Spider taxonomy

Paintings of Araneus angulatus from Svenska Spindlar of 1757, the first major work on 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.

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

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, 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

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 of Salticidae commonly called jumping spiders have a characteristic cephalothorax shapes, 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. They are perhaps the only family of spiders who will take cognizance of a human in their general area and then turn their bodies and elevate their cephalothoraxes to keep the human under observation. If approached closely, e.g., with the lens of a camera, some of them may choose to jump onto the nearby object to explore it. This behavior may be alarming but it never seems to be aggressive since these spiders are unwilling to attack prey that are very much larger than they are.

"Squared-off" cephalothorax of the jumping spiders.
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]
FamilyGeneraSpeciesCommon nameExample
Mesothelae
Liphistiidae896segmented spiders Kimura spider
Opisthothelae: Mygalomorphae
Actinopodidae347 Missulena (mouse spiders)
Antrodiaetidae235folding trapdoor spiders Antrodiaetus riversi
Atypidae351purse web spiders Red legged purseweb spider
Barychelidae42295trapdoor baboon spiders Sason sundaicum
Ctenizidae9130cork-lid trapdoor spiders Cteniza sauvagesi
Cyrtaucheniidae11107wafer trapdoor spiders Amblyocarenum nuragicus
Dipluridae24188funnel-web tarantulas Spruce-fir moss spider
Euctenizidae775 Aptostichus simus
Hexathelidae12113venomous funnel-web tarantulas Sydney funnel-web spider
Idiopidae22323 Black rugose trapdoor spider
Mecicobothriidae49dwarf tarantulas
Microstigmatidae716 Envia garciai
Migidae1091tree trapdoor spiders
Nemesiidae45390 Black wishbone spider
Paratropididae410baldlegged spiders
Theraphosidae131981tarantulas Goliath birdeater
Opisthothelae: Araneomorphae
Agelenidae701168araneomorph funnel-web spiders Hobo spider
Amaurobiidae51287tangled nest spiders Callobius claustrarius
Ammoxenidae418
Amphinectidae32159 Metaltella simoni
Anapidae38154
Anyphaenidae56542anyphaenid sac spiders Yellow ghost spider
Araneidae1693097orb-weaver spiders Zygiella x-notata
Archaeidae471pelican spiders Eriauchenius gracilicollis
Austrochilidae39 Tasmanian cave spider
Caponiidae1598 Diploglena capensis
Chummidae12
Cithaeronidae28
Clubionidae15587sac spiders Clubiona trivialis
Corinnidae67729dark sac spiders Castianeira sp.
Ctenidae41503tropical wolf spiders Brazilian wandering spiders
Cyatholipidae2358
Cybaeidae10179 Water spider
Cycloctenidae536
Deinopidae261net-casting spiders Rufous net-casting spider
Desidae38185intertidal spiders Phryganoporus candidus
Dictynidae52578 Nigma walckenaeri
Diguetidae215coneweb spiders
Drymusidae116false violin spiders
Dysderidae24534woodlouse hunter spiders Woodlouse spider
Eresidae997velvet spiders Eresus sandaliatus
Eutichuridae12343 Cheiracanthium mildei
Filistatidae18121crevice weavers Southern house spider
Gallieniellidae1055
Gnaphosidae1222178flat-bellied ground spiders Drassodes cupreus
Gradungulidae716large-clawed spiders Carrai cave spider
Hahniidae28249dwarf sheet spiders
Hersiliidae15179tree trunk spiders Hersilia savignyi
Holarchaeidae12
Homalonychidae13
Huttoniidae11 Huttonia palpimanoides
Hypochilidae212lampshade spiders Hypochilus thorelli
Lamponidae23192 White-tailed spider
Leptonetidae22272 Tooth cave spider
Linyphiidae6014533dwarf / money spiders Linyphia triangularis
Liocranidae32272liocranid sac spiders
Lycosidae1232403wolf spiders Lycosa tarantula
Malkaridae411shield spiders
Mecysmaucheniidae725
Micropholcommatidae1966
Mimetidae13158pirate spiders Oarces reticulatus
Miturgidae33159long-legged sac spiders
Mysmenidae13135spurred orb-weavers Mysmenopsis sp.
Nephilidae561golden-orb web spiders Nephila pilipes
Nesticidae13228cave cobweb spiders Nesticella marapu
Nicodamidae929
Ochyroceratidae15187midget ground weavers Theotima minutissima
Oecobiidae6110disc web spiders Oecobius navus
Oonopidae1121613dwarf hunting spiders Oonops domesticus
Orsolobidae30188
Oxyopidae9453lynx spiders Green lynx spider
Palpimanidae16139palp-footed spiders
Pararchaeidae735
Penestomidae19
Periegopidae13
Philodromidae30539philodromid crab spiders Philodromus dispar
Pholcidae791461daddy long-legs spiders Pholcus phalangioides
Phrurolithidae14197
Phyxelididae1464
Pimoidae438 Pimoa cthulhu
Pisauridae47335nursery web spiders Pisaura mirabilis
Plectreuridae231
Prodidomidae31309long-spinneret ground spiders Lygromma anops
Psechridae257
Salticidae5885841jumping spiders Zebra spider
Scytodidae5232spitting spiders Scytodes thoracica
Segestriidae4120tubeweb spiders Segestria florentina
Selenopidae10256wall spiders Selenops radiatus
Senoculidae131
Sicariidae2139recluse spiders Brown recluse
Sinopimoidae11
Sparassidae851180huntsman spiders Avondale spider
Stenochilidae213
Stiphidiidae22135 Tartarus mullamullangensis
Symphytognathidae871dwarf orb-weavers Patu digua
Synaphridae313
Synotaxidae1482
Telemidae962long-legged cave spiders
Tetrablemmidae31160armored spiders
Tetragnathidae47973long jawed orb-weavers Orchard spider
Theridiidae1222459cobweb spiders Redback spider
Theridiosomatidae18109ray spiders Theridiosoma gemmosum
Thomisidae1752150crab spiders Goldenrod spider
Titanoecidae553 Goeldia obscura
Trachelidae16208
Trechaleidae16120
Trochanteriidae19152
Trogloraptoridae11 Trogloraptor marchingtoni
Udubidae412
Uloboridae18272hackled orb-weaversUloborus walckenaerius
Viridasiidae29
Zodariidae811095 Zodarion germanicum
Zoropsidae25177 Zoropsis spinimana

Notes

  1. 1 2 Species counts from Platnick & Raven (2013, Table 1), family classification from Coddington (2005, p. 20).
  2. Currently accepted families and counts based on the World Spider Catalog version 16.5 as of 6 November 2015.[25] Assignment to sub- and infraorders based on Coddington (2005, p. 20) (when given).

References

Bibliography

External links

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