Araneidae

Basilica spider Mecynogea lemniscata, copyright Volcano.

Belongs within: Araneoidea.
Contains: Nephilinae, Micratheninae, Gasteracanthinae, Cyclosini, Mangora, Cyrtophora, Argiope, Cyrtarachninae, Neoscona, Araneus.

The Araneidae are the largest family of orb-weaving spiders, members of which construct an orb web with a sticky spiral, or a modification thereof (Jocqué & Dippenaar-Schoeman 2007). Members of the family are morphologically and behaviourally diverse and their classification has long been confused. As recognised below, araneids are united by the presence of modified setae (the sustentaculum) on the tip of the fourth tarsi and the presence of a radix in the embolic division of the male palp (Scharff et al. 2020) and may be divided between three major clades. The Zygiellinae are characterised by spinnerets with aggregate spigots apart from flagelliform spigots, and construct webs using doubled radial threads and incorporating a silken tube retreat. A second clade unites the Nephilinae with the southern Australian Paraplectanoides crassipes.

The remaining araneids are united by molecular data into a clade informally dubbed the ‘ARA clade’ by Scharff et al. (2020). Historical subfamilial divisions within this clade remain of uncertain validity. In particular, the subfamily Araneinae, as historically defined by the presence of tubercles on the femur of the male pedipalp and an epigynal scape in the female, appears to be highly polyphyletic. The Caerostrinae are morphologically aberrant araneids that lack a radix on the male pedipalp (Scharff et al. 2020). Members of the Cyrtophorinae have the piriform spinning field on the anterior lateral spinnerets reduced to a narrow ribbon, and construct their web as a horizontal, tightly woven mesh with an extremely tight, permanent non-sticky spiral (Framenau & Scharff 2009). The Argiopeae have a procurved posterior eye row with a reduced canoe-shaped tapetum in the lateral eyes, and the male palp possesses a thread-like spur on the median apophysis (Scharff & Coddington 1997).

Araneidae: with web and scent
Published 30 June 2008
The St. Andrew’s Cross Argiope keyserlingi. Photo by Louise Docker.

The orb-weavers are undoubtedly the best-known of all spiders. Ask anyone to imagine a spider and they will probably picture an orb-weaver (they may also have transcribed the words “some pig” in the web). This is something of an unfair characterisation—of the more than 100 recognised families of spiders, less than ten are orb-weavers. Still, it is one of the orb-weaving genera that holds the name of “spider”, Araneus, which, as the only generic name used in Clerck’s (1757) Aranei Svecici, the only taxonomic work recognised by the ICZN that predates the 1758 tenth edition of Linnaeus’ Systema Naturae, is officially the oldest generic name in zoological nomenclature*. That’s right—spiders came before humans. Nyeh nyeh nyeh.

*Admittedly Clerck did use the name Araneus for all spiders, not just species included in the modern Araneus.

The Araneidae are the largest family of orb-weaving spiders, with a little less than 3000 described species. They are actually a lot more numerous than you might realise—many species build their webs only at night, taking them down in the morning before hiding during the day and rebuilding the web every evening. The family is decidedly diverse in appearance—from the gaudy colours and spines of the Christmas spiders to the idiosyncratic figures of the tailed spiders to one group whose common name describes their appearance perfectly—the bird-dropping spiders.

The classic orb-web is made by first floating a line of sticky silk horizontally across a space between two anchoring points (such as a pair of branches), then running a second looser non-sticky strand along the initial strand. The spider then drops herself* from the centre-point of the second strand, trailing a third strand behind her, so that the second and third strands form a Y-shape. The vertex of the Y will be the centre of the web. The spider next constructs an outer frame, as shown above in a diagram by Ed Nieuwenhuys (the page linked to has diagrams of each of the stages in orb-web construction), then runs a series of spokes from the centre of the web to the outside. She then runs a broad spiral of non-sticky thread from the centre of the web until she reaches the outer edge. After that, it travels back to the centre laying a much tighter spiral of sticky thread, removing the non-sticky scaffold as she goes. As the sticky thread is stretched, the sticky coating breaks into a series of globules of coiled thread, which is how the web is able to be so elastic and stand up to the thrashings of captured prey. The spider herself is able to move about without being trapped by means of secretions produced by glands near the mouth with which she coats her legs. Forster & Forster (1999) refer to an experiment where the tips of a spider’s legs were dipped in solvent before the spider was returned to its web. The spider initially showed great difficulty in moving due to the removal of its protective coating, though it was able to renew the covering and regain mobility. After the web has been completed, the spider will take up residence at the central hub, legs resting on the radiating spokes in order to feel for any vibrations. Araneid eyesight is almost non-existent, and orb-weavers are incapable of hunting without a web. They are perhaps the closest thing to a terrestrial filter-feeder, filtering the air for small animals.

*All spiders are referred to as female unless proven otherwise, like ships and birds of prey. It’s another one of those things that make the English language so damn confusing.

An unidentified member of the genus Gasteracantha. These spiders come in a dazzling array of colours and ornamentations, and unlike many other araneids are often visible during the day, earning them such names as “jewel spider” or “Christmas spider”. Colour patterns can vary significantly even between members of the same species. Photo from here.

Many araneids may vary the basic orb-web design further. Ladder-web spiders, for instance, have a long narrow web instead of the usual circle. The function of these is not really understood, though it has been suggested as a specialisation for catching moths – moths have a covering of loose scales on their wings which would normally allow them to shake off a web and escape, but it is suggested that the elongate shape of a ladder-web means that as the moth shakes off its scales, it falls onto a lower part of the web until eventually it is no longer able to escape. Many orb-weavers construct a stabilimentum, a zig-zag ladder-shaped structure that extends upwards from the central hub. Again, despite being such a distinctive structure, the function of the stabilimentum remains largely unknown, though subject to intense debate – suggested roles include strengthening the web (the original source of the name), disguising the position of the spider from predators or making the spider look bigger, attracting prey or even making the web more visible for larger animals and so reducing the risk of them walking or flying through it. One large and striking araneid found here in Australia, the St. Andrew’s cross Argiope keyserlingi, shown at the top of this section, gets its name because it builds four stabilimenta radiating from the central hub, while the spider itself sits with the front two and rear two pairs of legs held alongside each other, so the spider itself forms the eponymous cross shape.

Perhaps the most remarkable thing about araneids, however, is that despite the total dependence of most species on their webs for survival, some species no longer make them. The aforementioned bird-dropping spider Celaenia is so-called because its lumpy brown-and-white-splotched abdomen really does look like a lump of bird poo, offering excellent camouflage from discerning predators. Instead of constructing a full web, Celaenia simply hang from a leaf or a thread with their legs outstreched. There they catch and feed on moths (excellent pictures of hunting Celaenia can be seen at Esperance Blog). It used to be a mystery how this seemingly limited and haphazard means of capture could possibly feed the spider (after all, how many moths could reasonably be expected to pass by one point over the course of a night) until it was observed that a surprising proportion of the moths being caught (that is, all of them) were males, and that, far from passing by the spider accidentally, male moths will actually approach the spider and remain close by it until caught. It seems that the spider actually emits pheromones that mimic those of a female moth, luring their prey in with the false promise of sexual gratification (like a Trojan virus attached to a spam e-mail). The bolas spiders of the tribe Mastophoreae have refined this process further—as well as producing attractive pheromones, they also dangle a single sticky thread below themselves. When a moth approaches close enough, the spider spins the sticky thread around in the air until it sticks to the moth and they are able to draw it in. How bird-dropping and bolas spiders make their living until they become large enough to handle moths seems a little confused—Brunet (1996) claims that Celaenia construct standard orb-webs until they reach maturity, while bolas spiders produce different pheromones for attracting different-sized moths when at different ages. Forster & Forster (1999) and Yeargan (1994), in contrast, both claim that Celaenia spiderlings produce pheromones to attract psychodid midges. Interestingly, while bird-dropping and bolas spiders are both members of the subfamily Araneinae, it is debatable whether they are each other’s closest relatives within the subfamily (Yeargan 1994), so it is possible that their amazing pheromone-capture techniques could have arisen separately of each other!

Systematics of Araneidae

Characters (from Jocqué & Dippenaar-Schoeman 2007): Small to large; eight eyes present in two rows with lateral eyes widely separated from median eyes; labium rebordered; legs with three tarsal claws, usually with numerous spines and sustentaculum on tarsus IV; abdomen globose, overhanging the carapace; ecribellate; male palp with mesal cymbium, median apophysis and radix in embolic division; entelegyne.

<==Araneidae [Anepsieae, Araneinae, Araneini, Argiopinae, Epeiridae, Epeirides, Mangoreae, Mangorini, Poltyeae, Poltyinae]
    |--Zygiellinae [Phonognatheae, Phonognathidae, Zygielleae]SC20
    |    |  i. s.: LeviellusSC20
    |    |--+--Deliochus Simon 1894SC20, K02
    |    |  `--Phonognatha Simon 1894SC20, K02 [incl. SingotypaHAA07]
    |    |       |--P. graeffei (Rainbow 1896)B96
    |    |       |--P. melaniaB96
    |    |       `--P. sylvicolaB96
    |    `--Zygiella Pickard-Cambridge 1902SC20, PVD10 [=Zygia Koch 1834 non Fabricius 1775G76]
    |         |--*Z. atrica (Koch 1845) (see below for synonymy)G76
    |         |--Z. kochi (Thorell 1870) [=Zilla kochi]G76
    |         |--Z. montana (Koch 1839) [=Zilla montana]G76
    |         |--Z. stroemi (Thorell 1870)G76 [=Zilla stroemiG76, Aranea (Zilla) stroemiS06, Stroemiellus stroemiAM11]
    |         |--Z. thorelli (Ausserer 1871) [=Zilla thorelli]G76
    |         `--Z. x-notata (Clerck 1758)PVD10 [=Zilla x-notataG76]
    `--+--+--NephilinaeSC20
       |  `--Paraplectanoides Keyserling 1886SC20
       |       |--*P. crassipes Keyserling 1886KK90, SC20
       |       `--P. cerula Simon 1908M90
       `--+--CaerostrinaeSC20
          |    |--Testudinaria Taczanowski 1879 non Salisbury ex Burchell 1824 (ICBN) [Testudinareae]SC20
          |    |--Gnolus Simon 1879 [Gnoleae, Oarciinae]SC20
          |    |    `--G. cordiformisSC20
          |    `--Caerostris Thorell 1868SC20, GA11 [Caerostreae]
          |         |--C. darwini Kuntner & Agnarsson 2010GA11
          |         |--C. ‘paradoxa’ (Doleschall 1859)K92, D59 [=Epeira paradoxa non Lucas 1846K92]
          |         |--C. sexcuspidataSC97
          |         |--C. sumatrana Strand 1915GA11
          |         `--C. vinsoni Thorell 1868SC97
          `--+--‘Araneus’ necopinusSC20
             `--+--+--*Encyosaccus sexmaculatus Simon 1895SC97
                |  `--MicratheninaeSC20
                `--+--Spilasma Simon 1897SC20
                   |    `--S. duodecimguttataSC20
                   `--+--ChorizopeaeSC97
                      |    |--ArtonisSC97
                      |    `--Chorizopes Pickard-Cambridge 1871 [Chorizopinae]SC20
                      |         |--*C. frontalisK02
                      |         `--C. nipponicusSC20
                      `--+--TelaproceraSC20
                         |    |--T. joanaeSC20
                         |    `--T. maudaeSC20
                         `--+--+--Hingstepeira Levi 1995SC20, K02
                            |  |    `--H. folisecens (Hingston 1932)K02
                            |  `--+--+--+--Acanthepeira Marx 1883SC20, AM11
                            |     |  |  |    |--*A. stellata (Walckenaer 1805)SC97
                            |     |  |  |    `--A. venusta (Banks 1896)SC97
                            |     |  |  `--+--Lariniophora Framenau 2011F11
                            |     |  |     |    `--*L. ragnhildae (Strand 1917) (see below for synonymy)F11
                            |     |  |     `--Eriophora Simon 1864SC20, F11
                            |     |  |          |--*E. ravilla (Koch 1844)F11
                            |     |  |          |--E. biapicata (Koch 1871)F11
                            |     |  |          |--E. bistriataD95
                            |     |  |          |--E. edax (Blackwall 1863)SC97
                            |     |  |          |--E. fuliginea (Koch 1838)B11
                            |     |  |          |--E. heroine (Koch 1871)PVD10 [incl. Epeira brounii Urquhart 1884FF99]
                            |     |  |          `--E. pustulosa (Walckenaer 1842)F11 (see below for synonymy)
                            |     |  `--+--Parawixia Pickard-Cambridge 1904SC20, AM11
                            |     |     |    |--*P. destricta Pickard-Cambridge 1889SC20
                            |     |     |    |--P. kochi (Taczanowski 1873)AM11
                            |     |     |    `--P. velutina (Taczanowski 1878)AM11
                            |     |     `--+--Wagneriana Pickard-Cambridge 1904SC20, AM11
                            |     |        |    |--W. tauricornisSC20
                            |     |        |    `--W. tayosSC97
                            |     |        `--Alpaida Pickard-Cambridge 1889SC20, AM11
                            |     |             |--A. bicornuta (Taczanowski 1878)SC97
                            |     |             |--A. championi (P.-Cambridge 1889)SC97
                            |     |             |--A. delicataSC20
                            |     |             |--A. grayi [=Aranea grayi]F09
                            |     |             |--A. leucogramma (White 1841)AM11
                            |     |             |--A. truncata (Keyserling 1865)AM11
                            |     |             `--A. veniliae (Keyserling 1865)AM11
                            |     `--+--+--Hypognatha Guérin 1839 [Eurycorminae, Hypognatheae]SC20
                            |        |  |    `--*H. scutata (Perty 1833)SC97
                            |        |  `--+--Eustala Simon 1895SC20, AM11
                            |        |     |    |--*E. anastera (Walckenaer 1841)SC97
                            |        |     |    `--E. sagana (Keyserling 1893) [=Araneus saganus]AM11
                            |        |     `--MetazygiaSC20
                            |        |          |--*M. wittfeldae P.-Cambridge 1903SC97
                            |        |          |--M. gregalis (Pickard-Cambridge 1889)P-GGJ08
                            |        |          `--M. rogenhoferi (Keyserling 1878)P-GGJ08
                            |        `--+--+--+--GasteracanthinaeSC20
                            |           |  |  `--Anepsion Strand 1929SC20, F11
                            |           |  |       |--A. depressum (Thorell 1877)SC97
                            |           |  |       |--A. peltoides (Thorell 1878)SC97
                            |           |  |       `--A. roeweri Chrysanthus 1961SC97
                            |           |  `--+--+--‘Araneus’ dimidiatusSC20
                            |           |     |  `--‘Araneus’ mulierariusSC20
                            |           |     `--+--CyclosiniSC20
                            |           |        `--DolophoneaeSC20
                            |           |             |--PitharatusSC20
                            |           |             `--Dolophones Walckenaer 1837SC20
                            |           |                  |--D. bituberculata Lamb 1911M72
                            |           |                  |--D. coniferaM90
                            |           |                  `--D. maximusB96
                            |           `--+--+--‘Parawixia’ dehaaniSC20
                            |              |  `--Backobourkia Framenau, Dupérré et al. 2010SC20, F11
                            |              |       |--B. brounii (Koch 1871)PVD10
                            |              |       |--B. collina (Keyserling 1886)T13
                            |              |       `--B. heroineSC20
                            |              `--+--Novakiella Court & Forster 1993SC20, F11
                            |                 |    `--*N. trituberculosa (Roewer 1942)PVD10
                            |                 `--+--+--‘Eriophora’ transmarina (Keyserling 1865)SC20, F11 (see below for synonymy)
                            |                    |  `--+--Carepalxis Koch 1872SC20, KK90
                            |                    |     |    |--C. bilobata Koch & Keyserling 1886KK90
                            |                    |     |    |--C. furcula Koch & Keyserling 1886KK90
                            |                    |     |    `--C. tuberculata Koch & Keyserling 1886KK90
                            |                    |     `--Singa Koch 1836SC20, G76
                            |                    |          |--*S. hamata (Clerck 1757)G76 (see below for synonymy)
                            |                    |          |--S. atra (Kulcz. 1885) [=Aranea (Singa) atra]S06
                            |                    |          `--S. nitidula Koch 1845G76 [=Epeira (Singa) nitidulaK01]
                            |                    `--+--Plebs Joseph & Framenau 2012SC20
                            |                       |    |--P. bradleyiSC20
                            |                       |    |--P. cyphoxisSC20
                            |                       |    `--P. eburnus (Keyserling 1886)SC20
                            |                       `--+--‘Aranea’ recherchensis Main 1954M54 [=Araneus recherchensisSC20]
                            |                          `--+--‘Araneus’ senicaudatusSC20
                            |                             `--Acroaspis Karsch 1878SC20, PVD10 (see below for synonymy)
                            |                                  |--*A. olorinaSC20
                            |                                  |--A. decorosa (Urquhart 1894)PVD10 (see below for synonymy)
                            |                                  `--*Heurodes’ turrita Koch & Keyserling 1886KK90
                            `--+--MangoraSC20
                               `--+--+--+--Acusilas Simon 1895SC20, K02
                                  |  |  |    |--A. coccineus Simon 1895K02
                                  |  |  |    `--A. malaccensisJD-S07
                                  |  |  `--+--WiticaSC97 [incl. PhysiolaSC20, SalassinaSC97; Physioleae]
                                  |  |     |    `--*W. crassicauda (Keyserling 1865)SC97
                                  |  |     `--Arachnura Vinson 1863SC20, PVD10 [Arachnureae]
                                  |  |          |--A. feredayi (Koch 1872)PVD10 (see below for synonymy)
                                  |  |          `--A. higginsii (Koch 1873) [=Epeira higginsii]M54
                                  |  `--+--Cyrtophorinae [Cyrtophoreae]FS09
                                  |     |    |--CyrtophoraSC20
                                  |     |    |--Kapogea Levi 1997FS09
                                  |     |    |--Manogea Levi 1997FS09
                                  |     |    |--Cyrtobill Framenau & Scharff 2009FS09
                                  |     |    |    `--*C. darwini Framenau & Scharff 2009FS09
                                  |     |    `--Mecynogea Simon 1903 [incl. Hentzia McCook 1894]FS09
                                  |     |         |--M. lemniscata (Walckenaer 1841)B11
                                  |     |         `--‘Hentzia’ mitrata (Marx 1883)TF98
                                  |     `--ArgiopeaeSC97
                                  |          |--*Neogea egregia (Kulczynski 1911)SC97
                                  |          `--+--ArgiopeBS09
                                  |             `--+--Gea Koch 1843SC97, FS09
                                  |                |    |--*G. spinipes Koch 1843SC97
                                  |                |    |--G. effE95
                                  |                |    `--G. heptagon (Hentz 1850)SC97
                                  |                `--+--‘Argiope’ aurantia Lucas 1833SC20, B11
                                  |                   `--Lariniaria Grasshoff 1970SC20
                                  |                        `--*L. argiopiformis (Bösenberg & Strand 1906)SC20, AM11
                                  `--+--+--Poltys Koch 1843SC20, K92 [incl. Pleuromma Doleschall 1859K92]
                                     |  |    |--*P. illepidus Koch 1843SC20, T94 (see below for synonymy)
                                     |  |    |--P. bimaculatus Koch in Koch & Keyserling 1886KK90
                                     |  |    |--P. columnaris Thor. 1889-1890K92
                                     |  |    |--P. coronatus Koch in Koch & Keyserling 1886KK90
                                     |  |    |--P. keyserlingi Koch in Koch & Keyserling 1886KK90
                                     |  |    |--P. laciniosus Koch in Koch & Keyserling 1886KK90
                                     |  |    `--P. mammeatus Koch in Koch & Keyserling 1886KK90
                                     |  `--+--Perilla Thorell 1895SC20, K02
                                     |     |    `--*P. teres Thorell 1895 [incl. P. cylindrogaster Simon 1909]K02
                                     |     `--+--+--‘Araneus’ mitificus (Simon 1886)SC20, AM11 (see below for synonymy)
                                     |        |  `--‘Araneus’ praesignisSC20
                                     |        `--+--‘Araneus’ rotundulusSC20
                                     |           `--EriovixiaSC20
                                     |                |--E. excelsa (Simon 1889)HU08
                                     |                `--E. laglaizei (Simon 1877)GA11
                                     `--+--+--Metepeira F.O.P.-Cambridge 1903SC20, B11
                                        |  |    |--M. arizonica Chamberlin & Ivie 1942SC97
                                        |  |    |--M. foxiSC97
                                        |  |    |--M. incrassata Pickard-Cambridge 1903B11
                                        |  |    |--M. labyrinthea (Hentz 1847)B11
                                        |  |    `--M. spinipes Pickard-Cambridge 1903B11
                                        |  `--+--Kaira Pickard-Cambridge 1889SC20
                                        |     |    `--*K. alba (Hentz 1850)SC97
                                        |     `--Aculepeira Chamberlain & Ivie 1942SC20, G76
                                        |          |--*A. packardi (Thorell 1875)SC97 [=Epeira packardiiS06]
                                        |          |--A. aculeata (Emerton 1877) [=Epeira aculeata]G76
                                        |          |--A. carbonaria (Koch 1869)G76 (see below for synonymy)
                                        |          `--A. ceropegia (Walckenaer 1802)G76 (see below for synonymy)
                                        `--+--CyrtarachninaeSC20
                                           `--+--+--‘Araneus’ bicentenariusSC20
                                              |  `--+--Araniella Chamberlain & Ivie 1942SC20, G76
                                              |     |    |--*A. displicata (Hentz 1847)G76 (see below for synonymy)
                                              |     |    |--A. alpica (Koch 1869)G76 [=Araneus alpicusG76, Aranea alpicaS06]
                                              |     |    |--A. crispula Tullgren 1952G76
                                              |     |    |--A. cucurbitina (Clerck 1757)SC20, TB81 (see below for synonymy)
                                              |     |    |--A. inconspicua (Simon 1874) [=Araneus inconspicuus]G76
                                              |     |    |--A. opistographa (Kulczynski 1905) (see below for synonymy)G76
                                              |     |    `--A. silesiaca (Fickert 1876)G76
                                              |     `--+--Pycnacantha Blackwall 1865SC20
                                              |        |    `--P. fuscosaSC20
                                              |        `--Larinia Simon 1874SC20, F11
                                              |             |--*L. lineata Lucas 1846SC20
                                              |             |--L. acuticauda Simon 1906 (see below for synonymy)J98
                                              |             |--L. bonnetiSC20
                                              |             |--L. borealis (Hentz 1847)SC97
                                              |             |--L. directa (Hentz 1847)AM11
                                              |             |--L. eburneiventris Simon 1908M90
                                              |             |--L. montagui Hogg 1914T13
                                              |             `--L. phthisica (Koch 1871)F11
                                              `--+--+--Hyposinga Ausserer 1871SC20, G76
                                                 |  |    |--*H. sanguinea (Koch 1845) [=Singa sanguinea]G76
                                                 |  |    |--H. albovittata (Westring 1851) [=Singa albovittata]G76
                                                 |  |    |--H. groenlandica (Simon 1889) [=Aranea groenlandica]S06
                                                 |  |    |--H. heri (Hahn 1836) [=Singa heri]G76
                                                 |  |    |--H. pygmaea (Sundevall 1831)G76 (see below for synonymy)
                                                 |  |    `--H. rubens (Hentz 1847)SC97
                                                 |  `--+--‘Larinia’ jeskoviSC20
                                                 |     `--+--Larinioides Caporiacco 1934SC20, AM11
                                                 |        |    |--L. cornutus (Clerck 1757)SC20, G76 (see below for synonymy)
                                                 |        |    |--L. patagiatus (Clerck 1757)AM11
                                                 |        |    `--L. sclopetarius (Clerck 1757) [=Epeira sclopetaria]AM11
                                                 |        `--Nuctenea Simon 1864SC20, G76 [incl. Cyphepeira Archer 1951G76]
                                                 |             |  i. s.: ‘Cyphepeira’ gracilisG76
                                                 |             |--*N. umbratica (Clerck 1757)G76 (see below for synonymy)
                                                 |             |--N. silvicultrix (Koch 1835) (see below for synonymy)G76
                                                 |             `--+--N. folium (Schrank 1803)G76 (see below for synonymy)
                                                 |                |--N. ixobola (Thorell 1873)G76 (see below for synonymy)
                                                 |                |--N. patagiata (Clerck 1757)G76 (see below for synonymy)
                                                 |                `--N. sclopetaria (Clerck 1757) (see below for synonymy)G76
                                                 `--+--+--Agalenatea redii (Scopoli 1763)SC20, G76 (see below for synonymy)
                                                    |  `--+--‘Araneus’ eburneiventrisSC20
                                                    |     `--NeosconaSC20
                                                    `--+--Gibbaranea Archer 1951SC20, G76
                                                       |    `--*G. bituberculata (Walckenaer 1802)G76, TB81 (see below for synonymy)
                                                       `--+--+--AraneusSC20
                                                          |  `--+--*Pronous tuberculifer Keyserling 1880SC97
                                                          |     `--Cercidia Thorell 1869SC20
                                                          |          `--*C. prominens (Westring 1851)SC97 (see below for synonymy)
                                                          `--+--+--‘Araneus’ talipedatusSC20
                                                             |  `--Paralarinia Grasshoff 1970SC20
                                                             `--+--‘Araneus’ albotriangulusSC20
                                                                `--+--Cryptaranea Court & Forster 1988SC20, PVD10 (see below for synonymy)
                                                                   |    |--*C. invisibilis (Urquhart 1892)PVD10 (see below for synonymy)
                                                                   |    |--C. albolineata (Urquhart 1893)PVD10 (see below for synonymy)
                                                                   |    |--C. atrihastula (Urquhart 1891)PVD10
                                                                   |    |--C. stewartensis Court & Forster 1988PVD10
                                                                   |    |--C. subalpina Court & Forster 1988PVD10
                                                                   |    |--C. subcompta (Urquhart 1887)PVD10 (see below for synonymy)
                                                                   |    `--C. venustula (Urquhart 1891)PVD10 (see below for synonymy)
                                                                   `--+--Colaranea Court & Forster 1988SC20, PVD10
                                                                      |    |--*C. viriditas (Urquhart 1887)PVD10 (see below for synonymy)
                                                                      |    |--C. brunnea Court & Forster 1988PVD10
                                                                      |    |--C. melanoviridis Court & Forster 1988PVD10
                                                                      |    `--C. verutum (Urquhart 1887)PVD10 (see below for synonymy)
                                                                      `--Zealaranea Court & Forster 1988SC20, PVD10
                                                                           |--*Z. crassa (Walckenaer 1841)PVD10 (see below for synonymy)
                                                                           |--Z. prina Court & Forster 1988PVD10
                                                                           |--Z. saxitalis (Urquhart 1887)PVD10 (see below for synonymy)
                                                                           `--Z. trinotata (Urquhart 1890)PVD10 (see below for synonymy)
Araneidae incertae sedis:
  Milonia Thorell 1890K02
    `--M. trifasciata Thorell 1890K02
  Novaranea Court & Forster 1988PVD10
    `--*N. queribunda (Keyserling 1887) [incl. N. laevigata (Urquhart 1891)]PVD10
  CyphalonotusHU08
  Megaraneus Lawrence 1968FS09
    `--M. gabonensis (Lucas 1858) [=Cyrtophora gabonensis]FS09
  Alenatea fuscocolorata (Bösenberg & Strand 1906) [=Araneus fuscocoloratus]AM11
  Edricus Pickard-Cambridge 1890AM11
  Prasonica Simon 1895PVD10
    `--P. plagiata (de Dalmas 1917)PVD10
  Dicrostichus furcatusN91
  Xylethrus [Xylethreae]SC97
    |--*X. superbus Simon 1895SC97
    `--X. scupeus Simon 1895SC97
  Ursa [Urseae]SC97
  Alcimosphenus licinusC01
  CyclosomaP92
  ElucusP92
  EustaloidesP92
  SigaP92
  TheridiometaP92
  MirometaP92
  AraneometaP92
  FossilaraneusP92
  PycnosingaP92
  Singafrotypa Benoit 1962K02
    |--S. acanthopus (Simon 1907)K02
    `--S. okavangoK02
  Zilla Koch 1834G76
    |--Z. diodia (Walckenaer 1802) [=Aranea diodia; incl. *Z. albimacula Koch 1834]G76
    |--Z. californica (Banks 1896) [=Aranea (Zilla) californica]S06
    |--Z. decolorata Keyserling 1893 (n. d.) [=Araneus decoloratus, Zygiella decolorata]L05
    `--Z. dispar (Kulcz. 1885) [=Aranea (Zilla) dispar]S06
  Atea Koch 1837G76
    |--*A. sturmi (Hahn 1831) [=Epeira sturmi]G76
    `--A. triguttata (Fabricius 1775)G76 [=Araneus triguttatusG76, Epeira triguttataV09]
  Mesozygiella dunlopiSC20
  *Colphepeira catawba (Banks 1911)SC97
  BertraneaeSC97
    |--SpintharidiusSC97
    `--Bertrana Keyserling 1884SC97, AM11
         |--*B. striolata Keyserling 1884SC97
         `--B. laselva Levi 1989SC97
  AspidolasiusSC97
    |--*A. branickii (Taczanowski 1879)SC97
    `--A. bifurcatus Simon 1897SC97
  PsylloSC20

Acroaspis Karsch 1878SC20, PVD10 [incl. Heurodes Koch & Keyserling 1886SC20, KK90]

Acroaspis decorosa (Urquhart 1894)PVD10 [=Epeira decorosaNS00, Eriophora decorosaPVD10]

Aculepeira carbonaria (Koch 1869)G76 [=Araneus carbonariusG76, Aranea carbonariaS06]

Aculepeira ceropegia (Walckenaer 1802)G76 [=Araneus ceropegiusG76, Aranea ceropegiaS06, Araneus (Epeira) ceropegiaS95]

Agalenatea redii (Scopoli 1763)SC20, G76 [=Aranea rediiG76, Araneus rediiG76]

Arachnura feredayi (Koch 1872)PVD10 [incl. A. longicauda Urquhart 1885NS00, A. longicauda var. nigritia Urquhart 1885NS00, A. obtusa Urquhart 1885NS00]

Araneus’ mitificus (Simon 1886)SC20, AM11 [=Epeira mitificaTB81; incl. Zilla nawazi Dyal 1935TB81]

Araniella cucurbitina (Clerck 1757)SC20, TB81 [=Aranea cucurbitinaAM11, Araneus cucurbitinusG76, Epeira cucurbitinaV09]

*Araniella displicata (Hentz 1847)G76 [=Epeira displicataG76, Aranea displicataS06, Araneus diplicatusG76]

Araniella opistographa (Kulczynski 1905) [=Araneus cucurbitinus opisthographus]G76

*Cercidia prominens (Westring 1851)SC97 [=Epeira prominensG76, Aranea (Cercidia) prominensS06]

Colaranea verutum (Urquhart 1887)PVD10 [=Epeira verutumNS00; incl. E. verutum var. hastatum Urquhart 1887NS00, E. verutum var. lineola Urquhart 1887NS00]

*Colaranea viriditas (Urquhart 1887)PVD10 [incl. Epeira flavomaculata Urquhart 1890NS00]

Cryptaranea Court & Forster 1988SC20, PVD10 [incl. Holaranea Court & Forster 1988 (n. n.)NS00]

Cryptaranea albolineata (Urquhart 1893)PVD10 [=Epeira albolineataNS00, Holaranea albolineataNS00]

*Cryptaranea invisibilis (Urquhart 1892)PVD10 [=Epeira invisibilisNS00]

Cryptaranea subcompta (Urquhart 1887)PVD10 [=Epeira subcomptaNS00; incl. E. peronginia Urquhart 1894NS00]

Cryptaranea venustula (Urquhart 1891)PVD10 [=Holaranea venustulaNS00; incl. Epeira munda var. inversa Urquhart 1893NS00, E. munda Urquhart 1893NS00]

Eriophora pustulosa (Walckenaer 1842)F11 [=Araneus pustulosusFF99, Aranea pustulosaM54; incl. Epeira blattea Urquhart 1893NS00, Ep. helveoguttata Urquhart 1888NS00, Ep. ostribrunnea Urquhart 1890NS00]

Eriophora’ transmarina (Keyserling 1865)SC20, F11 [=Aranea transmarinaFF99]

*Gibbaranea bituberculata (Walckenaer 1802)G76, TB81 [=Aranea bituberculataTB81, Araneus bituberculatusG76; incl. Aranea alboarcuata Panzer 1804TB81, Epeira bicornis Menge 1866TB81, E. dromedaria Koch 1845TB81, Araneus dromedariusTB81]

Hyposinga pygmaea (Sundevall 1831)G76 [=Araneus (Singa) pygmaeusS95, Epeira (Singa) pygmaeaK01, G76]

Larinia acuticauda Simon 1906 [=Drexelia acuticauda; incl. D. camerunica Strand 1908, Larinia camerunica]J98

Larinioides cornutus (Clerck 1757)SC20, G76 [=Araneus cornutusG76, Cyphepeira cornutaG76, Epeira cornutaK01, Nuctenea cornutaG76; incl. Aranea umbratica Olivier non Araneus umbraticus Clerck 1757K01, Epeira vicaria Kulczyński 1885K01, Araneus (Epeira) vicariusS95]

*Lariniophora ragnhildae (Strand 1917) [=Araneus ragnhildae; incl. Epeira gracilis Hogg 1896 nec Walckenaer 1805 nec Keyserling 1865 nec Menge 1879, A. gracilis]F11

Nuctenea folium (Schrank 1803)G76 [=Araneus foliumG76, Aranea foliumK01, Epeira foliumK01; incl. Aranea foliataK01]

Nuctenea ixobola (Thorell 1873)G76 [=Araneus ixobolusG76, Cyphepeira ixobolaG76, A. (Epeira) ixobolaS95]

Nuctenea patagiata (Clerck 1757)G76 [=Araneus patagiatusG76, Cyphepeira patagiataG76, Epeira patagiataK01; incl. Aranea apoclisaK01, Epeira apoclisaS69]

Nuctenea sclopetaria (Clerck 1757) [=Araneus sclopetarius, Cyphepeira sclopetaria]G76

Nuctenea silvicultrix (Koch 1835) [=Araneus silvicultrix, *Cyphepeira silvicultrix, Epeira silvicultrix]G76

*Nuctenea umbratica (Clerck 1757)G76 [=Araneus umbraticusG76, Chinestela umbraticaG76, Epeira umbraticaV09]

*Poltys illepidus Koch 1843SC20, T94 [incl. *Pleuromma moluccum Doleschall 1859D59, T94, Po. moluccumT94]

*Singa hamata (Clerck 1757)G76 [=Araneus hamatusG76, Epeira (Singa) hamataK01]

*Zealaranea crassa (Walckenaer 1841)PVD10 [incl. Epeira albiscutum Urquhart 1890NS00, E. albostricta Urquhart 1890NS00, E. angusticlava Urquhart 1893NS00, E. bialbimaculata Urquhart 1887NS00, E. dubitabilis Urquhart 1890NS00, E. dumicola Urquhart 1889NS00, E. lineaacuta Urquhart 1887NS00, E. purpura Urquhart 1887NS00]

Zealaranea saxitalis (Urquhart 1887)PVD10 [=Epeira saxitalisNS00]

Zealaranea trinotata (Urquhart 1890)PVD10 [=Epeira trinotataNS00; incl. E. aestiva Urquhart 1890NS00, E. trinotata var. olivinia Urquhart 1890NS00]

*Zygiella atrica (Koch 1845) [=Eucharia atrica, Zilla atrica, Zygia calophylla non Aranea calophylla Walckenaer 1802]G76

*Type species of generic name indicated

References

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