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.[1]

Arachnologists currently divide spiders into two suborders with about 114 families.

Due to constant research, with new species being discovered every month and others being recognized as synonyms, the number of species in the families is bound to change and can never reflect the present status with total accuracy. Nevertheless, the species numbers given here are useful as a guideline – see the table of families at the end of the article.

History

Spider taxonomy can be traced to the work of Swedish naturalist Carl Alexander Clerck, who in 1757 published the first binomial scientific names of some 67 spiders species in his Svenska Spindlar ("Swedish Spiders"), one year before Linnaeus named over 30 spiders in his Systema Naturae. In the ensuing 250 years, thousands more species have been described by researchers around the world, yet only a dozen taxonomists are responsible for more than a third of all species described. The most prolific authors include Eugène Simon of France, Norman Platnick and Herbert Walter Levi of the United States, Embrik Strand of Norway, and Tamerlan Thorell of Sweden, each having described well over 1,000 species.[2]

Overview of phylogeny

At the very top level, there is broad agreement on the phylogeny and hence classification of spiders, which is summarized in the cladogram below. The three main clades into which spiders are divided are shown in bold; as of 2015, they are usually treated as one suborder, Mesothelae, and two infraorders, Mygalomorphae and Araneomorphae, grouped into the suborder Opisthothelae.[3][4] The Mesothelae, with only 9 species, make up an insignificant proportion of the total of around 45,000 known species. Mygalomorphae species comprise around 6% of the total, the remaining 94% being in the Araneomorphae.[note 1]

Araneae (spiders)

Mesothelae

Opisthothelae

Mygalomorphae

Araneomorphae

Hypochiloidea

Austrochiloidea

Haplogynae

Entelegynae

The Araneomorphae are divided into two main groups: the Haplogynae and the Entelegynae. The Haplogynae make up about 10% of the total number of spider species, the Entelegynae about 83%.[note 1] The phylogenetic relationships of the Haplogynae, Entelegynae and the two smaller groups Hypochiloidea and Austrochiloidea remain uncertain as of 2015. Some analyses place both Hypochiloidea and Austrochiloidea outside Haplogynae;[5] others place the Austrochiloidea between the Haplogynae and the Entelegynae;[6][7] the Hypochiloidea have also been grouped with the Haplogynae.[8] Earlier analyses regarded the Hypochiloidea as the sole representatives of a group called the Paleocribellatae, with all other araneomorphs placed in the Neocribellatae.[9]

The Haplogynae are a group of araneomorph spiders with simpler male and female reproductive anatomy than the Entelegynae. Like the mesotheles and mygalomorphs, females have only a single genital opening (gonopore), used both for copulation and egg-laying;[10] males have less complex palpal bulbs than those of the Entelegynae.[11] Although some studies based on both morphology and DNA suggest that the Haplogynae form a monophyletic group (i.e. they comprise all the descendants of a common ancestor),[12][8] this hypothesis has been described as "weakly supported", with most of the distinguishing features of the group being inherited from ancestors shared with other groups of spiders, rather than being clearly indicative of a separate common origin (i.e. being synapomorphies).[13] One phylogenetic hypothesis based on molecular data shows the Haplogynae as a paraphyletic group leading to the Austrochilidae and Entelegynae.[14]

The Entelegynae have a more complex reproductive anatomy: females have two "copulatory pores" in addition to the single genital pore of other groups of spiders; males have complex palpal bulbs, matching the female genital structures (epigynes).[12] The monophyly of the group is well supported in both morphological and molecular studies. The internal phylogeny of the Entelegynae has been the subject of much research. Two groups within this clade contain the only spiders that make vertical orb webs: the Deinopoidea are cribellate – the adhesive properties of their webs are created by packets of thousands of extremely fine loops of dry silk; the Araneoidea are ecribellate – the adhesive properties of their webs are created by fine droplets of "glue". In spite of these differences, the webs of the two groups are similar in their overall geometry.[15] The evolutionary history of the Entelegynae is thus intimately connected with the evolutionary history of orb webs. One hypothesis is that there is a single clade, Orbiculariae, uniting the orb web makers, in whose ancestors orb webs evolved. A review in 2014 concluded that there is strong evidence that orb webs evolved only once, although only weak support for the monophyly of the Orbiculariae.[16] One possible phylogeny is shown below; the type of web made is shown for each terminal node in order of the frequency of occurrence.[17]

Entelegynae

Eresoidea, RTA clade – no web; substrate-defined web

Orbiculariae

Deinopoideaorb web

Nicodamidaeaerial sheet web

Araneoideaorb web; aerial sheet web; cobweb; no web

If this is correct, the earliest members of the Entelegynae made webs defined by the substrate on which they were placed (e.g. the ground) rather than suspended orb webs. True orb webs evolved once, in the ancestors of the Orbiculariae, but were then modified or lost in some descendants.

An alternative hypothesis, supported by some molecular phylogenetic studies, is that the Orbiculariae are paraphyletic, with the phylogeny of the Entelegynae being as shown below.[18]

Entelegynae

Araneoideaorb web; aerial sheet web; cobweb; no web

RTA clade – no web; substrate-defined web

Deinopoidea, Oecobiidaeorb web; substrate-defined web

On this view, orb webs evolved earlier, being present in the early members of the Entelegynae, and were then lost in more groups,[19] making web evolution more convoluted, with different kinds of web having evolved separately more than once.[16] Future advances in technology, including whole-genome sampling, should lead to "a clearer image of the evolutionary chronicle and the underlying diversity patterns that have resulted in one of the most extraordinary radiations of animals".[16]

Suborder Mesothelae

Mesothelae resemble the Solifugae ("wind scorpions" or "sun scorpions") in having segmented plates on their abdomens that create the appearance of the segmented abdomens of these other arachnids. They are both few in number and also limited in geographical range.

Suborder Opisthothelae

Suborder Opisthothelae contains the spiders that have no plates on their abdomens. It can be somewhat difficult on casual inspection to determine whether the chelicerae of members are of the sort that would classify them as mygalomorphs or araneomorphs. The spiders that are called "tarantulas" in English are so large and hairy that inspection of their chelicerae is hardly necessary to categorize one of them as a mygalomorph. Other, smaller, members of this suborder, however, look little different from the araneomorphs. (See the picture of Sphodros rufipes below.) Many araneomorphs are immediately identifiable as such since they are found on webs designed for the capture of prey or exhibit other habitat choices that eliminate the possibility that they could be mygalomorphs.

Infraorder Mygalomorphae

Spiders in infraorder Mygalomorphae are characterized by the vertical orientation of their chelicerae and the possession of four book lungs.

Infraorder Araneomorphae

Most, if not all, of the spiders one is likely to encounter in everyday life belong to infraorder Araneomorphae. It includes a wide range from the spiders that weave their distinctive orb webs in the garden, the more chaotic-looking webs of the cobweb spiders that frequent window frames and the corners of rooms, the crab spiders that lurk waiting for nectar- and pollen-gathering insects on flowers, to the jumping spiders that patrol the outside walls of a dwelling, and so on. They are characterized by having chelicerae whose tips approach each other as they bite, and (usually) having one pair of book lungs.

Some important spider families are :

These spiders are frequently seen in cellars. When light contact disturbs their web their characteristic response is to set the entire web moving the way a person would jump up and down on a trampoline. It is unclear why they cause their webs to vibrate in this way; moving their webs back and forward may increase the possibility that insects flying close by may be ensnared, or the rapid gyrations caused by the spider in its web may make the spider harder to target by predators.

The family Salticidae, commonly called jumping spiders, have a characteristic cephalothorax shape, as shown in the diagram below. They have eight eyes, two of them very prominent, and excellent vision. Their maximum size is perhaps 13/16 inch (20 mm), but many species are much smaller than that. The largest North American species such as Phidippus regius, P. octopunctatis, etc., are so heavy bodied that they cannot jump far. The smaller species of jumping spider can jump many times their own body length. They hunt by first getting within range of a prey animal such as a fly, securing a silken "climbing rope" to their current perch, and then jumping onto their prey and biting it. Many seem to take unerring aim at the neck of their prey. Should they jump from one twig to another in an attempt to capture prey and miss or get knocked off the second twig by their struggling prey then they are protected from falling by their silken lifeline. At night these spiders usually retreat to a silken "puptent" that they construct for their own protection and, when needed, as a place to deposit their eggs. They are frequently seen in sunlit areas on walls, tree trunks, and other such vertical surfaces.

Classification above families

Spiders were long classified into families that were then grouped into superfamilies, some of which were in turn placed into a number of higher taxa below the level of infraorder. When more rigorous approaches, such as cladistics, were applied to spider classification, it became clear that most of the major groupings used in the 20th century were not supported. Many were based on shared characters inherited from the ancestors of multiple clades (plesiomorphies), rather than being distinctive characters originating in the ancestors of that clade only (apomorphies). According to Jonathan A. Coddington in 2005, "books and overviews published prior to the last two decades have been superseded".[20] Listings of spiders, such as the World Spider Catalog, currently ignore classification above the family level.[20][21]

At the higher level, the phylogeny of spiders is now often discussed using informal clade names, such as the "RTA clade",[22] the "Oval Calmistrum" clade or the "Divided Cribellum" clade.[23] Older names previously used formally are used as clade names, e.g. Entelegynae and Orbiculariae.[24]

Table of families

Key
Genera1≥2≥10≥100
Species1–9≥10≥100≥1000
Spider families[note 2]
FamilyGeneraSpeciesCommon nameExample
Mesothelae
Liphistiidae8116segmented spidersHeptathela kimurai (Kimura spider)
Opisthothelae: Mygalomorphae
Actinopodidae369Missulena bradleyi (eastern mouse spider)
Antrodiaetidae235folding trapdoor spidersAntrodiaetus riversi
Atracidae335Australian funnel-web spidersAtrax robustus (Sydney funnel-web spider)
Atypidae354purse web spiders Sphodros rufipes (red legged purseweb spider)
Barychelidae42295trapdoor baboon spidersSason sundaicum
Ctenizidae353cork-lid trapdoor spidersCteniza sauvagesi
Cyrtaucheniidae11107wafer trapdoor spidersAmblyocarenum nuragicus
Dipluridae25189funnel-web tarantulasMicrohexura montivaga (spruce-fir moss spider)
Euctenizidae776Aptostichus simus
Halonoproctidae684Bothriocyrtum californicum (California trapdoor spider)
Hexathelidae745funnel-web tarantulasPorrhothele antipodiana (black tunnelweb spider)
Idiopidae22344armored trapdoor spidersIdiosoma nigrum (black rugose trapdoor spider)
Macrothelidae129Macrothele calpeiana (Spanish funnel-web spider)
Mecicobothriidae49dwarf tarantulasMegahexura fulva
Microstigmatidae717Envia garciai
Migidae1197tree trapdoor spidersCalathotarsus simoni
Nemesiidae46413Aname atra (black wishbone spider)
Paratropididae411baldlegged spidersParatropis tuxtlensis
Porrhothelidae15Porrhothele antipodiana (black tunnelweb spider)
Theraphosidae144974tarantulasTheraphosa blondi (Goliath birdeater)
Opisthothelae: Araneomorphae
Agelenidae781282araneomorph funnel-web spidersHobo spider
Amaurobiidae49274tangled nest spidersCallobius claustrarius
Ammoxenidae418
Anapidae58223
Anyphaenidae56563anyphaenid sac spidersYellow ghost spider
Araneidae1743128orb-weaver spidersZygiella x-notata
Archaeidae590pelican spidersEriauchenius gracilicollis
Arkyidae237
Austrochilidae310Tasmanian cave spider
Caponiidae18119Diploglena capensis
Cithaeronidae28
Clubionidae15618sac spidersClubiona trivialis
Corinnidae67779dark sac spidersCastianeira sp.
Ctenidae47525tropical wolf spidersBrazilian wandering spiders
Cyatholipidae2358
Cybaeidae19259
Cycloctenidae880
Deinopidae265net-casting spidersRufous net-casting spider
Desidae60297intertidal spidersPhryganoporus candidus
Dictynidae52464Nigma walckenaeri
Diguetidae215coneweb spiders
Drymusidae217false violin spiders
Dysderidae24564woodlouse hunter spidersWoodlouse spider
Eresidae998velvet spidersEresus sandaliatus
Eutichuridae12351Cheiracanthium mildei
Filistatidae19164crevice weaversSouthern house spider
Gallieniellidae1056
Gnaphosidae[note 3]1582532flat-bellied ground spidersDrassodes cupreus
Gradungulidae716large-clawed spidersCarrai cave spider
Hahniidae23346dwarf sheet spiders
Hersiliidae16181tree trunk spidersHersilia savignyi
Homalonychidae13
Huttoniidae11Huttonia palpimanoides
Hypochilidae212lampshade spidersHypochilus thorelli
Lamponidae23192White-tailed spider
Leptonetidae21346Tooth cave spider
Linyphiidae6074566dwarf / money spidersLinyphia triangularis
Liocranidae31272liocranid sac spiders
Lycosidae1242419wolf spidersLycosa tarantula
Malkaridae1146shield spiders
Mecysmaucheniidae725
Megadictynidae22
Mimetidae12152pirate spidersOarces reticulatus
Miturgidae29130long-legged sac spiders
Mysmenidae13137spurred orb-weavers
Nesticidae16278cave cobweb spidersNesticella marapu
Nicodamidae727
Ochyroceratidae20216midget ground weaversTheotima minutissima
Oecobiidae6113disc web spidersOecobius navus
Oonopidae1141801dwarf hunting spidersOonops domesticus
Orsolobidae30188
Oxyopidae9457lynx spidersGreen lynx spider
Pacullidae438
Palpimanidae18150palp-footed spiders
Penestomidae19
Periegopidae13
Philodromidae30539philodromid crab spidersPhilodromus dispar
Pholcidae771666daddy long-legs spidersPholcus phalangioides
Phrurolithidae13205
Physoglenidae1372
Phyxelididae1464
Pimoidae441Pimoa cthulhu
Pisauridae51356nursery web spidersPisaura mirabilis
Plectreuridae231
Psechridae261
Salticidae6356080jumping spidersZebra spider
Scytodidae5248spitting spidersScytodes thoracica
Segestriidae4130tubeweb spidersSegestria florentina
Selenopidae10257wall spidersSelenops radiatus
Senoculidae131
Sicariidae31623recluse spidersBrown recluse
Sparassidae881224huntsman spidersAvondale spider
Stenochilidae213
Stiphidiidae20125Tartarus mullamullangensis
Symphytognathidae873dwarf orb-weaversPatu digua
Synaphridae313
Synotaxidae111
Telemidae1079long-legged cave spiders
Tetrablemmidae27129armored spiders
Tetragnathidae48996long jawed orb-weaversOrchard spider
Theridiidae1242503cobweb spidersRedback spider
Theridiosomatidae19124ray spidersTheridiosoma gemmosum
Thomisidae1702171crab spidersGoldenrod spider
Titanoecidae553Goeldia obscura
Toxopidae1482
Trachelidae18232
Trechaleidae16120
Trochanteriidae19153
Trogloraptoridae11Trogloraptor marchingtoni
Udubidae415
Uloboridae19283hackled orb-weaversUloborus walckenaerius
Viridasiidae29
Xenoctenidae433
Zodariidae851141ant spidersZodarion germanicum
Zoropsidae26180Zoropsis spinimana

Notes

  1. Species counts from Platnick & Raven (2013, Table 1), family classification from Coddington (2005, p. 20).
  2. Unless otherwise shown, currently accepted families and counts based on the World Spider Catalog version 19.0 as of 11 July 2018.[25] In the World Spider Catalog, "species" counts include subspecies. Assignment to sub- and infraorders based on Coddington (2005, p. 20) (when given there).
  3. June 2019 data

References

Bibliography

  • 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.
  • Blackledge, Todd A.; Scharff, Nikolaj; Coddington, Jonathan A.; Szüts, Tamas; Wenzel, John W.; Hayashi, Cheryl Y. & Agnarsson, Ingi (2009). "Reconstructing web evolution and spider diversification in the molecular era". Proceedings of the National Academy of Sciences. 106 (13): 5229–5234. Bibcode:2009PNAS..106.5229B. doi:10.1073/pnas.0901377106. PMC 2656561. PMID 19289848.
  • Bond, Jason E.; Garrison, Nicole L.; Hamilton, Chris A.; Godwin, Rebecca L.; Hedin, Marshal & Agnarsson, Ingi (2014). "Phylogenomics Resolves a Spider Backbone Phylogeny and Rejects a Prevailing Paradigm for Orb Web Evolution". Current Biology. 24 (15): 1765–1771. doi:10.1016/j.cub.2014.06.034. PMID 25042592.
  • 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.
  • Coddington, Jonathan A. & Levi, Herbert W. (1991). "Systematics and evolution of spiders (Araneae)". Annual Review of Ecology and Systematics. 22: 565–592. doi:10.1146/annurev.es.22.110191.003025. JSTOR 2097274.
  • Eberhard, W.G. & Huber, B.A. (2010). "Spider genitalia: precise maneouvers with a numb structure in a complex lock" (PDF). In Leonard, Janet L. & Córdoba-Aguilar, Alex (eds.). The evolution of primary sexual characters in animals. Oxford University Press. ISBN 978-0-19-971703-3. Retrieved 2015-09-20.
  • Griswold, C.E.; Ramirez, M.J.; Coddington, J.A. & Platnick, N.I. (2005). "Atlas of phylogenetic data for entelegyne spiders (Araneae: Araneomorphae: Entelegynae) with comments on their phylogeny" (PDF). Proceedings of the California Academy of Sciences. 56 (Suppl. 2): 1–324. Retrieved 2015-10-11.
  • Hormiga, Gustavo & Griswold, Charles E. (2014). "Systematics, Phylogeny, and Evolution of Orb-Weaving Spiders". Annual Review of Entomology. 59 (1): 487–512. doi:10.1146/annurev-ento-011613-162046. PMID 24160416.
  • 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. PMID 24907603.
  • Platnick, Norman I. & Raven, Robert J. (2013). "Spider Systematics: Past and Future". Zootaxa. 3683 (5): 595–600. doi:10.11646/zootaxa.3683.5.8.
  • Ramírez, Martín J. (2014). The morphology and phylogeny of dionychan spiders (Araneae, Araneomorphae). Bulletin of the American Museum of Natural History. 390. hdl:2246/6537.
  • World Spider Catalog (2018). "World Spider Catalog version 19.0". Natural History Museum Bern. Retrieved 2018-07-11.
This article is issued from Wikipedia. The text is licensed under Creative Commons - Attribution - Sharealike. Additional terms may apply for the media files.