Studies published in 2014 and 2015 suggest that a smaller group of families does form a clade.

Contents

The reflective attributes of the tapeta vary significantly betwixt lycosoid spiders. [3]

Phylogeny

The phylogeny of the Lycosoidea and related groups has been studied using both morphological and molecular data. Comparing these studies is complicated by the redrawing of family boundaries. Thus potential lycosoid genera placed in the family Miturgidae in 1993, such as Uliodon , [4] have since been placed in the Zoropsidae. The family Ctenidae, placed in the Lycosoidea in many analyses, has been reduced by moving some genera into the new family Viridasiidae. [5]

The "classical" circumscription of the Lycosoidea was based on the possession of a grate-shaped tapetum in some or all of the indirect eyes. In 1993, Griswold concluded that the Lycosoidea so defined was monophyletic, although some of the lycosoid families (as then circumscribed) were not. [4] (See the table below for the families included in this and later studies.) Stiphidiidae was later excluded from the Lycosoidea. [6] Later studies have recovered a version of Lycosoidea as a monophyletic group, although with varying compositions and placements within clades. [1] [7] Ramírez, as part of a study primarily directed at dionychans, suggested that a reduced Lycosoidea, consisting of only five families, was monophyletic. Ctenidae and Zoropsidae were excluded from the lycosoid clade. [8] (Trechaleidae was not included in the study.) An alternative hypothesis explored by Ramírez also placed Thomisidae in the Lycosoidea. [9] Polotow et al. (2015) concluded that a seven-family Lycosoidea was monophyletic. Of the "classical" lycosoid families, they put back Ctenidae left out by Ramírez (although saying that the family is not monophyletic), but additionally excluded Senoculidae. They supported Ramírez' alternative of placing Thomisidae in Lycosoidea. [10]

Some circumscriptions of Lycosoidea
FamilyGriswold (1993) [4] Ramírez (2014) [8] Polotow et al. (2015) [10]
Ctenidae yesyes
Lycosidae yesyesyes
Oxyopidae yesyesyes
Pisauridae yesyesyes
Psechridae yesyesyes
Senoculidae yesyes
Stiphidiidae yes
Thomisidae ?yes
Trechaleidae yes*yes
Zoropsidae yes**
* Not included in the study.
** Some genera included were then placed in Miturgidae.

Lycosoid families are located within the large RTA clade of araneomorph spiders. [2] [11] Within this clade, Polotow et al. place them within a "grate-shaped tapetum clade", with the family Zoropsidae as their sister: [12]

RTA clade

basal groups

Dionycha

Oval calamistrum clade

Udubidae

Grateshaped tapetum clade

Zoropsidae

Lycosoidea

Related Research Articles

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.

Zorocratidae is a formerly accepted family of spiders. Most of the genera formerly placed in this family have been transferred to the family Udubidae. The type genus, Zorocrates, is now placed in the Zoropsidae.

Tengellidae Family of spiders

Tengellidae is a former family of spiders that has been merged into the family Zoropsidae. Genera formerly placed in Tengellidae now in Zoropsidae include:

Zoropsidae Family of spiders

Zoropsidae, also known as false wolf spiders for their physical similarity to wolf spiders, is a family of cribellate araneomorph spiders first described by Philipp Bertkau in 1882. They can be distinguished from wolf spiders by their two rows of eyes that are more equal in size than those of Lycosidae.

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.

Dionycha Clade of spiders

The Dionycha are a clade of spiders (Araneomorphae:Entelegynae), characterized by the possession of two tarsal claws with tufts of hairs (setae) beside them, which produce strong adhesion, enabling some species to climb glass. The circumscription of the group has varied widely; a 2014 analysis resulted in about 20 families, including Salticidae, Thomisidae, and Clubionidae. Spiders in this group have better senses than others, some even show courtship dances and songs.

Gnaphosoidea Superfamily of spiders

The Gnaphosoidea or gnaphosoids are a superfamily of araneomorph spiders with seven families. A 2014 study did not find the group to be monophyletic.

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.

Araneoidea Superfamily of spiders

Araneoidea is a taxon of araneomorph spiders, termed "araneoids", treated as a superfamily. As with many such groups, its circumscription has varied; in particular some families that had at one time moved to the Palpimanoidea have more recently been restored to Araneoidea. A 2014 treatment includes 18 families, with the araneoids making up about 26% of the total number of known spider species; a 2016 treatment includes essentially the same taxa, but now divided into 17 families.

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.

Entelegynae Clade of spiders

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.

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.

RTA clade Clade of spiders

The RTA clade is a clade of araneomorph spiders, united by the possession of a retrolateral tibial apophysis – a backward-facing projection on the tibia of the male pedipalp. The clade contains over 21,000 species, almost half the current total of about 46,000 known species of spider. Most of the members of the clade are wanderers and do not build webs. Despite making up approximately half of all modern spider diversity, there are no unambiguous records of the group from the Mesozoic and molecular clock evidence suggests that the group began to diversify during the Late Cretaceous.

Udubidae Family of spiders

Udubidae is a family of araneomorph spiders, most of whose members were formerly placed in the family Zorocratidae, which is no longer accepted.

Zorodictyna is a genus of spiders in the family Udubidae native to Madagascar. It has been described as an intermediate genus between Zoropsidae and Dictynidae, though it is now placed in Udubidae. This genus was originally placed in the family Zoropsidae, but it has been reassigned several times since. In 1967, Lehtinen moved it to Miturgidae. In 1999, it was moved back to Zoropsidae, and in 2015, it was moved to Udubidae.

Campostichomma is a genus of spiders in the family Udubidae native to Sri Lanka. Many of its species were moved to either Griswoldia or Devendra. This genus was originally placed in the family Agelenidae. It was moved to Miturgidae in 1967, to Zoropsidae in 1999, then to Udubidae in 2015.

<i>Viridasius</i> Genus of spiders

Viridasius is a monotypic genus of East African araneomorph spiders in the family Viridasiidae, containing the single species, Viridasius fasciatus. It was first described by Eugène Simon in 1889, and has only been found in Madagascar.

Liocranoides is a genus of American false wolf spiders that was first described by Eugen von Keyserling in 1881. They live in habitats with cold surfaces, such as caves. It was transferred from the sac spiders to the Tengellidae in 1967, which was later merged with Zoropsidae.

References

  1. 1 2 Ramírez (2014), p. 282.
  2. 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.
  3. Benson, Kari; Suter, Robert B. (2013). "Reflections on the tapetum lucidum and eyeshine in lycosoid spiders". The Journal of Arachnology. 41 (1): 43–52. doi:10.1636/B12-35.1. JSTOR   23610234. S2CID   36757386. These results demonstrate that the reflective qualities of the tapeta, and perhaps the absorptive qualities of other tissues and media that the light must traverse, vary widely among lycosoid spiders.
  4. 1 2 3 Griswold, C. E. (1993). "Investigations into the phylogeny of Lycosoid spiders and their kin (Arachnida: Araneae: Lycosoidea)". Smithsonian Contributions to Zoology. 539 (539): 1–39. doi:10.5479/si.00810282.539 . Retrieved 2015-10-17.
  5. Polotow, Carmichael & Griswold (2015), p. 151-152.
  6. Davila, Diana Silva (2003). Higher-level relationships of the spider family Ctenidae (Araneae, Ctenoidea). Bulletin of the American Museum of Natural History. hdl:2246/441.
  7. Polotow, Carmichael & Griswold (2015), p. 125.
  8. 1 2 Ramírez (2014), p. 290.
  9. Ramírez (2014), pp. 319–320.
  10. 1 2 Polotow, Carmichael & Griswold (2015), p. 134.
  11. Nentwig, Wolfgang, ed. (2013). "Appendix : Spider Phylogeny" (PDF). Spider Ecophysiology. Springer. ISBN   978-3-642-33988-2 . Retrieved 2015-11-03.
  12. Polotow, Carmichael & Griswold (2015), p. 132.

Bibliography