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== Spider sociality ==
== Spider sociality ==
Most species of social spiders live in the tropical regions of the world where insect size and density is highest but several species reach into the eastern United States and other temperate areas. By building a communal web, it is thought that the spiders approximately maximize total biomass capture per spider.<ref>{{Cite journal | url=http://www.pnas.org/content/early/2008/08/07/0710603105.abstract | title=Cooperative capture of large prey solves scaling challenge faced by spider societies | journal=By Eric C. Yip, Kimberly S. Powers, and Leticia Avilés | doi=10.1073/pnas.0710603105 | volume=105 | pages=11818–11822| pmc=2575263 | pmid=18689677 | year=2008 | author=Yip EC, Powers KS, Avilés L| issue=33 }}</ref> Having a larger web and multiple spiders to work together to subdue prey allows them to prey on larger organisms than would be possible if they led a solitary existence. The colonies can grow large enough to take down birds and bats, as well as very large insects.
Most species of social spiders live in the tropical regions of the world where insect size and density is highest but several species reach into the eastern United States and other temperate areas. By building a communal web, it is thought that the spiders approximately maximize total biomass capture per spider.<ref>{{Cite journal | url=http://www.pnas.org/content/early/2008/08/07/0710603105.abstract | title=Cooperative capture of large prey solves scaling challenge faced by spider societies | journal= Proceedings of the National Academy of Sciences| doi=10.1073/pnas.0710603105 | volume=105 | pages=11818–11822| pmc=2575263 | pmid=18689677 | year=2008 | author=Yip EC, Powers KS, Avilés L| issue=33 | bibcode=2008PNAS..10511818Y }}</ref> Having a larger web and multiple spiders to work together to subdue prey allows them to prey on larger organisms than would be possible if they led a solitary existence. The colonies can grow large enough to take down birds and bats, as well as very large insects.


Living in a colony also has another major benefit for spiders: cooperative nest maintenance. Nest maintenance does not rely solely on an individual in a colony setting and thus saves on a per-capita investment in maintaining silk structures. Predator defense is also increased in a colony with a large web and multiple individuals analogous to schools of fish or herds of mammals.<ref>{{Cite web|url=http://www.americanarachnology.org/JoA_free/JoA_v38_n3/arac-38-03-446.pdf|title=Social organization of the colonial spider Leucauge sp in the Neotropics vertical stratification within colonies}}</ref>
Living in a colony also has another major benefit for spiders: cooperative nest maintenance. Nest maintenance does not rely solely on an individual in a colony setting and thus saves on a per-capita investment in maintaining silk structures. Predator defense is also increased in a colony with a large web and multiple individuals analogous to schools of fish or herds of mammals.<ref>{{Cite web|url=http://www.americanarachnology.org/JoA_free/JoA_v38_n3/arac-38-03-446.pdf|title=Social organization of the colonial spider Leucauge sp in the Neotropics vertical stratification within colonies}}</ref>
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===Levels===
===Levels===
[[File:2007-social-spider-1.jpg|thumb|Social spiders from [[Bengaluru]], [[India]]]]
[[File:2007-social-spider-1.jpg|thumb|Social spiders from [[Bengaluru]], [[India]]]]
Social spiders exhibit varying levels of [[Sociality|sociality]], of which there are six defined. Agnarsson ''et al''. estimate that spiders as a whole have independently evolved sociality 18 or 19 times.<ref name= "Agnarsson">{{cite journal |author1=Ingi Agnarsson |author2=Leticia Aviles |author3=Jonathan A. Coddington |author4=Wayne P. Maddison |name-list-style=amp | title= Sociality In Theridiid Sspiders: Repeated Origins Of An Evolutionary Dead End | journal=Evolution | volume=60 | issue=11 | year=2006 | pages=2342–51 | pmid= 17236425 | doi=10.1554/06-078.1}}</ref> Most of these social spiders broadly fit into the quasi-social definition of sociality, meaning they show cooperative brood care, use the same nest (web), and have some amount of generational overlap. Several permutations of social behavior exist amongst the 23 species of spider considered to be quasi-social out of some 45,000 known species of spider.<ref>{{cite web|title=World Spider Catalogue|url=http://www.wsc.nmbe.ch/|publisher=NMBE|access-date=31 October 2015}}</ref> These 23 species are [[Phylogenetics|phylogenetically]] scattered in 11 genera across eight widely separated families.<ref name="Agnarsson"/>
Social spiders exhibit varying levels of [[Sociality|sociality]], of which there are six defined. Agnarsson ''et al''. estimate that spiders as a whole have independently evolved sociality 18 or 19 times.<ref name= "Agnarsson">{{cite journal |author1=Ingi Agnarsson |author2=Leticia Aviles |author3=Jonathan A. Coddington |author4=Wayne P. Maddison |name-list-style=amp | title= Sociality In Theridiid Sspiders: Repeated Origins Of An Evolutionary Dead End | journal=Evolution | volume=60 | issue=11 | year=2006 | pages=2342–51 | pmid= 17236425 | doi=10.1554/06-078.1|s2cid=16995686 }}</ref> Most of these social spiders broadly fit into the quasi-social definition of sociality, meaning they show cooperative brood care, use the same nest (web), and have some amount of generational overlap. Several permutations of social behavior exist amongst the 23 species of spider considered to be quasi-social out of some 45,000 known species of spider.<ref>{{cite web|title=World Spider Catalogue|url=http://www.wsc.nmbe.ch/|publisher=NMBE|access-date=31 October 2015}}</ref> These 23 species are [[Phylogenetics|phylogenetically]] scattered in 11 genera across eight widely separated families.<ref name="Agnarsson"/>


The level of sociality often varies between species (interspecies) but can vary within a species (intraspecies) as well. Intraspecific variation is generally habitat dependant, where some populations within a species show all the characteristics of quasi-sociality, yet a population a mile away may be largely solitary<ref name="Lubin">{{cite journal |author1=Lubin, Yael D. |author2=Robinson Michael H. |name-list-style=amp | title= Dispersal by Swarming in a Social Spider | journal=Science | volume=216 | issue=4543 | year=1982 | pages=319–21 | doi= 10.1126/science.216.4543.319 | pmid= 17832747 }}</ref> because they inhabit a different environment. This facultative sociality allows them to survive periods of sub-optimal conditions, when sustaining large aggregations is not feasible. Some of these aggregations can contain as many as 50,000 individuals as in the case of ''[[Anelosimus eximius]]'' (in the [[family (biology)|family]] [[Theridiidae]]).<ref>{{cite journal | author=Vollrath, F. | year=1986 | title=Eusociality and extraordinary sex ratios in the spider Anelosimus eximius (Araneae: Theridiidae) | journal=Behavioral Ecology and Sociobiology | volume=18 | issue=4 | pages=283–287 | doi=10.1007/BF00300005 }}</ref> The genus ''Anelosimus'' has a strong tendency towards sociality: all known American species are social, and species in [[Madagascar]] are at least somewhat social.<ref>{{cite journal |author1=Agnarsson, I. |author2=Kuntner, M. |name-list-style=amp | title=Madagascar: an unexpected hotspot of social Anelosimus spider diversity (Araneae: Theridiidae) | journal=Systematic Entomology | volume=30 | issue=4| pages=575–592 |date=April 2005 | doi=10.1111/j.1365-3113.2005.00289.x}}</ref>
The level of sociality often varies between species (interspecies) but can vary within a species (intraspecies) as well. Intraspecific variation is generally habitat dependant, where some populations within a species show all the characteristics of quasi-sociality, yet a population a mile away may be largely solitary<ref name="Lubin">{{cite journal |author1=Lubin, Yael D. |author2=Robinson Michael H. |name-list-style=amp | title= Dispersal by Swarming in a Social Spider | journal=Science | volume=216 | issue=4543 | year=1982 | pages=319–21 | doi= 10.1126/science.216.4543.319 | pmid= 17832747 |bibcode=1982Sci...216..319L |s2cid=32470345 }}</ref> because they inhabit a different environment. This facultative sociality allows them to survive periods of sub-optimal conditions, when sustaining large aggregations is not feasible. Some of these aggregations can contain as many as 50,000 individuals as in the case of ''[[Anelosimus eximius]]'' (in the [[family (biology)|family]] [[Theridiidae]]).<ref>{{cite journal | author=Vollrath, F. | year=1986 | title=Eusociality and extraordinary sex ratios in the spider Anelosimus eximius (Araneae: Theridiidae) | journal=Behavioral Ecology and Sociobiology | volume=18 | issue=4 | pages=283–287 | doi=10.1007/BF00300005 | s2cid=44727810 }}</ref> The genus ''Anelosimus'' has a strong tendency towards sociality: all known American species are social, and species in [[Madagascar]] are at least somewhat social.<ref>{{cite journal |author1=Agnarsson, I. |author2=Kuntner, M. |name-list-style=amp | title=Madagascar: an unexpected hotspot of social Anelosimus spider diversity (Araneae: Theridiidae) | journal=Systematic Entomology | volume=30 | issue=4| pages=575–592 |date=April 2005 | doi=10.1111/j.1365-3113.2005.00289.x|s2cid=13871079 }}</ref>


Members of other species in the same family but several different genera have [[convergent evolution|independently]] developed social behavior. For example, although ''[[Theridion nigroannulatum]]'' belongs to a [[genus]] with no other social species, ''T. nigroannulatum'' build colonies that may contain several thousand individuals that co-operate in prey capture and share food.<ref>{{cite journal | author=Avilés, L., Maddison, W.P. and Agnarsson, I.| title=A New Independently Derived Social Spider with Explosive Colony Proliferation and a Female Size Dimorphism | journal=Biotropica | volume=38 | issue=6 | pages=743–753 |date=August 2006 | doi=10.1111/j.1744-7429.2006.00202.x }}</ref> Other communal spiders include several ''[[Philoponella]]'' species (family [[Uloboridae]]), ''[[Agelena consociata]]'' (family [[Agelenidae]]) and ''[[Mallos gregalis]]'' (family [[Dictynidae]]).<ref>{{cite journal | author=Matsumoto, T. | year=1998 | title=Cooperative prey capture in the communal web spider, ''Philoponella raffray'' (Araneae, Uloboridae)| journal=Journal of Arachnology | volume=26 | pages=392–396 | url=http://www.americanarachnology.org/JoA_free/JoA_v26_n3/JoA_v26_p392.pdf | access-date=2008-10-11 }}</ref> Social predatory spiders need to defend their prey against [[kleptoparasite]]s ("thieves"), and larger colonies are more successful in this.<ref>{{cite journal | author=Cangialosi, K.R. | title=Social spider defense against kleptoparasitism| journal=Behavioral Ecology and Sociobiology | volume=27 | issue=1 |date=July 1990 | doi=10.1007/BF00183313 }}</ref> The [[Herbivore|herbivorous]] spider ''[[Bagheera kiplingi]]'' lives in small colonies which helps to protect eggs and spiderlings.<ref>{{Cite conference | author=Meehan, C,J. Olson, E.J. and Curry, R.L. | date=21 August 2008 | title=Exploitation of the Pseudomyrmex–Acacia mutualism by a predominantly vegetarian jumping spider (''Bagheera kiplingi'') | conference=93rd ESA Annual Meeting | url=http://eco.confex.com/eco/2008/techprogram/P12401.HTM | access-date=2008-10-10 }}</ref> Even [[widow spider]]s (genus ''[[Latrodectus]]''), which are notoriously aggressive and cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.<ref>{{cite journal | author=Bertani, R., Fukushima, C.S., and Martins, R.| title=Sociable widow spiders? Evidence of subsociality in '' Latrodectus''Walckenaer, 1805 (Araneae, Theridiidae) | journal=Journal of Ethology | volume=26 | issue=2 |date=May 2008 | doi=10.1007/s10164-007-0082-8 | pages=299–302 }}</ref>
Members of other species in the same family but several different genera have [[convergent evolution|independently]] developed social behavior. For example, although ''[[Theridion nigroannulatum]]'' belongs to a [[genus]] with no other social species, ''T. nigroannulatum'' build colonies that may contain several thousand individuals that co-operate in prey capture and share food.<ref>{{cite journal | author=Avilés, L., Maddison, W.P. and Agnarsson, I.| title=A New Independently Derived Social Spider with Explosive Colony Proliferation and a Female Size Dimorphism | journal=Biotropica | volume=38 | issue=6 | pages=743–753 |date=August 2006 | doi=10.1111/j.1744-7429.2006.00202.x }}</ref> Other communal spiders include several ''[[Philoponella]]'' species (family [[Uloboridae]]), ''[[Agelena consociata]]'' (family [[Agelenidae]]) and ''[[Mallos gregalis]]'' (family [[Dictynidae]]).<ref>{{cite journal | author=Matsumoto, T. | year=1998 | title=Cooperative prey capture in the communal web spider, ''Philoponella raffray'' (Araneae, Uloboridae)| journal=Journal of Arachnology | volume=26 | pages=392–396 | url=http://www.americanarachnology.org/JoA_free/JoA_v26_n3/JoA_v26_p392.pdf | access-date=2008-10-11 }}</ref> Social predatory spiders need to defend their prey against [[kleptoparasite]]s ("thieves"), and larger colonies are more successful in this.<ref>{{cite journal | author=Cangialosi, K.R. | title=Social spider defense against kleptoparasitism| journal=Behavioral Ecology and Sociobiology | volume=27 | issue=1 |date=July 1990 | doi=10.1007/BF00183313 | s2cid=38307165}}</ref> The [[Herbivore|herbivorous]] spider ''[[Bagheera kiplingi]]'' lives in small colonies which helps to protect eggs and spiderlings.<ref>{{Cite conference | author=Meehan, C,J. Olson, E.J. and Curry, R.L. | date=21 August 2008 | title=Exploitation of the Pseudomyrmex–Acacia mutualism by a predominantly vegetarian jumping spider (''Bagheera kiplingi'') | conference=93rd ESA Annual Meeting | url=http://eco.confex.com/eco/2008/techprogram/P12401.HTM | access-date=2008-10-10 }}</ref> Even [[widow spider]]s (genus ''[[Latrodectus]]''), which are notoriously aggressive and cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.<ref>{{cite journal | author=Bertani, R., Fukushima, C.S., and Martins, R.| title=Sociable widow spiders? Evidence of subsociality in '' Latrodectus''Walckenaer, 1805 (Araneae, Theridiidae) | journal=Journal of Ethology | volume=26 | issue=2 |date=May 2008 | doi=10.1007/s10164-007-0082-8 | pages=299–302 | s2cid=36475912 }}</ref>


A few species, such as ''[[Anelosimus eximius]]'', are also known to have reproductive skew, where some of the females reproduce and others do not.<ref>{{cite journal | author= Ann Lundie Rypstra | title= Prey Size, Social Competition, and the Development of Reproductive Division of Labor in Social Spider Groups | journal=The American Naturalist | volume=142 | year=1993 | pages=868–880 | issue=5 | doi= 10.1086/285577 }}</ref> Even though these spiders cooperate by caring for each other's young, cooperating in prey capture and sharing food, they are not considered [[eusocial]] as they do not have defined castes. Any female has the potential to reproduce, therefore differences in the reproductive success is due to competition for scarce food resources rather than the existence of physically distinct castes.<ref name="Ingi anelosims phylogentic">{{cite journal|last=Agnarsson|first=Ingi|title=A revision of the New World eximius lineage of Anelosimus (Araneae, Theridiidae) and a phylogenetic analysis using worldwide exemplars|journal=Zoological Journal of the Linnean Society|year=2006|volume=146|issue=4|pages=453–593|doi=10.1111/j.1096-3642.2006.00213.x|doi-access=free}}</ref>
A few species, such as ''[[Anelosimus eximius]]'', are also known to have reproductive skew, where some of the females reproduce and others do not.<ref>{{cite journal | author= Ann Lundie Rypstra | title= Prey Size, Social Competition, and the Development of Reproductive Division of Labor in Social Spider Groups | journal=The American Naturalist | volume=142 | year=1993 | pages=868–880 | issue=5 | doi= 10.1086/285577 | s2cid= 85144990 }}</ref> Even though these spiders cooperate by caring for each other's young, cooperating in prey capture and sharing food, they are not considered [[eusocial]] as they do not have defined castes. Any female has the potential to reproduce, therefore differences in the reproductive success is due to competition for scarce food resources rather than the existence of physically distinct castes.<ref name="Ingi anelosims phylogentic">{{cite journal|last=Agnarsson|first=Ingi|title=A revision of the New World eximius lineage of Anelosimus (Araneae, Theridiidae) and a phylogenetic analysis using worldwide exemplars|journal=Zoological Journal of the Linnean Society|year=2006|volume=146|issue=4|pages=453–593|doi=10.1111/j.1096-3642.2006.00213.x|doi-access=free}}</ref>


===Sub-social===
===Sub-social===
Many more species of spider are considered to be sub-social than quasi-social – meaning they lack fixed or complex social organization. These species may only display social behaviors as a seasonal venture and have an obligate solitary phase. Some other species will establish territories within the colony and can even have discrete webs, narrowly connected to other webs within the colony (such is the case with ''Leucauge spp.''<ref>http://www.americanarachnology.org/JoA_free/JoA_v38_n3/arac-38-03-446.pdf</ref>). This is not a fully cooperative behavior as there is little to no cooperative nest maintenance or brood care occurring. The subsocial species, however, appear to be crucial for the evolution of sociality in spiders. Recent theory suggests that social spiders evolved along a restricted pathway through solitary subsocial ancestors.<ref name=Agnarsson /><ref>{{cite journal |author1=Johannesen J. |author2=Wickler W. |author3=Seibt U. |author4=Moritz R. F. A. | year = 2009 | title = Population history in social spiders repeated: Colony structure and lineage evolution in stegodyphus mimosarum (eresidae) | journal = Molecular Ecology | volume = 18 | issue = 13| pages = 2812–2818 | doi=10.1111/j.1365-294x.2009.04238.x | pmid=19500247}}</ref> It has been shown that the subsocial spiders of genus ''[[Stegodyphus]]'' tolerate a low level of inbreeding with low inbreeding depression, suggesting a possible stepping stone towards the fully inbred mating system found in social spiders.<ref>{{cite journal |author1=Bilde T. |author2=Lubin Y. |author3=Smith D. |author4=Schneider J. M. |author5=Maklakov A. A. | year = 2005 | title = The transition to social inbred mating systems in spiders: Role of inbreeding tolerance in a subsocial predecessor | journal = Evolution | volume = 59 | pages = 160–174 | doi=10.1554/04-361 | pmid = 15792236}}</ref><ref>{{cite journal |author1=Ruch J. |author2=Heinrich L. |author3=Bilde T. |author4=Schneider J. M. | year = 2009 | title = The evolution of social inbreeding mating systems in spiders: Limited male mating dispersal and lack of pre-copulatory inbreeding avoidance in a subsocial predecessor | journal = Biological Journal of the Linnean Society | volume = 98 | issue = 4| pages = 851–859 | doi=10.1111/j.1095-8312.2009.01322.x| doi-access = free }}</ref>
Many more species of spider are considered to be sub-social than quasi-social – meaning they lack fixed or complex social organization. These species may only display social behaviors as a seasonal venture and have an obligate solitary phase. Some other species will establish territories within the colony and can even have discrete webs, narrowly connected to other webs within the colony (such is the case with ''Leucauge spp.''<ref>http://www.americanarachnology.org/JoA_free/JoA_v38_n3/arac-38-03-446.pdf</ref>). This is not a fully cooperative behavior as there is little to no cooperative nest maintenance or brood care occurring. The subsocial species, however, appear to be crucial for the evolution of sociality in spiders. Recent theory suggests that social spiders evolved along a restricted pathway through solitary subsocial ancestors.<ref name=Agnarsson /><ref>{{cite journal |author1=Johannesen J. |author2=Wickler W. |author3=Seibt U. |author4=Moritz R. F. A. | year = 2009 | title = Population history in social spiders repeated: Colony structure and lineage evolution in stegodyphus mimosarum (eresidae) | journal = Molecular Ecology | volume = 18 | issue = 13| pages = 2812–2818 | doi=10.1111/j.1365-294x.2009.04238.x | pmid=19500247|s2cid=8229391 }}</ref> It has been shown that the subsocial spiders of genus ''[[Stegodyphus]]'' tolerate a low level of inbreeding with low inbreeding depression, suggesting a possible stepping stone towards the fully inbred mating system found in social spiders.<ref>{{cite journal |author1=Bilde T. |author2=Lubin Y. |author3=Smith D. |author4=Schneider J. M. |author5=Maklakov A. A. | year = 2005 | title = The transition to social inbred mating systems in spiders: Role of inbreeding tolerance in a subsocial predecessor | journal = Evolution | volume = 59 |issue=1 | pages = 160–174 | doi=10.1554/04-361 | pmid = 15792236|s2cid=13699137 }}</ref><ref>{{cite journal |author1=Ruch J. |author2=Heinrich L. |author3=Bilde T. |author4=Schneider J. M. | year = 2009 | title = The evolution of social inbreeding mating systems in spiders: Limited male mating dispersal and lack of pre-copulatory inbreeding avoidance in a subsocial predecessor | journal = Biological Journal of the Linnean Society | volume = 98 | issue = 4| pages = 851–859 | doi=10.1111/j.1095-8312.2009.01322.x| doi-access = free }}</ref>


===Swarming===
===Swarming===

Revision as of 12:27, 15 June 2021

A social spider is a spider species whose individuals form relatively long-lasting aggregations. Whereas most spiders are solitary and even aggressive toward other members of their own species, some hundreds of species in several families show a tendency to live in groups, often referred to as colonies.

Spider sociality

Most species of social spiders live in the tropical regions of the world where insect size and density is highest but several species reach into the eastern United States and other temperate areas. By building a communal web, it is thought that the spiders approximately maximize total biomass capture per spider.[1] Having a larger web and multiple spiders to work together to subdue prey allows them to prey on larger organisms than would be possible if they led a solitary existence. The colonies can grow large enough to take down birds and bats, as well as very large insects.

Living in a colony also has another major benefit for spiders: cooperative nest maintenance. Nest maintenance does not rely solely on an individual in a colony setting and thus saves on a per-capita investment in maintaining silk structures. Predator defense is also increased in a colony with a large web and multiple individuals analogous to schools of fish or herds of mammals.[2]

Levels

Social spiders from Bengaluru, India

Social spiders exhibit varying levels of sociality, of which there are six defined. Agnarsson et al. estimate that spiders as a whole have independently evolved sociality 18 or 19 times.[3] Most of these social spiders broadly fit into the quasi-social definition of sociality, meaning they show cooperative brood care, use the same nest (web), and have some amount of generational overlap. Several permutations of social behavior exist amongst the 23 species of spider considered to be quasi-social out of some 45,000 known species of spider.[4] These 23 species are phylogenetically scattered in 11 genera across eight widely separated families.[3]

The level of sociality often varies between species (interspecies) but can vary within a species (intraspecies) as well. Intraspecific variation is generally habitat dependant, where some populations within a species show all the characteristics of quasi-sociality, yet a population a mile away may be largely solitary[5] because they inhabit a different environment. This facultative sociality allows them to survive periods of sub-optimal conditions, when sustaining large aggregations is not feasible. Some of these aggregations can contain as many as 50,000 individuals as in the case of Anelosimus eximius (in the family Theridiidae).[6] The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social.[7]

Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food.[8] Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae).[9] Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this.[10] The herbivorous spider Bagheera kiplingi lives in small colonies which helps to protect eggs and spiderlings.[11] Even widow spiders (genus Latrodectus), which are notoriously aggressive and cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.[12]

A few species, such as Anelosimus eximius, are also known to have reproductive skew, where some of the females reproduce and others do not.[13] Even though these spiders cooperate by caring for each other's young, cooperating in prey capture and sharing food, they are not considered eusocial as they do not have defined castes. Any female has the potential to reproduce, therefore differences in the reproductive success is due to competition for scarce food resources rather than the existence of physically distinct castes.[14]

Sub-social

Many more species of spider are considered to be sub-social than quasi-social – meaning they lack fixed or complex social organization. These species may only display social behaviors as a seasonal venture and have an obligate solitary phase. Some other species will establish territories within the colony and can even have discrete webs, narrowly connected to other webs within the colony (such is the case with Leucauge spp.[15]). This is not a fully cooperative behavior as there is little to no cooperative nest maintenance or brood care occurring. The subsocial species, however, appear to be crucial for the evolution of sociality in spiders. Recent theory suggests that social spiders evolved along a restricted pathway through solitary subsocial ancestors.[3][16] It has been shown that the subsocial spiders of genus Stegodyphus tolerate a low level of inbreeding with low inbreeding depression, suggesting a possible stepping stone towards the fully inbred mating system found in social spiders.[17][18]

Swarming

Several social spiders including Parasteatoda wau and Anelosimus eximius also swarm in an analogous way to the eusocial ants, bees and wasps.[5] These species disperse and establish new colonies by means of synchronized emigrations of adult and sub-adult females. After courtship and copulation, but prior to oviposition, many females will emigrate to a new nesting site and deposit their eggs, forming a new colony. In this way social spiders are also extremely inbred, as there is limited migration of males or juveniles to different colonies, forcing the offspring to mate with one another decreasing genetic variation within the colonies.[5] Occasionally males will emigrate with females or will emigrate from one colony to another but this is a rare event and has not been studied sufficiently to quantify for any social spider species. Females can also out-number males as much as 10:1 in many species; this too acts as a genetic bottleneck and further decreases the genetic variation of the species.

Quasi-social spider families and genera

Species include:[19]

References

  1. ^ Yip EC, Powers KS, Avilés L (2008). "Cooperative capture of large prey solves scaling challenge faced by spider societies". Proceedings of the National Academy of Sciences. 105 (33): 11818–11822. Bibcode:2008PNAS..10511818Y. doi:10.1073/pnas.0710603105. PMC 2575263. PMID 18689677.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  2. ^ "Social organization of the colonial spider Leucauge sp in the Neotropics vertical stratification within colonies" (PDF).
  3. ^ a b c Ingi Agnarsson; Leticia Aviles; Jonathan A. Coddington & Wayne P. Maddison (2006). "Sociality In Theridiid Sspiders: Repeated Origins Of An Evolutionary Dead End". Evolution. 60 (11): 2342–51. doi:10.1554/06-078.1. PMID 17236425. S2CID 16995686.
  4. ^ "World Spider Catalogue". NMBE. Retrieved 31 October 2015.
  5. ^ a b c Lubin, Yael D. & Robinson Michael H. (1982). "Dispersal by Swarming in a Social Spider". Science. 216 (4543): 319–21. Bibcode:1982Sci...216..319L. doi:10.1126/science.216.4543.319. PMID 17832747. S2CID 32470345.
  6. ^ Vollrath, F. (1986). "Eusociality and extraordinary sex ratios in the spider Anelosimus eximius (Araneae: Theridiidae)". Behavioral Ecology and Sociobiology. 18 (4): 283–287. doi:10.1007/BF00300005. S2CID 44727810.
  7. ^ Agnarsson, I. & Kuntner, M. (April 2005). "Madagascar: an unexpected hotspot of social Anelosimus spider diversity (Araneae: Theridiidae)". Systematic Entomology. 30 (4): 575–592. doi:10.1111/j.1365-3113.2005.00289.x. S2CID 13871079.
  8. ^ Avilés, L., Maddison, W.P. and Agnarsson, I. (August 2006). "A New Independently Derived Social Spider with Explosive Colony Proliferation and a Female Size Dimorphism". Biotropica. 38 (6): 743–753. doi:10.1111/j.1744-7429.2006.00202.x.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ Matsumoto, T. (1998). "Cooperative prey capture in the communal web spider, Philoponella raffray (Araneae, Uloboridae)" (PDF). Journal of Arachnology. 26: 392–396. Retrieved 2008-10-11.
  10. ^ Cangialosi, K.R. (July 1990). "Social spider defense against kleptoparasitism". Behavioral Ecology and Sociobiology. 27 (1). doi:10.1007/BF00183313. S2CID 38307165.
  11. ^ Meehan, C,J. Olson, E.J. and Curry, R.L. (21 August 2008). Exploitation of the Pseudomyrmex–Acacia mutualism by a predominantly vegetarian jumping spider (Bagheera kiplingi). 93rd ESA Annual Meeting. Retrieved 2008-10-10.{{cite conference}}: CS1 maint: multiple names: authors list (link)
  12. ^ Bertani, R., Fukushima, C.S., and Martins, R. (May 2008). "Sociable widow spiders? Evidence of subsociality in LatrodectusWalckenaer, 1805 (Araneae, Theridiidae)". Journal of Ethology. 26 (2): 299–302. doi:10.1007/s10164-007-0082-8. S2CID 36475912.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  13. ^ Ann Lundie Rypstra (1993). "Prey Size, Social Competition, and the Development of Reproductive Division of Labor in Social Spider Groups". The American Naturalist. 142 (5): 868–880. doi:10.1086/285577. S2CID 85144990.
  14. ^ Agnarsson, Ingi (2006). "A revision of the New World eximius lineage of Anelosimus (Araneae, Theridiidae) and a phylogenetic analysis using worldwide exemplars". Zoological Journal of the Linnean Society. 146 (4): 453–593. doi:10.1111/j.1096-3642.2006.00213.x.
  15. ^ http://www.americanarachnology.org/JoA_free/JoA_v38_n3/arac-38-03-446.pdf
  16. ^ Johannesen J.; Wickler W.; Seibt U.; Moritz R. F. A. (2009). "Population history in social spiders repeated: Colony structure and lineage evolution in stegodyphus mimosarum (eresidae)". Molecular Ecology. 18 (13): 2812–2818. doi:10.1111/j.1365-294x.2009.04238.x. PMID 19500247. S2CID 8229391.
  17. ^ Bilde T.; Lubin Y.; Smith D.; Schneider J. M.; Maklakov A. A. (2005). "The transition to social inbred mating systems in spiders: Role of inbreeding tolerance in a subsocial predecessor". Evolution. 59 (1): 160–174. doi:10.1554/04-361. PMID 15792236. S2CID 13699137.
  18. ^ Ruch J.; Heinrich L.; Bilde T.; Schneider J. M. (2009). "The evolution of social inbreeding mating systems in spiders: Limited male mating dispersal and lack of pre-copulatory inbreeding avoidance in a subsocial predecessor". Biological Journal of the Linnean Society. 98 (4): 851–859. doi:10.1111/j.1095-8312.2009.01322.x.
  19. ^ Ingi Agnarsson, PhD.: Social Spiders.
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