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Tangle-web spiders Temporal range: Cretaceous–present PreꞒ Ꞓ O S D C P T J K Pg N
Enoplognatha ovata
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Subphylum:	Chelicerata
Class:	Arachnida
Order:	Araneae
Infraorder:	Araneomorphae
Family:	Theridiidae Sundevall, 1833
Diversity
124 genera, 3028 species

Theridiidae, also known as the tangle-web spiders, cobweb spiders and comb-footed spiders, is a large family of araneomorph spiders first described by Carl Jakob Sundevall in 1833. This diverse, globally distributed family includes over 3,000 species in 124 genera, and is the most common arthropod found in human dwellings throughout the world.

Theridiid spiders are both entelegyne, meaning that the females have a genital plate, and ecribellate, meaning that they spin sticky capture silk instead of woolly silk. They have a comb of serrated bristles (setae) on the tarsus of the fourth leg.

The family includes some model organisms for research, including the medically important widow spiders. They are important to studies characterizing their venom and its clinical manifestation, but widow spiders are also used in research on spider silk and sexual biology, including sexual cannibalism. Anelosimus are also model organisms, used for the study of sociality, because it has evolved frequently within the genus, allowing comparative studies across species, and because it contains species varying from solitary to permanently social. These spiders are also a promising model for the study of inbreeding because all permanently social species are highly inbred.

The Hawaiian Theridion grallator is used as a model to understand the selective forces and the genetic basis of color polymorphism within species. T. grallator is known as the "happyface" spider, as certain morphs have a pattern resembling a smiley face or a grinning clown face on their yellow body.

Webs

They often build tangle space webs, hence the common name, but Theridiidae has a large diversity of spider web forms. Many trap ants and other ground dwelling insects using elastic, sticky silk trap lines leading to the soil surface. Webs remain in place for extended periods and are expanded and repaired, but no regular pattern of web replacement has been observed.

The well studied kleptoparasitic members of Argyrodinae (Argyrodes, Faiditus, and Neospintharus) live in the webs of larger spiders and pilfer small prey caught by their host's web. They eat prey killed by the host spider, consume silk from the host web, and sometimes attack and eat the host itself.

Theridiid gumfoot-webs consist of frame lines that anchor them to surroundings and of support threads, which possess viscid silk. These can either have a central retreat (Achaearanea-type) or a peripheral retreat (Latrodectus-type). Building gum-foot lines is a unique, stereotyped behaviour, and is likely homologous for Theridiidae and its sister family Nesticidae.

Among webs without gumfooted lines, some contain viscid silk (Theridion-type) and some that are sheet-like, which do not contain viscid silk (Coleosoma-type). However, there are many undescribed web forms.

Genera

The largest genus is Theridion with over 600 species, but it is not monophyletic. Parasteatoda, previously Achaearanea, is another large genus that includes the North American common house spider. As of April 2019, the World Spider Catalog accepts the following genera:

• Achaearanea Strand, 1929 – Africa, Asia, Australia, South America, Central America
• Achaearyopa Barrion & Litsinger, 1995 – Philippines
• Achaeridion Wunderlich, 2008 – Turkey
• Allothymoites Ono, 2007 – China, Japan
• Ameridion Wunderlich, 1995 – Central America, Caribbean, Mexico, South America
• Anatea Berland, 1927 – Australia
• Anatolidion Wunderlich, 2008 – Africa, Europe, Turkey
• Anelosimus Simon, 1891 – Asia, Africa, North America, South America, Oceania, Central America, Caribbean
• Argyrodella Saaristo, 2006 – Seychelles
• Argyrodes Simon, 1864 – Africa, Asia, Oceania, North America, South America, Jamaica
• Ariamnes Thorell, 1869 – Costa Rica, South America, Asia, Africa, Oceania, Mexico, Cuba
• Asagena Sundevall, 1833 – North America, Asia, Europe, Algeria
• Asygyna Agnarsson, 2006 – Madagascar
• Audifia Keyserling, 1884 – Guinea-Bissau, Congo, Brazil
• Bardala Saaristo, 2006 – Seychelles
• Borneoridion Deeleman & Wunderlich, 2011 – Indonesia
• Brunepisinus Yoshida & Koh, 2011 – Indonesia
• Cabello Levi, 1964 – Venezuela
• Cameronidion Wunderlich, 2011 – Malaysia
• Campanicola Yoshida, 2015 – Asia
• Canalidion Wunderlich, 2008 – Russia
• Carniella Thaler & Steinberger, 1988 – Europe, Angola, Asia
• Cephalobares O. Pickard-Cambridge, 1871 – Sri Lanka, China
• Cerocida Simon, 1894 – Brazil, Venezuela, Guyana
• Chikunia Yoshida, 2009 – Asia
• Chorizopella Lawrence, 1947 – South Africa
• Chrosiothes Simon, 1894 – North America, South America, Central America, Caribbean, Asia
• Chrysso O. Pickard-Cambridge, 1882 – North America, South America, Central America, Asia, Trinidad, Europe
• Coleosoma O. Pickard-Cambridge, 1882 – United States, South America, Seychelles, Asia, New Zealand
• Coscinida Simon, 1895 – Asia, Africa
• Craspedisia Simon, 1894 – Brazil
• Crustulina Menge, 1868 – Ukraine, United States, Africa, Oceania, Asia
• Cryptachaea Archer, 1946 – South America, North America, Oceania, Central America, Asia, Trinidad, Belgium
• Cyllognatha L. Koch, 1872 – Samoa, Australia, India
• Deelemanella Yoshida, 2003 – Indonesia
• Dipoena Thorell, 1869 – North America, Oceania, Asia, Central America, South America, Caribbean, Africa, Europe
• Dipoenata Wunderlich, 1988 – Panama, South America, Malta
• Dipoenura Simon, 1909 – Asia, Sierra Leone
• Echinotheridion Levi, 1963 – South America
• Emertonella Bryant, 1945 – North America, Asia, Papua New Guinea
• Enoplognatha Pavesi, 1880 – Asia, Europe, Australia, Africa, North America, South America
• Episinus Walckenaer, 1809 – Asia, South America, Europe, North America, New Zealand, Central America, Africa, Caribbean
• Euryopis Menge, 1868 – Asia, North America, South America, Jamaica, Europe, Oceania, Africa, Panama
• Eurypoena Wunderlich, 1992 – Canary Is.
• Exalbidion Wunderlich, 1995 – Central America, South America, Mexico
• Faiditus Keyserling, 1884 – South America, North America, Central America, Caribbean, Asia
• Gmogala Keyserling, 1890 – Papua New Guinea, Australia
• Grancanaridion Wunderlich, 2011 – Canary Is.
• Guaraniella Baert, 1984 – Brazil, Paraguay
• Hadrotarsus Thorell, 1881 – Oceania, Belgium, Taiwan
• Helvibis Keyserling, 1884 – South America, Panama, Trinidad
• Helvidia Thorell, 1890 – Indonesia
• Hentziectypus Archer, 1946 – Caribbean, Panama, North America, South America
• Heterotheridion Wunderlich, 2008 – Turkey, Russia, China
• Hetschkia Keyserling, 1886 – Brazil
• Histagonia Simon, 1895 – South Africa
• Icona Forster, 1955 – New Zealand
• Jamaitidion Wunderlich, 1995 – Jamaica
• Janula Strand, 1932 – Asia, South America, Australia, Panama, Trinidad
• Keijiella Yoshida, 2016 – Asia
• Kochiura Archer, 1950 – Chile, Turkey, Brazil
• Landoppo Barrion & Litsinger, 1995 – Philippines
• Lasaeola Simon, 1881 – Europe, North America, Panama, South America, Asia
• Latrodectus Walckenaer, 1805 – South America, North America, Asia, Europe, Oceania, Africa
• Macaridion Wunderlich, 1992 – Europe
• Magnopholcomma Wunderlich, 2008 – Australia
• Meotipa Simon, 1894 – Asia, Papua New Guinea
• Molione Thorell, 1892 – Asia
• Moneta O. Pickard-Cambridge, 1871 – Oceania, Asia, Seychelles
• Montanidion Wunderlich, 2011 – Malaysia
• Nanume Saaristo, 2006 – Seychelles
• Neopisinus Marques, Buckup & Rodrigues, 2011 – Panama, Caribbean, South America, North America
• Neospintharus Exline, 1950 – North America, Asia, South America, Central America
• Neottiura Menge, 1868 – Asia, Europe, Algeria
• Nesopholcomma Ono, 2010 – Japan
• Nesticodes Archer, 1950 – Asia, New Zealand
• Nihonhimea Yoshida, 2016 – Asia, Seychelles, Oceania, Mexico
• Nipponidion Yoshida, 2001 – Japan
• Nojimaia Yoshida, 2009 – China, Japan
• Ohlertidion Wunderlich, 2008 – Greenland, Russia
• Okumaella Yoshida, 2009 – Japan
• Paidiscura Archer, 1950 – Europe, Algeria, Asia
• Parasteatoda Archer, 1946 – Asia, Oceania, Cuba, North America, Argentina, Seychelles
• Paratheridula Levi, 1957 – United States, Chile
• Pholcomma Thorell, 1869 – Oceania, North America, Asia, South America
• Phoroncidia Westwood, 1835 – Asia, Africa, North America, Caribbean, South America, Oceania, Europe, Costa Rica
• Phycosoma O. Pickard-Cambridge, 1879 – North America, Asia, Africa, Jamaica, Panama, Brazil, New Zealand
• Phylloneta Archer, 1950 – Asia, United States, Spain
• Platnickina Koçak & Kemal, 2008 – North America, Asia, Africa
• Proboscidula Miller, 1970 – Angola, Rwanda
• Propostira Simon, 1894 – India, Sri Lanka
• Pycnoepisinus Wunderlich, 2008 – Kenya
• Rhomphaea L. Koch, 1872 – Asia, Africa, South America, Oceania, North America, Europe, Central America, Saint Vincent and the Grenadines
• Robertus O. Pickard-Cambridge, 1879 – Europe, North America, Asia, Congo
• Ruborridion Wunderlich, 2011 – India
• Rugathodes Archer, 1950 – Asia, North America
• Sardinidion Wunderlich, 1995 – Africa, Europe
• Selkirkiella Berland, 1924 – Chile, Argentina
• Sesato Saaristo, 2006 – Seychelles
• Seycellesa Koçak & Kemal, 2008 – Seychelles
• Simitidion Wunderlich, 1992 – Europe, Asia, Canada
• Spheropistha Yaginuma, 1957 – Japan, China
• Spinembolia Saaristo, 2006 – Seychelles
• Spintharus Hentz, 1850 – Pakistan, Caribbean, Mexico, Brazil
• Steatoda Sundevall, 1833 – Oceania, North America, Asia, Europe, South America, Africa
• Stemmops O. Pickard-Cambridge, 1894 – South America, North America, Central America, Caribbean, Asia
• Stoda Saaristo, 2006 – Seychelles
• Styposis Simon, 1894 – United States, South America, Central America, Congo
• Takayus Yoshida, 2001 – Asia
• Tamanidion Wunderlich, 2011 – Malaysia
• Tekellina Levi, 1957 – United States, Brazil, Asia
• Theonoe Simon, 1881 – Tanzania, Europe, North America
• Theridion Walckenaer, 1805 – Asia, North America, Central America, Europe, South America, Africa, Oceania, Caribbean
• Theridula Emerton, 1882 – Spain, Africa, North America, Central America, Asia, South America
• Thwaitesia O. Pickard-Cambridge, 1881 – Panama, South America, Africa, Asia, Oceania, Trinidad
• Thymoites Keyserling, 1884 – South America, Central America, Asia, North America, Caribbean, Greenland, Tanzania
• Tidarren Chamberlin & Ivie, 1934 – Africa, Yemen, North America, Argentina, Costa Rica
• Tomoxena Simon, 1895 – Indonesia, India
• Wamba O. Pickard-Cambridge, 1896 – North America, South America, Panama
• Wirada Keyserling, 1886 – Mexico, South America
• Yaginumena Yoshida, 2002 – Asia
• Yoroa Baert, 1984 – Papua New Guinea, Australia
• Yunohamella Yoshida, 2007 – Asia, Europe
• Zercidium Benoit, 1977 – St. Helena

About 35 extinct genera have also been placed in the family. The oldest known stem-group member of the family is Cretotheridion from the Cenomanian aged Burmese amber of Myanmar.

See also

• List of Theridiidae species

References

1. Sundevall, C. J. (1833). Conspectus Arachnidum.
2. ^ "Family: Theridiidae Sundevall, 1833". World Spider Catalog. Natural History Museum Bern. Retrieved 2019-04-25.
3. Leong, Misha; et al. (2017). "The Habitats Humans Provide: Factors affecting the diversity and composition of arthropods in houses". Scientific Reports. 7 (15347): 15347. Bibcode:2017NatSR...715347L. doi:10.1038/s41598-017-15584-2. PMC 5681556. PMID 29127355.
4. Agnarsson, I. (2006). "Asymmetric female genitalia and other remarkable morphology in a new genus of cobweb spiders (Theridiidae, Araneae) from Madagascar". Biological Journal of the Linnean Society. 87 (2): 211–232. doi:10.1111/j.1095-8312.2006.00569.x.
5. Purcell, J.; Aviles, L. (2007). "Smaller colonies and more solitary living mark higher elevation populations of a social spider". Journal of Animal Ecology. 76 (3): 590–597. Bibcode:2007JAnEc..76..590P. doi:10.1111/j.1365-2656.2007.01228.x. PMID 17439475.
6. Agnarsson, I. (2006). "A revision of the New World eximius lineage of Anelosimus (Araneae, Theridiidae) and a phylogenetic analysis using worldwide exemplars" (PDF). Zoological Journal of the Linnean Society. 146 (4): 453–593. doi:10.1111/j.1096-3642.2006.00213.x. Archived from the original (PDF) on 2007-09-27.
7. Oxford, G.S.; Gillespie, R.G. (1996). "Quantum Shifts in the Genetic Control of a Colour Polymorphism in Theridion Grallator (Araneae: Theridiidae), the Hawaiian Happy-Face Spider". Heredity. 76 (3): 249–256. doi:10.1038/hdy.1996.38. S2CID 824014.
8. Gillespie, R.G.; Tabashnik, B.E. (1989). "What makes a happy face? Determinants of color pattern in the Hawaiian happy face spider Theridion grallator (Araneae, Theridiidae)". Heredity. 62 (3): 355–364. doi:10.1038/hdy.1989.50.
9. Benjamin, S.P.; Zschokke, S. (2003). "Webs of theridiid spiders: construction, structure and evolution". Biological Journal of the Linnean Society. 78 (3): 293–305. doi:10.1046/j.1095-8312.2003.00110.x.
10. Benjamin, Suresh P.; Zschokke, Samuel (2002). "Untangling the tangle-web: web building behaviour of the comb-footed spider Steatoda triangulosa and comments on phylogenetic implications (Araneae: Theridiidae)". Journal of Insect Behavior. 15 (6): 791–809. doi:10.1023/A:1021175507377. S2CID 26850827.
11. Vollrath, F. (1979). "Behavior of the Kleptoparasitic Spider Argyrodes-Elevatus (Araneae, Theridiidae)". Animal Behaviour. 27: 515–521. doi:10.1016/0003-3472(79)90186-6. S2CID 53177663.
12. Grostal, P.; Walter, D.E. (1997). "Kleptoparasites or commensals? Effects of Argyrodes antipodianus (Araneae: Theridiidae) on nephila plumipes (Araneae: Tetragnathidae)". Oecologia. 111 (4): 570–574. Bibcode:1997Oecol.111..570G. doi:10.1007/s004420050273. PMID 28308120. S2CID 11217593.
13. Blackledge, T.A.; Swindeman, J.E.; Hayashi, C.Y. (2005). "Quasistatic and continuous dynamic characterization of the mechanical properties of silk from the cobweb of the black widow spider Latrodectus hesperus". Journal of Experimental Biology. 208 (10): 1937–1949. doi:10.1242/jeb.01597. PMID 15879074.
14. Blackledge, T.A.; Zevenbergen, J.M. (2007). "Condition dependent spider web architecture in the western black widow Latrodectus hesperus". Animal Behaviour. 73 (5): 855–864. doi:10.1016/j.anbehav.2006.10.014. S2CID 53182880.
15. Benjamin, Suresh P.; Zschokke, Samuel (2003). "Webs of theridiid spiders: construction, structure and evolution". Biological Journal of the Linnean Society. 78 (3): 293–305. doi:10.1046/j.1095-8312.2003.00110.x.
16. Dunlop, J.A.; Penney, D.; Jekel, D. (2015). "A summary list of fossil spiders and their relatives" (PDF). World Spider Catalog. Natural History Museum Bern. Retrieved 2015-11-06.
17. Magalhaes, Ivan L. F.; Azevedo, Guilherme H. F.; Michalik, Peter; Ramírez, Martín J. (February 2020). "The fossil record of spiders revisited: implications for calibrating trees and evidence for a major faunal turnover since the Mesozoic". Biological Reviews. 95 (1): 184–217. doi:10.1111/brv.12559. ISSN 1464-7931. PMID 31713947. S2CID 207937170.

Further reading

• Agnarsson I. 2006c. Phylogenetic placement of Echinotheridion (Araneae: Theridiidae) – do male sexual organ removal, emasculation, and sexual cannibalism in Echinotheridion and Tidarren represent evolutionary replicas? Invertebrate Systematics 20: 415–429. PDF Archived 2007-09-27 at the Wayback Machine
• Agnarsson I. 2004. Morphological phylogeny of cobweb spiders and their relatives (Araneae, Araneoidea, Theridiidae). Zoological Journal of the Linnean Society 141: 447–626. PDF Archived 2007-01-01 at the Wayback Machine
• Arnedo, M.A., Coddington, J., Agnarsson, I. & Gillespie, R.G. (2004). From a comb to a tree: phylogenetic relationships of the comb-footed spiders (Araneae, Theridiidae) inferred from nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 31:225–245. PDF
• Arnedo MA, Agnarsson I, Gillespie RG. In Press. Molecular insights into the phylogenetic structure of the spider genus Theridion (Araneae, Theridiidae) and the origin of the Hawaiian Theridion-like fauna. Zoologica Scripta.
• Aviles, L., Maddison, W.P. and Agnarsson, I. 2006. A new independently derived social spider with explosive colony proliferation and a female size dimorphism. Biotropica, 38: 743–753.
• Gillespie, R.G. and Tabashnik, B.E. 1994. Foraging Behavior of the Hawaiian Happy Face Spider (Araneae, Theridiidae). Annals of the Entomological Society of America, 87: 815–822.
• Oxford, G.S. and Gillespie, R.G. 1996. Genetics of a colour polymorphism in Theridion grallator (Araneae: Theridiidae), the Hawaiian happy-face spider, from greater Maui. Heredity, 76: 238–248.

External links

• Platnick, N.I. 2006. World Spider Catalog
• Key to several Theridiidae genera
• Tree of Life: Theridiidae
• Theridiidae research

• Theridiidae
• Extant Cretaceous first appearances
• Araneomorphae families