Idiopoidea

Cantuaria sp., copyright Steve Reekie.


Belongs within: Rastelloidina.
Contains: Ctenizinae.

The Idiopoidea are a clade of mygalomorph spiders in which the legs bear tarsal spines, females lack scopulae on the first and second tarsi, and the anterior metatarsi and tibiae of the female possess digging spines (Jocqué & Dippenaar-Schoeman 2007).

<==Idiopoidea JD-S07
    |--Ctenizidae [Ctenizoidea] JD-S07
    |    |  i. s.: Eucyrtops latior K77
    |    |         Bolostromus S93
    |    |           |--B. destructus Wunderlich 1988 S93
    |    |           `--B. suspectus V09
    |    |--Ctenizinae JD-S07
    |    `--Pachylomerinae JD-S07
    |         |--Conothele Thorell 1878 JD-S07
    |         |--Ummidia Thorell 1875 JD-S07
    |         `--Hebestatis Simon 1903 JD-S07
    |              `--H. theveneti JD-S07
    `--Idiopidae JD-S07
         |--Neocteniza mexicana P.-Cambridge 1897 R95
         |--Galeosoma JD-S07
         |--Genysa R14 [Genyseae, Genysinae JD-S07]
         |--Ctenolophus JD-S07
         |--Misgolas rapax (Rainbow 1914) FF99, B14 [incl. Dyarcyops fuscipes B96]
         |--Idiops Perty 1833 [Idiopinae] JD-S07
         |    `--I. bersebaensis Strand 1917 J98
         |--Idiosoma JD-S07
         |    |--I. nigrum Main 1952 DH10
         |    `--I. sigillatum K77
         |--Arbanitis Koch 1874 R14 [Arbanitinae JD-S07, Arbaniteae]
         |    |--*A. longipes Koch 1874 R14
         |    |--A. fuscipes Rainbow 1914 R14
         |    |--A. gracilis (Rainbow & Pulleine 1918) B96
         |    |--A. hoggi K77
         |    |--A. inornatus Y95
         |    `--A. maculipes R14
         `--Cantuaria Hogg 1902 PVD10
              |--*C. dendyi (Hogg 1901) PVD10
              |--C. abdita Forster 1968 PVD10
              |--C. allani Forster 1967 PVD10
              |--C. aperta Forster 1968 PVD10
              |--C. apica Forster 1968 PVD10
              |--C. assimilis Forster 1968 [=Misgolas assimilis] PVD10
              |--C. borealis Forster 1968 PVD10
              |--C. catlinensis Forster 1968 PVD10
              |--C. cognata Forster 1968 PVD10
              |--C. collensis (Todd 1945) PVD10
              |--C. delli Forster 1968 PVD10
              |--C. depressa Forster 1968 PVD10
              |--C. dunedinensis Forster 1968 PVD10
              |--C. gilliesii (Pickard-Cambridge 1878) PVD10
              |--C. grandis Forster 1968 PVD10
              |--C. huttonii (Pickard-Cambridge 1879) PVD10 [=Misgolas huttoni H04]
              |--C. insulana Forster 1968 PVD10 [=Misgolas insulanus NS00]
              |--C. isolata Forster 1968 PVD10
              |--C. johnsi Forster 1968 PVD10 [=Misgolas johnsi NS00]
              |--C. kakahuensis Forster 1968 PVD10
              |--C. kakanuiensis Forster 1968 PVD10 [=Misgolas kakanuiensis NS00]
              |--C. lomasi Forster 1968 PVD10
              |--C. magna Forster 1968 PVD10
              |--C. marplesi (Todd 1945) PVD10
              |--C. maxima Forster 19068 PVD10
              |--C. medialis Forster 1968 PVD10
              |--C. minor Forster 1968 PVD10
              |--C. myersi Forster 1968 PVD10
              |--C. napua (Forster 1968) PVD10 [=Misgolas napua H04]
              |--C. orepukiensis Forster 1968 PVD10
              |--C. parrotti Forster 1968 PVD10
              |--C. pilama Forster 1968 PVD10
              |--C. prina Forster 1968 PVD10
              |--C. reducta Forster 1968 PVD10 [=Misgolas reductus NS00]
              |--C. secunda Forster 1968 PVD10
              |--C. sinclairi Forster 1968 PVD10
              |--C. stephenensis Forster 1968 PVD10 [=Misgolas stephenensis NS00]
              |--C. stewarti (Todd 1945) PVD10
              |--C. sylvatica Forster 1968 PVD10
              |--C. toddae Forster 1968 PVD10 [=Misgolas toddae FF99]
              |--C. vellosa Forster 1968 PVD10 [=Misgolas vellosa FF99]
              `--C. wanganuiensis (Todd 1945) PVD10

*Type species of generic name indicated

REFERENCES

[B96] Brunet, B. 1996. Spiderwatch: A Guide to Australian Spiders. Reed New Holland: Sydney.

[DH10] Durrant, B. J., M. S. Harvey, V. W. Framenau, R. Ott & J. M. Waldock. 2010. Patterns in the composition of ground-dwelling spider communities in the Pilbara bioregion, Western Australia. Records of the Western Australian Museum Supplement 78 (1): 185–204.

[FF99] Forster, R., & L. Forster. 1999. Spiders of New Zealand and their World-wide Kin. University of Otago Press: Dunedin (New Zealand).

[H04] Harris, A. C. 2004. Priocnemis (Trichochurgus) monachus (Smith) (Hymenoptera: Pompilidae) preying on Misgolas napua Forster (Mygalomorphae: Idiophidae) after opening the lids of the prey’s burrows, and notes of the affinities of Trichocurgus Haupt. Weta 27: 13–14.

[J98] Jäger, P. 1998. Das Typenmaterial der Spinnentiere (Arachnida: Acari, Amblypygi, Araneae, Opiliones, Pseudoscorpiones, Scorpiones, Uropygi) aus dem Museum Wiesbaden. Jahrbuecher des Nassauischen Vereins fuer Naturkunde 119: 81–91.

[JD-S07 Jocqué, R., & A. S. Dippenaar-Schoeman. 2007. Spider Families of the World. Royal Museum for Central Africa: Tervuren (Belgium).

[K77] Koch, L. E. 1977. The taxonomy, geographic distribution and evolutionary radiation of Australo-Papuan scorpions. Records of the Western Australian Museum 5 (2): 83–367.

[NS00] Nicholls, D. C., P. J. Sirvid, S. D. Pollard & M. Walker. 2000. A list of arachnid primary types held in Canterbury Museum. Records of the Canterbury Museum 14: 37–48.

[PVD10] Paquin, P., C. J. Vink & N. Dupérré. 2010. Spiders of New Zealand: annotated family key and species list. Manaaki Whenua Press: Lincoln (New Zealand).

[R14] Rainbow, W. J. 1914. Studies in Australian Araneidae, no. 6, the Terretelariae. Records of the Australian Museum 10 (8): 187–270.

[R95] Raven, R. J. 1995. Coxal glands of spiders of the genera Bymainiella, Atrax and Namea (Hexathelidae, Dipluridae, Mygalomorphae). Records of the Western Australian Museum Supplement 52: 67–71.

[S93] Selden, P. A. 1993. Arthropoda (Aglaspidida, Pycnogonida and Chelicerata). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 297–320. Chapman & Hall: London.

[V09] Verdcourt, B. (ed.) 2009. Additions to the Wild Fauna and Flora of the Royal Botanic Gardens, Kew. XXVI. Miscellaneous records. Kew Bulletin 64 (1): 183–194.

[Y95] Yen, A. L. 1995. Australian spiders: an opportunity for conservation. Records of the Western Australian Museum Supplement 52: 39–47.

Basalhaplogynae

Desert bush spider Diguetia canities, from the National Park Service.


Belongs within: Araneida.
Contains: Scytodoidea, Pholcidae, Segestriidae, Dysderidae, Orsolobidae, Oonopidae.

The Basalhaplogynae are a clade of spiders in which the chelicerae are fused at the base and provided with a lamina; members of the clade also have the tegulum and subtegulum fused and the spinnerets lack tartipores (Jocqué & Dippenaar-Schoeman 2007).

<==Basalhaplogynae [Haplogynae, Pholcoidea] W19
    |--Filistatidae [Filistatomorpha] JD-S07
    |    |  i. s.: Misionella P05
    |    |           |--M. didicostae Penney 2005 P05
    |    |           `--M. mendensis R14
    |    |         Microfilistata Zohnstein 1990 G95
    |    |--Filistatinae G95
    |    |    |  i. s.: Zaituna Lehtinen 1967 G95
    |    |    |--Sahastata Benoit 1968 G95
    |    |    |    |--S. nigra G95
    |    |    |    `--S. sabaea Brignoli 1982
    |    |    `--+--Kukulcania Lehtinen 1967 G95
    |    |       |    `--K. hibernalis G95
    |    |       `--Filistata Latreille 1810 G95, JD-S07
    |    |            |--F. hibernalis B06
    |    |            `--F. insidiatrix (Forskål 1775) MG03
    |    `--Prithinae G95
    |         |  i. s.: Malalistata Mello-Leitão 1946 G95
    |         |         Tricalamus Wang 1987 G95
    |         |--+--Filistatinella Gertsch & Ivie 1936 G95
    |         |  `--+--Filistatoides P.-Cambridge 1899 G95
    |         |     `--Pikelinia Mello-Leitão 1946 G95
    |         |          `--P. tambilloi R14
    |         `--+--+--Andoharano Lehtinen 1967 G95
    |            |  `--Afrofilistata Benoit 1968 G95
    |            `--+--Pritha Lehtinen 1967 G95, MG03
    |               |    |--*P. nana (Simon 1868) MG03
    |               |    `--P. crosbyi (Spassky 1938) [=Filistata crosbyi] MG03
    |               `--+--Yardiella Gray 1994 G95
    |                  `--Wandella Gray 1994 G95
    |                       |--W. orana G95
    |                       `--W. waldockae MC13
    `--Synspermiata [Scytodiformes] W19
         |--+--Scytodoidea JD-S07
         |  `--+--Pholcidae JD-S07
         |     `--+--Plectreuridae JD-S07
         |        |    |--Kibramoa Chamberlin 1924 JD-S07
         |        |    `--Plectreurys Simon 1893 JD-S07
         |        |         `--P. tristis Simon 1893 HF08
         |        `--Diguetidae JD-S07
         |             |--Segestrioides Keyserling 1883 JD-S07
         |             |    `--S. bicolor JD-S07
         |             `--Diguetia Simon 1895 JD-S07
         |                  |--D. canities JD-S07
         |                  `--D. signata Gertsch 1958 HF08
         `--+--+--Dysderoidea JD-S07
            |  |    |  i. s.: Trogloraptor [Trogloraptoridae] R14
            |  |    |--Segestriidae JD-S07
            |  |    `--+--Dysderidae JD-S07
            |  |       `--+--Orsolobidae JD-S07
            |  |          `--Oonopidae JD-S07
            |  `--Tetrablemmidae JD-S07
            |       |--Shearella JD-S07
            |       |--Afroblemma JD-S07
            |       |--Tetrablemma P.-Cambridge 1873 [Tetrablemminae] JD-S07
            |       |--Paculla [Pacullinae] JD-S07
            |       `--Monoblemma spinosum Wunderlich 1988 S93
            `--Caponiidae JD-S07
                 |--Caponia Simon 1887 [Caponiinae] JD-S07
                 |--Calponia Platnick 1993 JD-S07
                 |--Diploglena Purcell 1904 JD-S07
                 |--Nopsides Chamberlin 1924 JD-S07
                 |--Notnops Platnick 1994 JD-S07
                 |--Orthonops Chamberlin 1924 JD-S07
                 |--Taintnops Platnick 1994 JD-S07
                 |--Tarsonops Platnick 1994 JD-S07
                 |--Tisentnops Platnick 1994 JD-S07
                 |--Caponina Simon 1887 JD-S07
                 |    `--C. notabilis (Mello-Leitão 1939) FM11
                 `--Nops MacLeay 1839 [Nopinae] JD-S07
                      |--N. lobatus Wunderlich 1988 S93
                      `--N. segmentatus Wunderlich 1988 S93

*Type species of generic name indicated

REFERENCES

[B06] Banks, N. 1906. Arachnida from the Bahamas. Bulletin of the American Museum of Natural History 22: 185–189.

[FM11] Freire-Jr, G. de B., & P. C. Motta. 2011. Effects of experimental fire regimes on the abundance and diversity of cursorial arachnids of Brazilian savannah (cerrado biome). Journal of Arachnology 39 (2): 263–272.

[G95] Gray, M. R. 1995. Morphology and relationships within the spider family Filistatidae (Araneae: Araneomorphae). Records of the Western Australian Museum Supplement 52: 79–89.

[HF08] Huber, B. A., & N. Fleckenstein. 2008. Comb-hairs on the fourth tarsi in pholcid spiders. Journal of Arachnology 36 (2): 232–240.

[JD-S07] Jocqué, R., & A. S. Dippenaar-Schoeman. 2007. Spider Families of the World. Royal Museum for Central Africa: Tervuren (Belgium).

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[MG03] Marusik, Yu. M., & E. F. Guseinov. 2003. Spiders (Arachnida: Aranei) of Azerbaijan. 1. New family and genus records. Arthropoda Selecta 12 (1): 29–46.

[P05] Penney, D. 2005. Fossil blood droplets in Miocene Dominican amber yield clues to speed and direction of resin secretion. Palaeontology 48 (5): 925–927.

[R14] Ramírez, M. J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374.

[S93] Selden, P. A. 1993. Arthropoda (Aglaspidida, Pycnogonida and Chelicerata). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 297–320. Chapman & Hall: London.

[W19] Wunderlich, J. 2019. What is a spider? Cretaceous fossils modify strongly phylogenetics as well as diagnoses of families, superfamilies and even suborders of spiders (Araneida) and other arthropods. Beiträge zur Araneologie 12: 1–32.

Scytodidae

Scytodes thoracica, copyright Tone Killick.


Belongs within: Scytodoidea.

The Scytodidae, spitting spiders, are a cosmopolitan (mostly tropical) group of spiders that capture prey by spraying gluey silk over it (Jocqué & Dippenaar-Schoeman 2007).

Characters (from Jocqué & Dippenaar-Schoeman 2007): Small to medium-sized; three tarsal claws; ecribeelate; haplogyne; six eyes in three diads; thoracic region domed, containing prosomal glands producing gluey silk; legs long and slender; body decorated with symmetrical dark pattern.

Scytodidae [Retitelariae]
    |--Phalangopus subtilis Menge 1854 D06
    |--Scyloxes Dunin 1992 JD-S07
    |--Soeuria Saaristo 1997 JD-S07
    |--Stedocys Ono 1995 JD-S07
    |    `--S. leopoldi R14
    |--Dictis Koch 1872 JD-S07
    |    |--D. domestica (Doleschall 1859) [=Scytodes domestica; incl. D. fumida Thorell 1891] T94
    |    `--D. striatipes Koch 1872 PVD10
    `--Scytodes Latreille 1804 JD-S07
         |--*S. thoracica (Latreille 1804) [=Aranea thoracica] C01
         |--S. bergeri Strand 1915 J98
         |--S. distincta Lucas 1846 E12
         |--S. fusca D95
         |--S. intricata R14
         |--S. itapevi Brescovit & Rheims 2000 FM11
         |--S. longipes B06
         |--S. marmorata Koch 1872 S99
         |--S. pallida Doleschall 1859 D59
         |--S. piliformis Wunderlich 1988 S93
         |--S. planithorax Wunderlich 1988 S93
         |--S. propinqua Stoliczka 1869 D59
         `--S. stridulans Wunderlich 1988 S93

*Type species of generic name indicated

REFERENCES

[B06] Banks, N. 1906. Arachnida from the Bahamas. Bulletin of the American Museum of Natural History 22: 185–189.

[C01] Cambridge, F. O. P. 1901. A revision of the genera of the Araneae or spiders with reference to their type species. Annals and Magazine of Natural History, series 7, 7: 51–65.

[D59] Doleschall, C. L. 1859. Tweede Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Verhandelingen der Natuurkundige Vereeniging in Nederlandsch Indie [Acta Societatis Scientiarum Indo-Neêrlandicae] 5 (5): 1–60, pls 1–18.

[D95] Downes, M. F. 1995. Australasian social spiders: what is meant by 'social'? Records of the Western Australian Museum Supplement 52: 25–32.

[D06] Dunlop, J. A. 2006. Baltic amber harvestman types (Arachnida: Opiliones: Eupnoi and Dyspnoi). Fossil Record 9 (2): 167–182.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[FM11] Freire-Jr, G. de B., & P. C. Motta. 2011. Effects of experimental fire regimes on the abundance and diversity of cursorial arachnids of Brazilian savannah (cerrado biome). Journal of Arachnology 39 (2): 263–272.

[J98] Jäger, P. 1998. Das Typenmaterial der Spinnentiere (Arachnida: Acari, Amblypygi, Araneae, Opiliones, Pseudoscorpiones, Scorpiones, Uropygi) aus dem Museum Wiesbaden. Jahrbuecher des Nassauischen Vereins fuer Naturkunde 119: 81–91.

[JD-S07] Jocqué, R., & A. S. Dippenaar-Schoeman. 2007. Spider Families of the World. Royal Museum for Central Africa: Tervuren (Belgium).

[PVD10] Paquin, P., C. J. Vink & N. Dupérré. 2010. Spiders of New Zealand: annotated family key and species list. Manaaki Whenua Press: Lincoln (New Zealand).

[R14] Ramírez, M. J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374.

[S93] Selden, P. A. 1993. Arthropoda (Aglaspidida, Pycnogonida and Chelicerata). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 297–320. Chapman & Hall: London.

[S99] Simon, E. 1899. Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896–1897. Zoologische Jahrbücher, Abteilung für Systematik, Geographie un Biologie der Thiere 12 (4): 411–437.

[T94] Thorell, T. 1894. Förteckning öfver Arachnider från Java och närgränsande öar, insamlade af docenten D:r Carl Aurivillius; jemte beskrifingar å några sydasiatiska och sydamerikanska Spindlar. Bihang till K. Svenska Vetenskaps-Akademiens Handlingar 20 pt 4 (4): 1–63.

Neocribellatae

Lampshade spider Hypochilus thorelli, copyright Marshal Hedin.


Belongs within: Araneida.
Contains: Argyronetidae, Hahniidae, Leptonetoidea, Plectana, Palpimanoidea, Eresoidea, Nicodamidae, Araneoidea, Austrochiloidea, Deinopoidea, Dictynidae, Amaurobioidea, Phyxelididae.

The Neocribellatae are a clade of spiders in which a sieve-like spinning plate, the cribellum, evolved in front of the spinnerets, though this feature has been secondarily lost in many derived subgroups. Members of this clade have a relatively short life cycle and do not moult subsequent to attaining maturity (Wunderlich 2019). Many members of this clade (most members of the subclade Praeentelegynae) have an entelegyne female reproductive system, with an epigyne bearing separate pairs of ducts for sperm transport during insemination and fertilisation, though it is debated whether such a system has arisen more than once.

<==Neocribellatae [Cleistospermiata] W19
    |  i. s.: Spinipalpus [Pholcochyroceridae] W19
    |           `--S. veta Wunderlich 2015 W19
    |--Hypochilidae [Palaearchaeidea, Paleocribellatae] W19
    |    |--Ectatosticta Simon 1892 [Ectatostictidae] JD-S07
    |    |    `--E. davidi A99
    |    `--Hypochilus Marx 1888 JD-S07
    |         |--H. coylei HR01
    |         |--H. gertschi FF99
    |         |--H. pococki FF99
    |         |--H. sheari HR01
    |         `--H. thorelli Marx 1888 HF08
    `--Praeentelegynae (see below for synonymy) W19
         |  i. s.: Argyronetidae S02
         |         Hahniidae S02
         |         Eurocoelotes inermis (Koch 1855) SL14
         |         Homalonychus Marx 1891 [Homalonychidae] JD-S07
         |           |--H. selenopoides R14
         |           `--H. theologus JD-S07
         |         Leptonetoidea W19
         |         Plectana D59
         |         Notocentria Thorell 1894 T94
         |           `--*N. sexspinosa Thorell 1894 T94
         |         Talthybia depressa Thorell 1898 GA11
         |--Palpimanoidea R14
         `--+--Eresoidea R14
            `--+--+--Nicodamidae R14
               |  `--+--Araneoidea R14
               |     `--+--Austrochiloidea R14
               |        `--+--Deinopoidea R14
               |           `--Dictynidae R14
               `--+--Amaurobioidea R14
                  `--Titanoecoidea JD-S07
                       |--Phyxelididae JD-S07
                       `--Titanoecidae JD-S07
                            |--Goeldia Keyserling 1891 JD-S07
                            |--Anuvinda Lehtinen 1967 JD-S07
                            |--Nurscia Simon 1874 JD-S07
                            |--Pandava Lehtinen 1967 JD-S07
                            `--Titanoeca Thorell 1869 R14, JD-S07
                                 |--T. americana R14
                                 `--T. obscura JD-S07

Praeentelegynae [Dictynoidea, Dipneumonomorpha, Entelegynae, Euetrioidae, Orbitelae, Orbiculariae, Orbitelariae] W19

*Type species of generic name indicated

REFERENCES

[A99] Ax, P. 1999. Das System der Metazoa II. Ein Lehrbuch der phylogenetischen Systematik. Gustav Fisher Verlag: Stuttgart (transl. 2000. Multicellular Animals: The phylogenetic system of the Metazoa vol. 2. Springer).

[D59] Doleschall, C. L. 1859. Tweede Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Verhandelingen der Natuurkundige Vereeniging in Nederlandsch Indie [Acta Societatis Scientiarum Indo-Neêrlandicae] 5 (5): 1–60, pls 1–18.

[FF99] Forster, R., & L. Forster. 1999. Spiders of New Zealand and their World-wide Kin. University of Otago Press: Dunedin (New Zealand).

[GA11] Gregorič, M., I. Agnarsson, T. A. Blackledge & M. Kuntner. 2011. Darwin's bark spider: giant prey in giant orb webs (Caerostris darwini, Araneae: Araneidae)? Journal of Arachnology 39 (2): 287–295.

[HR01] Hale, R. J., & D. C. F. Rentz. 2001. The Gryllacrididae: an overview of the world fauna with emphasis on Australian examples. In: Field, L. H. (ed.) The Biology of Wetas, King Crickets and Their Allies pp. 95–110. CABI Publishing: Wallingford (UK).

[HF08] Huber, B. A., & N. Fleckenstein. 2008. Comb-hairs on the fourth tarsi in pholcid spiders. Journal of Arachnology 36 (2): 232–240.

[JD-S07] Jocqué, R., & A. S. Dippenaar-Schoeman. 2007. Spider Families of the World. Royal Museum for Central Africa: Tervuren (Belgium).

[R14] Ramírez, M. J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374.

[SL14] Samu, F., G. Lengyel, É. Szita, A. Bidló & P. Ódor. 2014. The effect of forest stand characteristics on spider diversity and species composition in deciduous-coniferous mixed forests. Journal of Arachnology 42 (2): 135–141.

[S02] Selden, P. A. 2002. Missing links between Argyroneta and Cybaeidae revealed by fossil spiders. Journal of Arachnology 30 (2): 189–200.

[T94] Thorell, T. 1894. Förteckning öfver Arachnider från Java och närgränsande öar, insamlade af docenten D:r Carl Aurivillius; jemte beskrifingar å några sydasiatiska och sydamerikanska Spindlar. Bihang till K. Svenska Vetenskaps-Akademiens Handlingar 20 pt. 4 (4): 1–63.

[W19] Wunderlich, J. 2019. What is a spider? Cretaceous fossils modify strongly phylogenetics as well as diagnoses of families, superfamilies and even suborders of spiders (Araneida) and other arthropods. Beiträge zur Araneologie 12: 1–32.

Araneida

Chimerarachne yingi, copyright Bo Yang.


Belongs within: Tetrapulmonata.
Contains: Mesothelae, Mygalomorphae, Basalhaplogynae, Neocribellatae.

The Araneida are a clade of arachnids including the spiders (Araneae) and the Cretaceous Chimerarachne, united by the organisation of the openings of the silk glands into a group of spinnerets at the end of the opisthosoma, and modification of the male pedipalp as an organ for sperm transfer. Chimerarachne yingi is known from the mid-Cretaceous Burmese amber and differs from modern spiders in retaining a long terminal flagellum (Wunderlich 2019). The tarsus of the pedipalp bears a pair of apical apophyses in some but not all of the known specimens; the modified pedipalp has been interpreted as a sperm transfer organ like that of modern male spiders. Modern spiders are also characterised by the modification of the chelicerae to bear venomous fangs. In most living spiders, forming the clade Opisthothelae, all trace of external segmentation has been lost from the abdomen. The fangs are plesiomorphically parallel with one another but in members of the clade Araneomorphae they close towards the midline of the body.

Araneida W19
    |--Chimerarachne Wang et al. 2018 [Chimerarachnida, Chimerarachnidae] W19
    |    `--C. yingi Wang et al. 2018 W19
    `--Araneae [Arachnides, Aranea, Aranina, Artharachnae, Parallelodontes] GD16
         |  i. s.: Archoleptona schusteri UB02
         |         Blabomma UB02
         |         Bryantella smaragdus (Crane 1945) [incl. Parnaenus convexus Chickering 1946] J02
         |         Pterinochilus WS02
         |           |--P. carnivorus Strand 1917 J98
         |           `--P. nigrofulvus (Pocock 1898) WS02
         |         Lophomma punctatum RKD02
         |         Tholia KK90
         |           |--T. conifera Koch & Keyserling 1886 KK90
         |           |--T. mammeata Koch & Keyserling 1886 KK90
         |           |--T. peltata Koch in Koch & Keyserling 1886 KK90
         |           |--T. pilosa Koch & Keyserling 1886 KK90
         |           |--T. simpla Koch & Keyserling 1886 KK90
         |           `--T. tuberculata Koch in Koch & Keyserling 1886 KK90
         |         Heurodes Koch & Keyserling 1886 KK90
         |           `--*H. turrita Koch & Keyserling 1886 KK90
         |         Carepalxis Koch 1872 KK90
         |           |--C. bilobata Koch & Keyserling 1886 KK90
         |           |--C. furcula Koch & Keyserling 1886 KK90
         |           `--C. tuberculata Koch & Keyserling 1886 KK90
         |         Ulesanis Koch 1872 KK90
         |           `--U. sextuberculata Koch & Keyserling 1890 KK90
         |         Wirada Keyserling 1886 KK90
         |           `--W. rotunda Koch & Keyserling 1890 KK90
         |         Tobesoa Koch & Keyserling 1890 KK90
         |           `--*T. theridioides Koch & Keyserling 1890 KK90
         |         Diaca insulana Koch & Keyserling 1890 KK90
         |         Gmogala Koch & Keyserling 1890 [=Imogala (l. c.)] KK90
         |           `--*G. scarabaea Koch & Keyserling 1890 [=Imogala (l. c.) scarabaea] KK90
         |         Euro permunda B96
         |         Daramulunia [=Daramuliana (l. c.)] C90
         |         Pyritaraneidae D07
         |           |--Pyritaranea tubifera F04
         |           `--Dinopilio Fritsch 1904 D07, F04
         |                `--*D. gigas Fritsch 1904 F04
         |         Amphitrogulus sternalis Gourret 1886 D07
         |         Phalangillum hirsutum Gourret 1886 D07
         |         Tidarren sisyphoides CM07
         |         Novakia ‘trituberculata’ (Urquhart 1887) [=Epeira trituberculata non Lucas 1846] NS00
         |         Phalaea marginata Strand 1907 J98
         |         Barusia maheni (Kratochvíl & Miller 1939) UO05
         |         Hadites tegenarioides Keyserling 1862 UO05
         |         Ariadne Doleschall 1857 D57
         |           `--*A. flagellum Doleschall 1857 D57
         |         Sarinda linda C08
         |         Perneria Fritsch 1904 F04
         |           `--*P. salticoides [=Arthrolycosa salticoides] F04
         |         Pleurolycosa Fritsch 1904 F04
         |           `--*P. prolifera [=Arthrolycosa prolifera] F04
         |         Acrosoma H04
         |           |--A. bifurcatum H04
         |           |--A. hexacanthum H04
         |           `--A. spinosum H04
         |         Troglodiplura lowryi Y95
         |         Desidiopsis racovitzai PP64
         |         Sanogasta backhauseni R14
         |         Cupa kalawitana R14
         |         Acanthodon P01
         |           |--A. flaveolum Pocock 1901 P01
         |           `--A. thorellii P01
         |         Heligmomerus P01
         |           |--H. deserti Pocock 1901 P01
         |           `--H. somalicus P01
         |--Mesothelae S02
         `--Opisthothelae S02
              |--Mygalomorphae S02
              `--Araneomorphae (see below for synonymy) S02
                   |  i. s.: Phrynarachne decipiens B96
                   |         Dendrolycosa Doleschall 1859 D59
                   |           `--*D. fusca Doleschall 1859 D59
                   |--Basalhaplogynae W19
                   `--Neocribellatae W19

Araneomorphae [Antiodontes, Arachnomorphae, Araneoclada, Hypochilomorphae, Inaequitelae, Tubitelariae] S02

*Type species of generic name indicated

REFERENCES

[B96] Brunet, B. 1996. Spiderwatch: A Guide to Australian Spiders. Reed New Holland: Sydney.

[C08] Ceccarelli, F. S. 2008. Behavioral mimicry in Myrmarachne species (Araneae, Salticidae) from north Queensland, Australia. Journal of Arachnology 36 (2): 344–351.

[C90] Coddington, J. A. 1990. Book review: Advances in spider taxonomy 1981–1987: a supplement to Brignoli’s A catalogue of the Araneae described between 1940 and 1981 (edited by P. Merrett), by Norman I. Platnick. Journal of Arachnology 18: 245–248.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[D57] Doleschall, C. L. 1857. Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch Indië, series 3, 3 (5–6): 399–434, pls 1–2.

[D59] Doleschall, C. L. 1859. Tweede Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Verhandelingen der Natuurkundige Vereeniging in Nederlandsch Indie [Acta Societatis Scientiarum Indo-Neêrlandicae] 5 (5): 1–60, pls 1–18.

[D07] Dunlop, J. A. 2007. Paleontology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 247–265. Harvard University Press: Cambridge (Massachusetts).

[F04] Fritsch, A. 1904. Palaeozoische Arachniden. Selestverlag: Prague.

[GD16] Garwood, R. J., J. A. Dunlop, P. A. Selden, A. R. T. Spencer, R. C. Atwood, N. T. Vo & M. Drakopoulos. 2016. Almost a spider: a 305-million-year-old fossil arachnid and spider origins. Proceedings of the Royal Society of London Series B—Biological Sciences 283: 20160125.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[J98] Jäger, P. 1998. Das Typenmaterial der Spinnentiere (Arachnida: Acari, Amblypygi, Araneae, Opiliones, Pseudoscorpiones, Scorpiones, Uropygi) aus dem Museum Wiesbaden. Jahrbuecher des Nassauischen Vereins fuer Naturkunde 119: 81–91.

[J02] Jocqué, R. 2002. Genitalic polymorphism—a challenge for taxonomy. Journal of Arachnology 30 (2): 298–306.

[KK90] Koch, L., & E. Keyserling. 1884–1890. Die Arachniden Australiens nach der Natur beschrieben und abgebildet vol. 2. Bauer & Raspe: Nürnberg.

[NS00] Nicholls, D. C., P. J. Sirvid, S. D. Pollard & M. Walker. 2000. A list of arachnid primary types held in Canterbury Museum. Records of the Canterbury Museum 14: 37–48.

[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d'Endoume, Bulletin 31 (27): 5–137.

[P01] Pocock, R. I. 1901. Descriptions of some new African Arachnida. Annals and Magazine of Natural History, series 7, 7: 284–288.

[R14] Ramírez, M. J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374.

[RKD02] Relys, V., S. Koponen & D. Dapkus. 2002. Annual differences and species turnover in peat bog spider communities. Journal of Arachnology 30 (2): 416–424.

[S02] Selden, P. A. 2002. First British Mesozoic spider, from Cretaceous amber of the Isle of Wight, southern England. Palaeontology 45 (5): 973–983.

[UB02] Ubick, D., & T. S. Briggs. 2002. The harvestman family Phalangodidae 4. A review of the genus Banksula (Opiliones, Laniatores). Journal of Arachnology 30 (2): 435–451.

[UO05] Ubick, D., & R. Ozimec. 2005. On the harvestman genus Lola Kratochvíl (Opiliones: Laniatores). Natura Croatica 14 (3): 161–174.

[WS02] Whitmore, C., R. Slotov, T. E. Crouch & A. S. Dippenaar-Schoeman. 2002. Diversity of spiders (Araneae) in a savanna reserve, Northern Province, South Africa. Journal of Arachnology 30 (2): 344–356.

[W19] Wunderlich, J. 2019. What is a spider? Cretaceous fossils modify strongly phylogenetics as well as diagnoses of families, superfamilies and even suborders of spiders (Araneida) and other arthropods. Beiträge zur Araneologie 12: 1–32.

[Y95] Yen, A. L. 1995. Australian spiders: an opportunity for conservation. Records of the Western Australian Museum Supplement 52: 39–47.

Last updated: 12 October 2019.

Micropholcommatidae

Rayforstia vulgaris, from Rix & Harvey (2010).


Belongs within: Palpimanoidea.

The Micropholcommatidae are a group of minute spiders with a strongly raised carapace found in South America and Australasia (Jocqué & Dippenaar-Schoeman 2007).

Characters (from Jocqué & Dippenaar-Schoeman 2007): Very small; three tarsal claws; ecribellate; entelegyne; six or eight eyes; cheliceral gland mound elevated; anterior booklungs reduced to either a few leaves or true tracheae.

<==Micropholcommatidae
    |  i. s.: Raveniella T13
    |--Taphiassa Simon 1880 PVD10 [incl. Parapua Forster 1959 PVD10, JD-S07; Taphiassinae PVD10]
    |    `--T. punctata (Forster 1959) [=Parapua punctata] PVD10
    |--Textricellinae [Textricellidae] PD07
    |    |--Tricella Forster & Platnick 1989 JD-S07
    |    |--Eterosonycha Butler 1932 JD-S07
    |    |--Cenotextricella Penney, Dierick et al. 2007 PD07
    |    |    `--*C. simoni Penney, Dierick et al. 2007 PD07
    |    `--Textricella Hickman 1945 JD-S07
    |         |--T. complexa PD07
    |         |--T. fulva S03
    |         |--T. luteola R14
    |         |--T. parva S03
    |         |--T. plebeia F64
    |         |--T. salmoni FF99
    |         |--T. vulgaris Forster 1959 S03
    |         `--T. wisei Forster 1964 [=T. campbellensis Forster 1964 (n. n.)] F64
    `--Micropholcommatinae JD-S07
         |--Tricellina Forster & Platnick 1989 S03
         |--Olgania Hickman 1979 JD-S07
         |--Pua Forster 1959 JD-S07
         |    `--*P. novaezealandiae Forster 1959 PVD10
         |--Teutoniella Brignoli 1981 JD-S07
         |    `--T. cekalovici JD-S07
         |--Algidiella Rix & Harvey 2010 PVD10
         |    `--*A. aucklandica (Forster 1955) [=Textricella aucklandica] PVD10
         |--Tinytrella Rix & Harvey 2010 PVD10
         |    `--*T. pusilla (Forster 1959) [=Textricella pusilla] PVD10
         |--Taliniella Rix & Harvey 2010 PVD10
         |    |--*T. nigra (Forster 1959) [=Textricella nigra] PVD10
         |    `--T. vinki Rix & Harvey 2010 PVD10
         |--Micropholcomma Crosby & Bishop 1927 [incl. Microlinypheus Butler 1932, Plectochetos Butler 1932] R08
         |    |--*M. caeligenum Crosby & Bishop 1927 R08
         |    |--M. bryophilum (Butler 1932) [=*Microlinypheus bryophilus] R08
         |    |--M. linnaei Rix 2008 R08
         |    |--M. longissimum (Butler 1932) [=*Plectochetos longissimus] R08
         |    |--M. mirum Hickman 1944 R08
         |    |--M. parmatum Hickman 1944 R08
         |    `--M. turbans Hickman 1981 R08
         `--Rayforstia Rix & Harvey 2010 PVD10
              |--*R. vulgaris (Forster 1959) PVD10
              |--R. antipoda (Forster 1959) PVD10
              |--R. insula (forster 1959) PVD10
              |--R. mcfarlanei (Forster 1959) PVD10
              |--R. plebeia (Forster 1959) PVD10
              |--R. propinqua (Forster 1959) PVD10
              |--R. salmoni (Forster 1959) PVD10
              |--R. scuta (Forster 1959) PVD10
              |--R. signata (Forster 1959) PVD10
              `--R. wisei (Forster 1964) PVD10

*Type species of generic name indicated

REFERENCES

[F64] Forster, R. R. 1964. The Araneae and Opiliones of the subantarctic islands of New Zealand. Pacific Insects Monograph 7: 58–115.

[FF99] Forster, R., & L. Forster. 1999. Spiders of New Zealand and their World-wide Kin. University of Otago Press: Dunedin (New Zealand).

[JD-S07] Jocqué, R., & A. S. Dippenaar-Schoeman. 2007. Spider Families of the World. Royal Museum for Central Africa: Tervuren (Belgium).

[PVD10] Paquin, P., C. J. Vink & N. Dupérré. 2010. Spiders of New Zealand: annotated family key and species list. Manaaki Whenua Press: Lincoln (New Zealand).

[PD07] Penney, D., M. Dierick, V. Cnudde, B. Masschaele, J. Vlassenbroeck, L. van Hoorebeke & P. Jacobs. 2007. First fossil Micropholcommatidae (Araneae), imaged in Eocene Paris amber using X-ray computed tomography. Zootaxa 1623: 47–53.

[R14] Ramírez, M. J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374.

[R08] Rix, M. G. 2008. A new species of Micropholcomma (Araneae: Araneoidea: Micropholcommatidae) from Western Australia. Records of the Western Australian Museum 24 (4): 343–348.

[S03] Schütt, K. 2003. Phylogeny of Symphytognathidae s.l. (Araneae, Araneoidea). Zoologica Scripta 32 (2): 129–151.

[T13] Taylor, C. K. 2013. Annotated bibliography for Barrow Island terrestrial invertebrates. Records of the Western Australian Museum Supplement 83: 135–144.

Amaurobioidea

Zorocrates aemulus, copyright Marshal Hedin.


Belongs within: Neocribellatae.
Contains: Stiphidiidae, Agelenidae, Amphinectidae, Desidae, Zodariidae, Amaurobiidae, Cycloctenidae, Pisauridae, Oxyopidae, Lycosidae, Dionycha.

The Amaurobioidea have historically been used to include cribellate spiders with unbranched median tracheae but such a grouping is now recognised as non-monophyletic. Instead, 'amaurobioids' represent basal members of what is commonly referred to as the RTA clade, referring to the presence of a retrolateral tibial apophysis on the male palp. Nested within this clade are the Lycosoidea, a clade of mostly actively hunting spiders united by a notably recurved posterior eyerow and grate-shaped tapeta in the anterior lateral and posterior median eyes (Ramírez 2014).

See also: Amaurobioidea: rummaging through a wastebasket.

Synapomorphies (from Ramírez 2014, as RTA clade): Carapace thoracic fovea a narrow dark longitudinal line; general metatarsal spination pattern x-x-x; more than two metatarsal trichobothria on a leg; tarsus with single trichobothrial row.

<==Amaurobioidea [Agelenoidea, Citigradae] R14
    |--+--Pimus Chamberlin 1947 R14, WZ08
    |  |    `--P. napa R14
    |  `--+--Stiphidiidae R14
    |     `--+--Agelenidae R14
    |        `--+--+--Amphinectidae R14
    |           |  `--Desidae R14
    |           `--+--Zodariidae JD-S07
    |              |--Amaurobiidae R14
    |              `--Cycloctenidae R14
    `--+--Lycosoidea R14
       |    |--Psechridae JD-S07
       |    |    |--Eomatachia latifrons Petrunkevitch 1942 S93
       |    |    |--Fecenia Simon 1887 JD-S07
       |    |    |    `--F. oblonga Rainbow 1913 R13
       |    |    `--Psechrus Thorell 1878 JD-S07 [incl. Lancaria Karsch 1879 K92]
       |    |         |--P. argentatus (Dol. 1857) (see below for synonymy) K92
       |    |         `--P. singaporensis JD-S07
       |    `--Lycosaeformia JD-S07
       |         |--+--Pisauridae R14
       |         |  `--+--Senoculus Taczanowski 1875 R14, JD-S07 [Senoculidae]
       |         |     `--Oxyopidae R14
       |         `--+--Lycosidae JD-S07
       |            `--Trechaleidae JD-S07
       |                 |--Syntrechalea brasilia Carico 2008 FM11
       |                 |--Enna JD-S07
       |                 `--Trechalea Thorell 1870 JD-S07
       |                      |--T. biocellata MC05
       |                      `--T. keyserlingi CM07
       `--+--Tengellidae R14
          |    |--Anachemmis Chamberlin 1919 JD-S07
          |    |--Calamistrula Dahl 1901 JD-S07
          |    |--Titiotus Simon 1897 JD-S07
          |    |--Austrotengella R14
          |    |--Anachemmis R14
          |    |--Socalchemmis Platnick & Ubick 2001 JD-S07
          |    |    `--S. idyllwild R14
          |    |--Tengella Dahl 1901 JD-S07
          |    |    `--T. radiata JD-S07
          |    `--Wiltona Koçak & Kemal 2008 PVD10 [=Haurokoa Forster & Wilton 1973 (preoc.) PVD10, JD-S07]
          |         `--*W. filicicola (Forster & Wilton 1973) [=*Haurokoa filicicola] PVD10
          `--+--*Ciniflella lutea R14
             `--+--Zorocratidae [Zorocratinae] BS09
                |    |--Zorodictyna Simon 1907 GC99, JD-S07
                |    |--Uduba Simon 1880 GC99, JD-S07
                |    |--Campostichomma Karsch 1891 GC99, JD-S07
                |    |    `--*C. manicatum Karsch 1892 K92
                |    `--Zorocrates Simon 1888 R14, JD-S07
                |         |--Z. aemulus R14
                |         |--Z. fuscus BS09
                |         |--Z. gnaphosoides R14
                |         `--Z. unicolor R14
                `--+--+--Dionycha R14
                   |  `--Liocranoides Keyserling 1881 R14, JD-S07
                   |       `--L. unicolor R14
                   `--+--Raecius Simon 1898 R14, JD-S07
                      |    `--R. jocquei JD-S07
                      `--Griswoldia Dippenaar-Schoeman & Jocqué 1997 R14, JD-S07 [Griswoldiinae]
                           |--G. acaenata R14
                           |--G. punctata R14
                           `--G. robusta R14

Amaurobioidea incertae sedis:
  Chumma Jocqué 2001 J01 [Chummidae JD-S07]
    |--*C. inquieta Jocqué 2001 J01
    `--C. gastroperforata Jocqué 2001 J01
  Phryganoporus candidus Do95
  Midgee Davies 1995 Da95
    |--*M. binnaburra Davies 1995 Da95
    |--M. alta Davies 1995 Da95
    |--M. bellendenker Davies 1995 Da95
    |--M. littlei Davies 1995 Da95
    |--M. minuta Davies 1995 Da95
    |--M. monteithi Davies 1995 Da95
    |--M. parva Davies 1995 Da95
    |--M. pumila Davies 1995 Da95
    `--M. thompsoni Davies 1995 Da95
  Manjala Da95
  Bakala Da95
  Malala Da95
  Insecutor [Insecutoridae] S93
    |--I. aculeatus Petrunkevitch 1942 S93
    |--I. mandibulatus Petrunkevitch 1942 S93
    `--I. rufus Petrunkevitch 1942 S93

Psechrus argentatus (Dol. 1857) [=Tegenaria argentata; incl. T. torva Cambr. 1870, Lancaria torva] K92

*Type species of generic name indicated

REFERENCES

[BS09] Blackledge, T. A., N. Scharff, J. A. Coddington, T. Szüts, J. W. Wenzel, C. Y. Hayashi & I. Agnarsson. 2009. Reconstructing web evolution and spider diversification in the molecular era. Proceedings of the National Academy of Sciences of the USA 106 (13): 5229–5234.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[Da95] Davies, V. T. 1995. A tiny litter spider (Araneae: Amaurobioidea) from Australian rainforests. Records of the Western Australian Museum Supplement 52: 119–129.

[Do95] Downes, M. F. 1995. Australasian social spiders: what is meant by 'social'? Records of the Western Australian Museum Supplement 52: 25–32.

[FM11] Freire-Jr, G. de B., & P. C. Motta. 2011. Effects of experimental fire regimes on the abundance and diversity of cursorial arachnids of Brazilian savannah (cerrado biome). Journal of Arachnology 39 (2): 263–272.

[GC99] Griswold, C. E., J. A. Coddington, N. I. Platnick & R. R. Forster. 1999. Towards a phylogeny of entelegyne spiders (Araneae, Araneomorphae, Entelegynae). Journal of Arachnology 27: 53–63.

[J01] Jocqué, R. 2001. Chummidae, a new spider family (Arachnida, Araneae) from South Africa. Journal of Zoology 254: 481–493.

[JD-S07] Jocqué, R., & A. S. Dippenaar-Schoeman. 2007. Spider Families of the World. Royal Museum for Central Africa: Tervuren (Belgium).

[K92] Karsch, F. 1892. Arachniden von Ceylon und von Minikoy gesammelt von den Herren Doctoren P. und F. Sarasin. Berliner Entomologische Zeitschrift 36 (2): 267–310.

[MC05] Machado, G., P. C. Carrera, A. M. Pomini & A. J. Marsaioli. 2005. Chemical defense in harvestmen (Arachnida, Opiliones): do benzoquinone secretions deter invertebrate and vertebrate predators? Journal of Chemical Ecology 31 (11): 2519–2539.

[PVD10] Paquin, P., C. J. Vink & N. Dupérré. 2010. Spiders of New Zealand: annotated family key and species list. Manaaki Whenua Press: Lincoln (New Zealand).

[R14] Ramírez, M. J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374.

[S93] Selden, P. A. 1993. Arthropoda (Aglaspidida, Pycnogonida and Chelicerata). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 297–320. Chapman & Hall: London.

[WZ08] Wang, X.-P., & M.-S. Zhu. 2008. Himalmartensus, a new genus of the spider family Amaurobiidae from Nepal (Araneae). Journal of Arachnology 36 (2): 241–250.