Spider taxonomy

Spider taxonomy is the taxonomy of the 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.

Araneus-angulatus-figure1757
Paintings of Araneus angulatus from Svenska Spindlar of 1757, the first major work on spider taxonomy

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

Sphodros rufipes non-crossing chel
Digitally enhanced image of a Sphodros rufipes that shows the nearly perfectly vertical orientation of the chelicerae, a prime characteristic of the Mygalomorphae.

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

Megaphobema robustum 1
Megaphobema robustum, one of the many kinds of spiders called "tarantulas"

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

Infraorder Araneomorphae

Cheiracanthium punctorium frei 1 17 Forst Jungfernhdeide Jg 46 070920
Photograph showing orientation of the chelicerae of the 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.

Salticidae cephalothorax di
"Squared-off" cephalothorax of the jumping spiders.
Salticidae eyes diag
Eye pattern of the jumping spiders.

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
Genera 1 ≥2 ≥10 ≥100
Species 1–9 ≥10 ≥100 ≥1000
Spider families[note 2]
Family Genera Species Common name Example
Mesothelae
Liphistiidae 8 116 segmented spiders Heptathela kimurai (Kimura spider)
Opisthothelae: Mygalomorphae
Actinopodidae 3 69 Missulena bradleyi (eastern mouse spider)
Antrodiaetidae 2 35 folding trapdoor spiders Antrodiaetus riversi
Atracidae 3 35 Australian funnel-web spiders Atrax robustus (Sydney funnel-web spider)
Atypidae 3 54 purse web spiders Sphodros rufipes (red legged purseweb spider)
Barychelidae 42 295 trapdoor baboon spiders Sason sundaicum
Ctenizidae 3 53 cork-lid trapdoor spiders Cteniza sauvagesi
Cyrtaucheniidae 11 107 wafer trapdoor spiders Amblyocarenum nuragicus
Dipluridae 25 189 funnel-web tarantulas Microhexura montivaga (spruce-fir moss spider)
Euctenizidae 7 76 Aptostichus simus
Halonoproctidae 6 84 Bothriocyrtum californicum (California trapdoor spider)
Hexathelidae 7 45 funnel-web tarantulas Porrhothele antipodiana (black tunnelweb spider)
Idiopidae 22 344 armored trapdoor spiders Idiosoma nigrum (black rugose trapdoor spider)
Macrothelidae 1 29 Macrothele calpeiana (Spanish funnel-web spider)
Mecicobothriidae 4 9 dwarf tarantulas Megahexura fulva
Microstigmatidae 7 17 Envia garciai
Migidae 11 97 tree trapdoor spiders Calathotarsus simoni
Nemesiidae 46 413 Aname atra (black wishbone spider)
Paratropididae 4 11 baldlegged spiders Paratropis tuxtlensis
Porrhothelidae 1 5 Porrhothele antipodiana (black tunnelweb spider)
Theraphosidae 144 974 tarantulas Theraphosa blondi (Goliath birdeater)
Opisthothelae: Araneomorphae
Agelenidae 78 1282 araneomorph funnel-web spiders Hobo spider
Amaurobiidae 49 274 tangled nest spiders Callobius claustrarius
Ammoxenidae 4 18
Anapidae 58 223
Anyphaenidae 56 563 anyphaenid sac spiders Yellow ghost spider
Araneidae 174 3128 orb-weaver spiders Zygiella x-notata
Archaeidae 5 90 pelican spiders Eriauchenius gracilicollis
Arkyidae 2 37
Austrochilidae 3 10 Tasmanian cave spider
Caponiidae 18 119 Diploglena capensis
Cithaeronidae 2 8
Clubionidae 15 618 sac spiders Clubiona trivialis
Corinnidae 67 779 dark sac spiders Castianeira sp.
Ctenidae 47 525 tropical wolf spiders Brazilian wandering spiders
Cyatholipidae 23 58
Cybaeidae 19 259
Cycloctenidae 8 80
Deinopidae 2 65 net-casting spiders Rufous net-casting spider
Desidae 60 297 intertidal spiders Phryganoporus candidus
Dictynidae 52 464 Nigma walckenaeri
Diguetidae 2 15 coneweb spiders
Drymusidae 2 17 false violin spiders
Dysderidae 24 564 woodlouse hunter spiders Woodlouse spider
Eresidae 9 98 velvet spiders Eresus sandaliatus
Eutichuridae 12 351 Cheiracanthium mildei
Filistatidae 19 164 crevice weavers Southern house spider
Gallieniellidae 10 56
Gnaphosidae 128 2220 flat-bellied ground spiders Drassodes cupreus
Gradungulidae 7 16 large-clawed spiders Carrai cave spider
Hahniidae 23 346 dwarf sheet spiders
Hersiliidae 16 181 tree trunk spiders Hersilia savignyi
Homalonychidae 1 3
Huttoniidae 1 1 Huttonia palpimanoides
Hypochilidae 2 12 lampshade spiders Hypochilus thorelli
Lamponidae 23 192 White-tailed spider
Leptonetidae 21 346 Tooth cave spider
Linyphiidae 607 4566 dwarf / money spiders Linyphia triangularis
Liocranidae 31 272 liocranid sac spiders
Lycosidae 124 2419 wolf spiders Lycosa tarantula
Malkaridae 11 46 shield spiders
Mecysmaucheniidae 7 25
Megadictynidae 2 2
Mimetidae 12 152 pirate spiders Oarces reticulatus
Miturgidae 29 130 long-legged sac spiders
Mysmenidae 13 137 spurred orb-weavers
Nesticidae 16 278 cave cobweb spiders Nesticella marapu
Nicodamidae 7 27
Ochyroceratidae 20 216 midget ground weavers Theotima minutissima
Oecobiidae 6 113 disc web spiders Oecobius navus
Oonopidae 114 1801 dwarf hunting spiders Oonops domesticus
Orsolobidae 30 188
Oxyopidae 9 457 lynx spiders Green lynx spider
Pacullidae 4 38
Palpimanidae 18 150 palp-footed spiders
Penestomidae 1 9
Periegopidae 1 3
Philodromidae 30 539 philodromid crab spiders Philodromus dispar
Pholcidae 77 1666 daddy long-legs spiders Pholcus phalangioides
Phrurolithidae 13 205
Physoglenidae 13 72
Phyxelididae 14 64
Pimoidae 4 41 Pimoa cthulhu
Pisauridae 51 356 nursery web spiders Pisaura mirabilis
Plectreuridae 2 31
Prodidomidae 31 309 long-spinneret ground spiders Lygromma anops
Psechridae 2 61
Salticidae 635 6080 jumping spiders Zebra spider
Scytodidae 5 248 spitting spiders Scytodes thoracica
Segestriidae 4 130 tubeweb spiders Segestria florentina
Selenopidae 10 257 wall spiders Selenops radiatus
Senoculidae 1 31
Sicariidae 3 1623 recluse spiders Brown recluse
Sparassidae 88 1224 huntsman spiders Avondale spider
Stenochilidae 2 13
Stiphidiidae 20 125 Tartarus mullamullangensis
Symphytognathidae 8 73 dwarf orb-weavers Patu digua
Synaphridae 3 13
Synotaxidae 1 11
Telemidae 10 79 long-legged cave spiders
Tetrablemmidae 27 129 armored spiders
Tetragnathidae 48 996 long jawed orb-weavers Orchard spider
Theridiidae 124 2503 cobweb spiders Redback spider
Theridiosomatidae 19 124 ray spiders Theridiosoma gemmosum
Thomisidae 170 2171 crab spiders Goldenrod spider
Titanoecidae 5 53 Goeldia obscura
Toxopidae 14 82
Trachelidae 18 232
Trechaleidae 16 120
Trochanteriidae 19 153
Trogloraptoridae 1 1 Trogloraptor marchingtoni
Udubidae 4 15
Uloboridae 19 283 hackled orb-weavers Uloborus walckenaerius
Viridasiidae 2 9
Xenoctenidae 4 33
Zodariidae 85 1141 ant spiders Zodarion germanicum
Zoropsidae 26 180 Zoropsis spinimana

Notes

  1. ^ a b 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).

References

  1. ^ Platnick & Raven (2013), p. 600.
  2. ^ Platnick & Raven (2013), p. 597.
  3. ^ Bond et al. (2014).
  4. ^ Coddington (2005).
  5. ^ Coddington (2005), p. 20.
  6. ^ Griswold et al. (2005).
  7. ^ Blackledge et al. (2009), p. 5232.
  8. ^ a b Bond et al. (2014), p. 1766.
  9. ^ Coddington & Levi (1991), p. 577.
  10. ^ Eberhard & Huber (2010), pp. 256–257.
  11. ^ Eberhard & Huber (2010), p. 250.
  12. ^ a b Coddington (2005), p. 22.
  13. ^ Michalik & Ramírez (2014), p. 312.
  14. ^ Agnarsson, Coddington & Kuntner (2013), p. 40.
  15. ^ Hormiga & Griswold (2014), p. 488.
  16. ^ a b c Hormiga & Griswold (2014), p. 505.
  17. ^ Blackledge et al. (2009), Fig. 3.
  18. ^ Bond et al. (2014), Fig 3. Web types defined as Blackledge et al. (2009, Fig. 3)
  19. ^ Bond et al. (2014), p. 1768.
  20. ^ a b Coddington (2005), p. 24.
  21. ^ World Spider Catalog (2018).
  22. ^ Hormiga & Griswold (2014), p. 491.
  23. ^ Ramírez (2014), p. 4.
  24. ^ Hormiga & Griswold (2014), pp. 490–491.
  25. ^ World Spider Catalog (2018), Currently valid spider genera and species.

Bibliography

  • Agnarsson, Ingi; Coddington, Jonathan A. & Kuntner, Matjaž (2013). "Systematics : Progress in the study of spider diversity and evolution". In Penney, David. 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. 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. 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. Retrieved 2015-09-24.
  • 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.
  • World Spider Catalog (2018). "World Spider Catalog version 19.0". Natural History Museum Bern. Retrieved 2018-07-11.

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