Spider taxonomy

Last updated
Paintings of Araneus angulatus from Svenska Spindlar of 1757, the first major work on spider taxonomy Araneus-angulatus-figure1757.jpg
Paintings of Araneus angulatus from Svenska Spindlar of 1757, the first major work on 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 more than 48,500 described species. [1] 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. [2]

Contents

Arachnologists currently divide spiders into two suborders with about 129 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 only reflects the present state of knowledge. 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. [3]

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. [4] [5] The Mesothelae, with about 140 species in 8 genera as of October 2020, make up a very small proportion of the total of around 49,000 known species. Mygalomorphae species comprise around 7% of the total, the remaining 93% 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; [6] others place the Austrochiloidea between the Haplogynae and the Entelegynae; [7] [8] the Hypochiloidea have also been grouped with the Haplogynae. [9] Earlier analyses regarded the Hypochiloidea as the sole representatives of a group called the Paleocribellatae, with all other araneomorphs placed in the Neocribellatae. [10]

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; [11] males have less complex palpal bulbs than those of the Entelegynae. [12] 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), [13] [9] 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). [14] One phylogenetic hypothesis based on molecular data shows the Haplogynae as a paraphyletic group leading to the Austrochilidae and Entelegynae. [15]

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). [13] 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. [16] 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. [17] One possible phylogeny is shown below; the type of web made is shown for each terminal node in order of the frequency of occurrence. [18]

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. [19]

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, [20] making web evolution more convoluted, with different kinds of web having evolved separately more than once. [17] Future advances in technology, including comparative genomics studies, [1] and 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". [17]

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.

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

Suborder Opisthothelae

Suborder Opisthothelae contains the spiders that have no plates on their abdomens. Opisthothelae is divided into two infraorders, Mygalomorphae and Araneomorphae, which can be distinguished by the orientation of their fangs. It can be somewhat difficult on casual inspection to determine whether the fang orientation would classify a spider as a mygalomorph or araneomorph. The spiders that are called "tarantulas" in English are so large and hairy that inspection of their fangs 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" Megaphobema robustum 1.jpg
Megaphobema robustum , one of the many kinds of spiders called "tarantulas"

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

Infraorder Araneomorphae

Photograph showing orientation of the fangs of the Araneomorphae. Cheiracanthium punctorium frei 1 17 Forst Jungfernhdeide Jg 46 070920.jpg
Photograph showing orientation of the fangs 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 of spider families, including the orb-weaver spiders that weave their distinctive webs in gardens, the cobweb spiders that frequent window frames and the corners of rooms, the crab spiders that lurk on flowers waiting for nectar- and pollen-gathering insects, the jumping spiders that patrol the outside walls of buildings, and so on. They are characterized by having fangs whose tips approach each other as they bite, and (usually) having one pair of book lungs.

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 characteristics inherited from the ancestors of multiple clades (plesiomorphies), rather than being distinct characteristics 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". [21] Listings of spiders, such as the World Spider Catalog, currently ignore classification above the family level. [21] [22]

At the higher level, the phylogeny of spiders is now often discussed using informal clade names, such as the "RTA clade", [23] the "Oval Calmistrum" clade or the "Divided Cribellum" clade. [24] Older names previously used formally are used as clade names, e.g. Entelegynae and Orbiculariae. [25]

Table of families

Key
Genera1≥2≥10≥100
Species1–9≥10≥100≥1000
Spider families [note 2]
FamilyGeneraSpeciesCommon nameExample
Mesothelae
Heptathelidae 7113 Kimura spider
Liphistiidae 170 Liphistius batuensis
Opisthothelae: Mygalomorphae
Actinopodidae 3118 Missulena (mouse spiders)
Anamidae 10111 Aname diversicolor (black wishbone spider)
Antrodiaetidae 437folding trapdoor spiders Atypoides riversi
Atracidae 336Australian funnel-web spiders Illawarra wisharti
Atypidae 356purseweb spiders Sphodros rufipes (red-legged purseweb spider)
Barychelidae 40282trapdoor baboon spiders Sason sundaicum
Bemmeridae 447 Spiroctenus personatus
Ctenizidae 25cork-lid trapdoor spiders Cteniza sauvagesi
Cyrtaucheniidae 693wafer trapdoor spiders Amblyocarenum nuragicus
Dipluridae 795funnel-web tarantulas Spruce-fir moss spider (Microhexura montivaga)
Entypesidae 642 Entypesa andohahela
Euagridae 1487 Euagrus formosanus
Euctenizidae 877 Aptostichus simus
Halonoproctidae 6130 Ummidia algarve
Hexathelidae 745venomous funnel-web tarantulas Hexathele hochstetteri
Hexurellidae 14
Idiopidae 23437armored trapdoor spiders Idiosoma nigrum (black rugose trapdoor spider)
Ischnothelidae 526
Macrothelidae 247
Mecicobothriidae 12dwarf tarantulas
Megahexuridae 11
Microhexuridae 12
Microstigmatidae 1138 Envia garciai
Migidae 11103tree trapdoor spiders Moggridgea rainbowi
Nemesiidae 10148funnel-web trapdoor spiders Aname atra (black wishbone spider)
Paratropididae 416baldlegged spiders Paratropis tuxtlensis
Porrhothelidae 15
Pycnothelidae 15137
Rhytidicolidae 214 Fufius lucasae
Stasimopidae 147
Theraphosidae 1561039tarantulas Goliath birdeater (Theraphosa blondi)
Opisthothelae: Araneomorphae
Agelenidae 901366araneomorph funnel-web spiders Hobo spider (Eratigena agrestis)
Amaurobiidae 50283tangled nest spiders Callobius claustrarius
Anapidae 58232
Anyphaenidae 58614anyphaenid sac spiders Hibana velox (yellow ghost spider)
Araneidae 1843097orb-weaver spiders Zygiella x-notata
Archaeidae 690pelican spiders Madagascarchaea gracilicollis
Archoleptonetidae 28 Archoleptoneta gertschi
Arkyidae 238
Austrochilidae 29 Hickmania troglodytes
Caponiidae 20139 Diploglena capensis
Cheiracanthiidae 14363
Cithaeronidae 29
Clubionidae 19662sac spiders Clubiona trivialis
Corinnidae 73824dark sac spiders Castianeira sp.
Ctenidae 48532wandering spiders Phoneutria fera
Cyatholipidae 2358
Cybaeidae 21303 Diving bell spider (Argyroneta aquatica)
Cycloctenidae 880
Deinopidae 367net-casting spiders Deinopis subrufa (rufous net-casting spider)
Desidae 60296intertidal spiders Phryganoporus candidus
Dictynidae 53475 Nigma walckenaeri
Diguetidae 215coneweb spiders
Drymusidae 217false violin spiders
Dysderidae 25591woodlouse hunter spiders Woodlouse spider (Dysdera crocata)
Eresidae 9102velvet spiders Eresus sandaliatus
Filistatidae 19189crevice weavers Southern house spider (Kukulcania hibernalis)
Gallieniellidae 541
Gnaphosidae 1452430ground spiders Drassodes cupreus
Gradungulidae 817large-clawed spiders Progradungula carraiensis (Carrai cave spider)
Hahniidae 24353dwarf sheet spiders
Hersiliidae 16187tree trunk spiders Hersilia savignyi
Homalonychidae 12
Huttoniidae 11 Huttonia palpimanoides
Hypochilidae 233lampshade spiders Hypochilus thorelli
Lamponidae 23192 White-tailed spider (Lampona spp.)
Leptonetidae 22370 Tooth Cave spider (Tayshaneta myopica)
Linyphiidae 6244724dwarf / money spiders Linyphia triangularis
Liocranidae 35311liocranid sac spiders
Lycosidae 1272457wolf spiders Lycosa tarantula
Malkaridae 1357shield spiders
Mecysmaucheniidae 725
Megadictynidae 22
Mimetidae 8159pirate spiders Oarces reticulatus
Miturgidae 29141long-legged sac spiders
Myrmecicultoridae 11
Mysmenidae 14158spurred orb-weavers
Nesticidae 18285scaffold web spiders Nesticella marapu
Nicodamidae 727
Ochyroceratidae 10177midget ground weavers Theotima minutissima
Oecobiidae 6120disc web spiders Oecobius navus
Oonopidae 1151888dwarf hunting spiders Oonops domesticus
Orsolobidae 30188
Oxyopidae 9443lynx spiders Peucetia viridans (green lynx spider)
Pacullidae 438
Palpimanidae 21165palp-footed spiders
Penestomidae 19
Periegopidae 13
Philodromidae 30535philodromid crab spiders Philodromus dispar
Pholcidae 971893daddy long-legs spiders Pholcus phalangioides
Phrurolithidae 20313
Physoglenidae 1372
Phyxelididae 1468
Pimoidae 285 Pimoa cthulhu
Pisauridae 51353nursery web spiders Pisaura mirabilis
Plectreuridae 231
Prodidomidae 23192
Psechridae 261
Psilodercidae 11224
Salticidae 6656433jumping spiders Zebra spider (Salticus scenicus)
Scytodidae 4241spitting spiders Scytodes thoracica
Segestriidae 5152tubeweb spiders Segestria florentina
Selenopidae 9262wall spiders Selenops radiatus
Senoculidae 131
Sicariidae 3172recluse spiders Brown recluse (Loxosceles reclusa)
Sparassidae 951337huntsman spiders Delena cancerides
Stenochilidae 213
Stiphidiidae 20125sheetweb spiders Tartarus mullamullangensis
Symphytognathidae 1098dwarf orb-weavers Patu digua
Synaphridae 313
Synotaxidae 111
Telemidae 16104long-legged cave spiders
Tetrablemmidae 27151armored spiders
Tetragnathidae 46982long jawed orb-weavers Leucauge venusta (orchard spider)
Theridiidae 1252538cobweb spiders Redback spider (Latrodectus hasselti)
Theridiosomatidae 20135ray spiders Theridiosoma gemmosum
Thomisidae 1712167crab spiders Misumena vatia (goldenrod crab spider)
Titanoecidae 556 Goeldia obscura
Toxopidae 1482
Trachelidae 20263
Trachycosmidae 20148
Trechaleidae 20148
Trochanteriidae 651
Trogloraptoridae 11 Trogloraptor marchingtoni
Udubidae 415
Uloboridae 19291hackled orb-weavers Uloborus walckenaerius
Viridasiidae 314
Xenoctenidae 433
Zodariidae 891251ant spiders Zodarion germanicum
Zoropsidae 27182 Zoropsis spinimana

Notes

  1. 1 2 Species counts from World Spider Catalog (2020 , Currently valid spider genera and species ), family classification from Coddington (2005 , p. 20).
  2. Unless otherwise shown, currently accepted families and counts based on the World Spider Catalog version 22.5 as of 5 November 2021. [26] In the World Spider Catalog, "species" counts include subspecies. Assignment to sub- and infraorders based on Coddington (2005, p. 20) (when given there).

Related Research Articles

<span class="mw-page-title-main">Mygalomorphae</span> Infraorder of arachnids (spiders)

The Mygalomorphae, or mygalomorphs, are an infraorder of spiders, and comprise one of three major groups of living spiders with over 3,000 species, found on all continents except Antarctica. Many members are known as trapdoor spiders due to their creation of trapdoors over their burrows. Other prominent groups include Australian funnel web spiders and tarantulas, with the latter accounting for around one third of all mygalomorphs.

<span class="mw-page-title-main">Araneomorphae</span> 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.

<span class="mw-page-title-main">Mesothelae</span> Suborder of spiders

The Mesothelae are a suborder of spiders that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral characters, such as a segmented abdomen with spinnerets in the middle and two pairs of book lungs. Members of Liphistiidae are medium to large spiders with eight eyes grouped on a tubercle. They are found only in China, Japan, and southeast Asia. The oldest known Mesothelae spiders are known from the Carboniferous, over 300 million years ago.

<span class="mw-page-title-main">Lampshade spider</span> Family of spiders

Lampshade spiders, family Hypochilidae, are among the most primitive of araneomorph spiders. There are two genera and twelve species currently recognized. Like mygalomorphs, most hypochilids have two pairs of book lungs, but like araneomorphs they have intersecting fangs, with the exception of some species which have chelicerae in an angle that is neither orthognathous or labidognathous. These long-legged spiders build typical "lampshade" style webs under overhangs and in caves. In the United States the fauna is primarily associated with the Appalachian, Rocky and California Mountains. Ten of the known species are found in these ranges, all in the genus Hypochilus. The genus Ectatosticta is found in China.

<span class="mw-page-title-main">Cribellum</span>

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

<span class="mw-page-title-main">Austrochilidae</span> Family of spiders

Austrochilidae is a small spider family with nine species in two genera. Austrochilus and Thaida are endemic to the Andean forest of central and southern Chile and adjacent Argentina.

<span class="mw-page-title-main">Phyxelididae</span> Family of spiders

Phyxelididae is a family of araneomorph spiders first described by Pekka T. Lehtinen in 1967 as a subfamily of Amaurobiidae, and later elevated to family status as a sister group of Titanoecidae.

<span class="mw-page-title-main">Micropholcommatinae</span> Subfamily of spiders

The Micropholcommatinae are a subfamily of araneomorph spiders in the family Anapidae. They were previously treated as the family Micropholcommatidae. Micropholcommatins are extremely small, with body lengths typically between 0.5 and 2 mm. They are usually found among leaf litter or moss.

<span class="mw-page-title-main">Dionycha</span> 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 2021 analysis resulted in about 20 families, including Salticidae, Gnaphosidae, and Clubionidae.

<span class="mw-page-title-main">Dysderoidea</span> 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.

<span class="mw-page-title-main">Eresoidea</span>

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.

<span class="mw-page-title-main">Deinopoidea</span> Superfamily of spiders

The Deinopoidea or deinopoids are group of cribellate araneomorph spiders that may be treated as a superfamily. As usually circumscribed, the group contains two families: Deinopidae and Uloboridae.

<span class="mw-page-title-main">Agelenoidea</span>

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.

<span class="mw-page-title-main">Titanoecoidea</span> Superfamily of spiders

The Titanoecoidea or titanoecoids are a proposed taxon of araneomorph spiders at the superfamily rank. The taxon contains two families of spiders, Phyxelididae and Titanoecidae. Although some phylogenetic studies have shown these two families to form a clade, other studies have not, placing Titanoecidae outside the RTA clade while Phyxelididae is placed inside it. A 2011 classification of spider families leaves both Phyxelididae and Titanoecidae outside the RTA clade as "unplaced non-Orbiculariae families". The status of the group remains unclear as of December 2015.

<span class="mw-page-title-main">Austrochiloidea</span> 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:

<span class="mw-page-title-main">Haplogynae</span> Infraorder of spiders

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.

<span class="mw-page-title-main">Entelegynae</span> 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.

<span class="mw-page-title-main">Opisthothelae</span> Suborder of spiders

Opisthothelae is a suborder of spiders within the order Araneae, containing Mygalomorphae and Araneomorphae, but excluding Mesothelae. The Opisthothelae are sometimes presented as an unranked clade and sometimes as a suborder of Araneae. In the latter case, Mygalomorphae and Araneomorphae are treated as infraorders.

<span class="mw-page-title-main">Orbiculariae</span>

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.

References

  1. 1 2 Dimitrov & Hormiga (2021).
  2. Platnick & Raven (2013), p. 600.
  3. Platnick & Raven (2013), p. 597.
  4. Bond et al. (2014).
  5. Coddington (2005).
  6. Coddington (2005), p. 20.
  7. Griswold et al. (2005).
  8. Blackledge et al. (2009), p. 5232.
  9. 1 2 Bond et al. (2014), p. 1766.
  10. Coddington & Levi (1991), p. 577.
  11. Eberhard & Huber (2010), pp. 256–257.
  12. Eberhard & Huber (2010), p. 250.
  13. 1 2 Coddington (2005), p. 22.
  14. Michalik & Ramírez (2014), p. 312.
  15. Agnarsson, Coddington & Kuntner (2013), p. 40.
  16. Hormiga & Griswold (2014), p. 488.
  17. 1 2 3 Hormiga & Griswold (2014), p. 505.
  18. Blackledge et al. (2009), Fig. 3.
  19. Bond et al. (2014), Fig 3. Web types defined as Blackledge et al. (2009, Fig. 3)
  20. Bond et al. (2014), p. 1768.
  21. 1 2 Coddington (2005), p. 24.
  22. World Spider Catalog (2020).
  23. Hormiga & Griswold (2014), p. 491.
  24. Ramírez (2014), p. 4.
  25. Hormiga & Griswold (2014), pp. 490–491.
  26. World Spider Catalog (2021), Currently valid spider genera and species.

Bibliography