Scytodes thoracica

Ge­o­graphic Range

Mem­bers of the genus Scy­todes are pre­dom­i­nately trop­i­cal or sub­trop­i­cal spi­ders. How­ever, spit­ting spi­ders are found scat­tered across neartic, paleartic, and neotrop­i­cal re­gions. This species is com­monly found in the east­ern United States, as well as in Britain, Swe­den and other Eu­ro­pean coun­tries. Spec­i­mens have also been found in Japan and Ar­gentina. It is un­clear which pop­u­la­tions are na­tive and which are in­tro­duced in these re­gions. The pres­ence of this species in more north­ern cli­mates is at­trib­uted to the avail­abil­ity of warm houses and build­ings, to which these spi­ders have adapted. ("The Spi­der Book", 1912; Emer­ton, 1902; Kas­ton, 1972; "Scy­todes tho­racica (Spit­ting spi­der)", 2011; Ubick, et al., 2005)

Habi­tat

Spit­ting spi­ders are found in tem­per­ate forests. When as­so­ci­ated with hu­mans, they are most com­monly found in dark cor­ners, cel­lars, cup­boards, and clos­ets of houses and other build­ings. ("Spit­ting spi­der Scy­todes tho­racica", 2012)

  • Average elevation
    Sea Level m
    ft

Phys­i­cal De­scrip­tion

This species has long, thin, legs, and are glabrous (hair­less), with the ex­cep­tion of short sen­sory setae scat­tered over the body. These spi­ders are also eas­ily iden­ti­fied by their over­sized cephalotho­rax (pro­soma), which slopes up­ward to­wards their pos­te­rior ends. Their ab­domens (which are roughly the same cir­cu­lar shape as the cephalotho­rax) slope down­wards and are only slightly smaller than the cephalotho­rax. Like all spi­ders, these two body tag­mata (seg­ments) are sep­a­rated by a thin pedi­cel (waist-like con­nec­tor). (Emer­ton, 1902; Goethals, 1997; Kas­ton, 1972; Larsen, 2005; Robin­son, 2005)

Large, well-de­vel­oped poi­son glands are lo­cated in the cephalotho­rax. These glands are di­vided into two parts: a smaller, an­te­rior com­part­ment which stores venom and a larger, pos­te­rior com­part­ment which con­tains a mu­cilagi­nous sub­stance. These spi­ders pro­duce a gummy sub­stance which is a mix of the two sub­stances and is ex­creted by their fused che­licerae, which can not be moved sep­a­rately. Scy­todes are ecribel­late, lack­ing the silk-spin­ning organ (cribel­lum) of some other spi­ders. They have a sin­gle tra­cheal spir­a­cle. (Emer­ton, 1902; Goethals, 1997; Kas­ton, 1972; Robin­son, 2005)

Spit­ting spi­ders have pale yel­low bod­ies with black speck­led mark­ings on the cephalotho­rax, which slightly re­sem­ble a lyre. Their legs, which slowly taper in size as they ex­tend from their bod­ies, are long with black bands. The most an­te­rior por­tion of the head, below the eyes, pro­jects for­wards, past the mandibles. This species ex­hibits sex­ual di­mor­phism, with males typ­i­cally rang­ing from 3.5-4 mm in length and fe­males rang­ing from 4-5.5 mm. (Emer­ton, 1902; Gertsch, 1949; Goethals, 1997; Kas­ton, 1972; Robin­son, 2005)

  • Sexual Dimorphism
  • female larger
  • Range length
    3.5 to 5.5 mm
    0.14 to 0.22 in

De­vel­op­ment

All spi­ders re­pro­duce sex­u­ally, and sex is de­ter­mined by meio­sis. Fol­low­ing cop­u­la­tion and egg fer­til­iza­tion, fe­males lay eggs in a co­coon car­ried under their bod­ies for 2-3 weeks until eggs hatch. Spi­der­lings re­main with their moth­ers until their first molts and then dis­perse to live soli­tary lives, reach­ing adult­hood after 5-7 molts. It can take 2-3 years for fe­males to reach ma­tu­rity. ("Spit­ting spi­der Scy­todes tho­racica", 2012; Gertsch, 1949; Guar­isco, 1991; Koh, et al., 2009; Larsen, 2005)

Re­pro­duc­tion

Spit­ting spi­ders are soli­tary, in­ter­act­ing only dur­ing mat­ing. These spi­ders are ag­gres­sive hunters, so males must ap­proach fe­males cau­tiously or else be mis­taken for prey. Both sexes pro­duce pheromones, which are de­tected through con­tact with chemosen­sory hairs cov­er­ing the pedi­palps and first pair of legs; fe­male mate choice is based on male pheromone pro­duc­tion. Lo­ca­tion of mates is not based on vi­sual cues and mat­ing usu­ally oc­curs fol­low­ing an ac­ci­den­tal meet­ing of a male and fe­male. Upon meet­ing a fe­male, male spit­ting spi­ders fill their pal­pal or­gans (ac­ces­sory re­pro­duc­tive parts on the ends of each pedi­palp) with sperm. They do this by draw­ing a sperm web across their gen­i­tal open­ings to ac­cu­mu­late sperm droplets, using their third pair of legs. In Scy­todes, the sperm web is a sin­gle thread which typ­i­cally takes a tri­an­gu­lar shape. From there, sperm droplets are drawn into the pal­pal organ (bulb). Males ap­proach fe­males an­te­ri­orly; fe­males raise their cephalotho­raxes to allow males to in­sert both pal­pal or­gans into their gen­i­tal open­ing. Sperm is then de­posited and stored in fe­males' sem­i­nal re­cep­ta­cles. After mat­ing, the pair sep­a­rates im­me­di­ately. Males and fe­males have mul­ti­ple part­ners through­out their life­times. ("Spit­ting spi­der Scy­todes tho­racica", 2012; Foelix, 1982; Koh, et al., 2009; Se­beok, 1977)

Spit­ting spi­ders are dioe­cious and fe­males re­quire 2-3 years to reach ma­tu­rity. Most mat­ing oc­curs in the warmer months (es­pe­cially Au­gust) but these spi­ders can ex­pand their breed­ing sea­sons when liv­ing in warm areas, such as within homes. Sperm can be stored by fe­males for months until eggs are laid. Com­pared to most spi­ders, spit­ting spi­ders lay rel­a­tively few eggs (20-35 eggs per co­coon) and 2-3 co­coons are typ­i­cally pro­duced by a fe­male each year. This species dis­plays ma­ter­nal care both be­fore (fe­males carry egg co­coons) and after hatch­ing, with newly hatched ju­ve­niles re­main­ing with their mother until their first molts. Speed of growth and, there­fore, rate of molt­ing, is closely re­lated to avail­abil­ity of prey and so the time that ju­ve­niles re­main with their mother, as well as the age at which males reach ma­tu­rity, is widely vari­able. These spi­ders can mate mul­ti­ple times and usu­ally die of hunger, ex­haus­tion, or pre­da­tion. (Koh, et al., 2009; Ramel, 2011; Suter and Strat­ton, 2011)

  • Breeding interval
    This species breeds 2-3 times a year.
  • Breeding season
    These spiders breed during warmer summer months.
  • Range number of offspring
    20 to 35
  • Average number of offspring
    28
  • Average age at sexual or reproductive maturity (female)
    2-3 years

Fe­male spit­ting spi­ders ex­hibit parental care. They do not make nests or sus­pend co­coons in webs or other struc­tures, but in­stead lay eggs in a co­coon that they carry ei­ther under their bod­ies or in their che­licerae. Fe­males are more vul­ner­a­ble to pre­da­tion dur­ing this egg-car­ry­ing pe­riod, as they are un­able to feed or de­fend them­selves by spit­ting. Eggs typ­i­cally hatch 2-3 weeks after being laid and nymphs (ju­ve­niles) re­main with their moth­ers until their first molts. On oc­ca­sion, fe­males con­sume egg sacs that have re­sulted from mat­ing with non-pre­ferred males or con­tain de­fec­tive eggs. Males dis­play no parental in­vest­ment after mat­ing. (Koh, et al., 2009; Larsen, 2005)

  • Parental Investment
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • protecting
      • female
  • pre-weaning/fledging
    • protecting
      • female
  • pre-independence
    • protecting
      • female

Lifes­pan/Longevity

Spit­ting spi­ders have a rel­a­tively long life span as they do not die fol­low­ing mat­ing; males live 1.5-2 years and fe­males live 2-4 years. Fe­males may be preyed upon dur­ing their egg-car­ry­ing pe­ri­ods, while males often die of hunger and ex­haus­tion while search­ing for and court­ing fe­males. (Guar­isco, 1991; Larsen, 2005; Ramel, 2011)

  • Range lifespan
    Status: wild
    1.5 to 4 years
  • Average lifespan
    Status: wild
    2.5 years

Be­hav­ior

Spit­ting spi­ders are pri­mar­ily noc­tur­nal. They are wan­der­ing and soli­tary spi­ders that ac­tively hunt their prey and, due to their long, thin legs, are slow mov­ing. They have ex­tremely poor eye­sight and there­fore walk around with their front legs, which are cov­ered with sen­sory setae, held up in order to sense their en­vi­ron­ments. ("Spit­ting Spi­der", 2004; Suter and Strat­ton, 2005)

When these spi­ders come into con­tact with prey, their at­tacks usu­ally pro­ceed in the fol­low­ing order: tap­ping of the front legs, spit­ting, bit­ing, wrap­ping, and then feed­ing. When a spi­der con­tacts a prey item, it ori­ents it­self to­wards the prey and slowly taps its front legs until the prey is cen­tered be­tween them. It then spits a string of a gluey, ven­omous sub­stance at the prey, cov­er­ing it in 5-17 par­al­lel, over­lap­ping bands. The ma­te­r­ial can travel at a rate of up to 28 m/s and the pat­tern of the bands is pro­duced as the spi­der lifts its che­licerae and os­cil­lates its fangs dur­ing ex­pul­sion. The spi­der then quickly ap­proaches its prey and uses its first and sec­ond pairs of legs to fur­ther en­tan­gle the prey in the dry­ing glue, silk and venom. The ven­omous glue par­a­lyzes the prey and, once it is dry, the spi­der bites its vic­tim, in­ject­ing venom to liquify its tis­sues. It then grooms its first two sets of legs, clean­ing any re­main­ing glue, be­fore draw­ing the prey into its che­licerae using its pedi­palps. The spi­der holds the prey with its third legs and wraps it in silk pro­duced by spin­nerets. It then con­sumes its prey by suck­ing up the dis­solved tis­sues. These spi­ders also use their spit­ting "web" as a de­fen­sive mea­sure against other spi­ders or threats. ("Spit­ting Spi­der", 2004; Suter and Strat­ton, 2005; Suter and Strat­ton, 2011)

  • Range territory size
    1 to 35 m^2

Home Range

Spit­ting spi­ders will typ­i­cally wan­der within a ter­ri­tory 30-35 m in size. ("Spit­ting Spi­der", 2004)

Com­mu­ni­ca­tion and Per­cep­tion

Al­though they can see, these spi­ders do not rely on vi­sion for mate or prey recog­ni­tion due to their poor eye­sight. How­ever, like most spi­ders, they use chem­i­cal sig­nals to de­tect and re­spond to preda­tors, prey, and mates. Spit­ting spi­ders use their raised front legs to de­tect prey and their en­vi­ron­ment using tri­choboth­ria (sen­sory hairs) on the metatarsi (sec­ond to last seg­ment of the leg). These sen­sory hairs also con­tain chemore­cep­tors, which are sen­si­tive to pheromones. Scy­todes are also known to em­ploy both acoustic and vi­bra­tional com­mu­ni­ca­tion by tap­ping on the ground with their first pair of legs. (Koh, et al., 2009; Se­beok, 1977)

Food Habits

Scy­todes are ac­tive noc­tur­nal wan­der­ers, not web-spin­ners. They are in­sec­ti­vores and when liv­ing in­doors, they mostly eat other in­sects and arthro­pods, such as moths (Order Lep­i­doptera), Flies (Order Diptera), other spi­ders (Order Araneae), and house­hold bugs (Order Het­eroptera). When liv­ing out­doors, they eat sim­i­lar food items and, in the Philip­pines, are known to eat in­sects such as green scale (Coc­cus viridis), black cit­rus aphids (Tox­optera au­ran­tii), cit­rus mealy­bugs (Planococ­cus citri), Philip­pine katy­dids (Phaneroptera fur­cifera) and lime swal­low­tails (Pa­pilio de­moleus). In Britain, mos­qui­tos (Fam­ily Culi­ci­dae) are often eaten. Many of their prey items are sig­nif­i­cantly larger than these spi­ders. Fe­males may also oc­ca­sion­ally con­sume their own egg sacs. ("Spi­der Fauna in Philip­pine cit­rus and their note as preda­tor in the con­trol of in­sect", 1989; "Spit­ting Spi­der", 2004; Gertsch, 1949; Gilbert and Rayor, 1985; Larsen, 2005; Med­lock and Snow, 2008)

  • Animal Foods
  • insects

Pre­da­tion

Since spit­ting spi­ders are slow mov­ing, they use their spit­ting ap­pa­ra­tus against preda­tors, rather than at­tempt­ing to flee. In­doors, the ma­jor­ity of its preda­tors are house­hold arthro­pods such as other spi­ders and cen­tipedes, and may in­clude mam­mals such as do­mes­tic cats. In the wild, spit­ting spi­ders may be prey for bats, toads, birds, and shrews. Ad­di­tion­ally, hu­mans play a major fac­tor in con­trol­ling pop­u­la­tions of this species through pest con­trol. (Atkin­son, 2002; Bris­towe, 1941; Suter and Strat­ton, 2005)

Ecosys­tem Roles

When as­so­ci­ated with hu­mans, this spi­der's largest ecosys­tem role is in the con­trol of in­sect pop­u­la­tions, mostly of house­hold pest species. They also serve as food for house cen­tipedes (Scutig­era coleop­trata) and other arthro­pod car­ni­vores found in homes. In the wild, they may be preyed upon by a num­ber of other species. (Gertsch, 1949; Kas­ton, 1948)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

This species con­tributes to the dis­posal of house­hold pest in­sects and could po­ten­tially save home­own­ers money on in­vest­ing in other meth­ods of pest con­trol. (Bris­towe, 1941; Gertsch, 1949)

  • Positive Impacts
  • controls pest population

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Though this species plays a role in con­trol­ling in­sect pop­u­la­tions, it may also be con­sid­ered a house­hold pest. Many home-own­ers in­vest in pest-con­trol in order to ex­ter­mi­nate these spi­ders. Ad­di­tion­ally, this spi­der is ven­omous, al­though its che­licerae, or fangs, are too small to pierce human flesh. (Bris­towe, 1941; Gertsch, 1949)

  • Negative Impacts
  • household pest

Con­ser­va­tion Sta­tus

This species is a com­mon house spi­der of the United States. It is less com­monly found in Eu­rope, Ar­gentina, and Japan, but its con­ser­va­tion sta­tus is not a con­cern. ("Spit­ting Spi­der", 2004)

Con­trib­u­tors

Jacque­line Brand (au­thor), Uni­ver­sity of Michi­gan-Ann Arbor, Je­remy Wright (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

delayed fertilization

a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

female parental care

parental care is carried out by females

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

insectivore

An animal that eats mainly insects or spiders.

internal fertilization

fertilization takes place within the female's body

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

nocturnal

active during the night

oceanic islands

islands that are not part of continental shelf areas, they are not, and have never been, connected to a continental land mass, most typically these are volcanic islands.

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

venomous

an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).

vibrations

movements of a hard surface that are produced by animals as signals to others

visual

uses sight to communicate

Ref­er­ences

1912. The Spi­der Book. Gar­den City, new York: Gar­den City, new York.

Myers En­ter­prises II 2009 - 2012. 2011. "Scy­todes tho­racica (Spit­ting spi­der)" (On-line). Global Species. Ac­cessed Jan­u­ary 26, 2012 at http://​globalspecies.​org/​ntaxa/​65738.

La­guna Col­lege. Spi­der Fauna in Philip­pine cit­rus and their note as preda­tor in the con­trol of in­sect. PCARRD-ASSP Proj. No. 89-38623. Taguig, Philip­pines: The Sci­ence and Tech­nol­ogy In­for­ma­tion of the Philip­pines. 1989. Ac­cessed Jan­u­ary 17, 2013 at http://​scinet.​dost.​gov.​ph/​union/​ShowSearchResult.​php?​s=2&​f=&​p=&​x=&​page=&​sid=1&​id=Spider+fauna+in+Philippine+citrus+and+their+note+as+predator+in+the+control+of+insect+pests.&​Mtype=REPORTS.

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2012. "Spit­ting spi­der Scy­todes tho­racica" (On-line). BBC. Ac­cessed Jan­u­ary 25, 2012 at http://​www.​bbc.​co.​uk/​nature/​wildfacts/​factfiles/​354.​shtml.

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Emer­ton, J. 1902. The Com­mon Spi­ders of the United States. Boston, Lon­don: Ginn & Com­pany, Pub­lish­ers.

Foelix, R. 1982. Bi­ol­ogy of Spi­ders. Cam­bridge, Mass­a­chu­setts, and Lon­don, Eng­land: Har­vard Uni­ver­sity Press.

Gertsch, J. 1949. Amer­i­can Spi­ders. New York: Van Nos­trand. Ac­cessed Jan­u­ary 25, 2012 at http://​babel.​hathitrust.​org/​cgi/​pt?​view=image;size=100;id=mdp.39015006146800;page=root;seq=327;num=235.

Gilbert, C., L. Rayor. 1985. PREDA­TORY BE­HAV­IOR OF SPIT­TING SPI­DERS (ARANEAE : SCY­TO­DI­DAE) AND THE EVO­LU­TION OF PREY WRAP­PING I. Jour­nal of Arach­nol­ogy, 13: 231-241.

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Guar­isco, H. 1991. Three Species of House Spi­ders First Recorded in Kansas. Trans­ac­tions of the Kansas Acad­emy of Sci­ence, 94: 73-76.

Kas­ton, B. 1972. How To Know: The Spi­ders. Dubuque, Iowa: WM. C. Brown Com­pany Pub­lish­ers.

Kas­ton, B. 1948. Spi­ders of Con­necti­cut. Hart­ford: Con­necti­cut.

Koh, T., W. Seah, L. Yap, D. Li. 2009. Pheromone-based fe­male mate choice and its ef­fect on re­pro­duc­tive in­vest­ment in a spit­ting spi­der. Be­hav­ioral Ecol­ogy So­cio­bi­ol­ogy, 63: 923–930. Ac­cessed March 23, 2012 at http://​dl2af5jf3e.​search.​serialssolutions.​com.​proxy.​lib.​umich.​edu/?​sid=jstor:​jstor&​genre=article&​issn=03405443&​eissn=14320762&​volume=63&​pages=+923-930&​spage=923&​epage=930&​atitle=Pheromone-Based+Female+Mate+Choice+and+Its+Effect+on+Reproductive+Investment+in+a+Spitting+Spider&​date=2009&​issue=6.

Larsen, N. 2005. "Fam­ily: Scy­to­di­dae (spit­ting spi­ders)" (On-line). Bio­di­ver­sity Ex­plorer. Ac­cessed Jan­u­ary 25, 2012 at http://​www.​biodiversityexplorer.​org/​arachnids/​spiders/​scytodidae/​index.​htm.

Med­lock, J., K. Snow. 2008. Nat­ural preda­tors and par­a­sites of British mos­qui­toes-a re­view. Eu­ro­pean Mos­quito Bul­letin, 25: 1-11. Ac­cessed Jan­u­ary 17, 2013 at http://​e-m-b.​org/​sites/​e-m-b.​org/​files/​European_​Mosquito_​Bulletin_​Publications811/​EMB25/​EMB25_​1.​pdf.

Ramel, G. 2011. "Re­pro­duc­tion in Spi­ders" (On-line). The Earth­life Web. Ac­cessed March 23, 2012 at http://​www.​earthlife.​net/​chelicerata/​s-reproduce.​html.

Robin­son, W. 2005. Hand­book of Urban In­sects and Arach­nids. Cam­bridge: Cam­bridge Uni­ver­sity Press.

Se­beok, T. 1977. How An­i­mals Com­mu­ni­cate. Bloom­ing­ton & Lon­don: In­di­ana Uni­ver­sity Press.

Suter, R., G. Strat­ton. 2011. Does al­lo­met­ric growth ex­plain the diminu­tive size of the fangs of Scy­todes (Araneae: Scy­to­di­dae)? (. The Jour­nal of Arach­nol­ogy, 39: 174-177. Ac­cessed Jan­u­ary 17, 2013 at http://​www.​americanarachnology.​org/​JoA_​free/​JoA_​v39_​n1/​arac-39-01-174.​pdf.

Suter, R., G. Strat­ton. 2005. Scy­todes vs. Schizo­cosa: Preda­tory Tech­niques and Their Mor­pho­log­i­cal Cor­re­lates. The Jour­nal of Arach­nol­ogy, 33: 7-15. Ac­cessed Feb­ru­ary 23, 2012 at http://​www.​jstor.​org.​proxy.​lib.​umich.​edu/​stable/​3706373?&​Search=yes&​searchText=scytodes&​list=hide&​searchUri=%2Faction%2FdoBasicSearch%3FQuery%3Dscytodes%26gw%3Djtx%26acc%3Don%26prq%3DTHE%2BEVOLUTION%2BOF%2BSOCIALITY%2BIN%2BTHE%2BSPITTING%2BSPIDER%2BSCYTODES-FUSCA%2BARANEAE%2BSCYTODIDAE%2BEVIDENCE%2BFROM%2BOBSERVATIONS%2BOF%2BINTRASPECIFIC%2BINTERACTIONS%26Search%3DSearch%26hp%3D25%26wc%3Don&​prevSearch=&​item=3&​ttl=110&​returnArticleService=showFullText.

Ubick, D., P. Paquin, P. Cush­ing, V. Roth. 2005. Spi­ders of North Amer­ica: an iden­ti­fi­ca­tion man­ual. Ox­ford Uni­ver­sity Press: Amer­i­can Arach­no­log­i­cal So­ci­ety.