Scytodes


Scytodes is a genus of spitting spiders that occur all around the world, with the most widely distributed species being Scytodes thoracica, originally having a palearctic distribution but has since been introduced to North America, Argentina, India, Australia, Asia and New Zealand.
Individuals of the genus are generally characterized by their predominant pale yellow coloration and black markings on their cephalothorax compounded with long, thin hairless legs with black bands.
Like other spitting spiders, they do not build a traditional web to catch prey, but instead spit a fluid on their prey that congeals on contact into a venomous and sticky mass.

Taxonomy

The first species of the genus was first described by Pierre André Latreille as Aranea thoraica in 1802 through "Histoire naturelle, générale et particulière des crustacés et des insectes" until the genus was later reclassified by Jean Victor Audouin as Scytodes in 1826.

Appearance

The genus exhibits sexual dimorphism. Males range in size from 3.5 to 4 mm while females are slightly larger, from 4 to 4.5 mm. Spitting spiders have pale yellow bodies with black spots on their wide cephalothorax, and legs characterized by black bands. Scytodes species have three paired eyes for a total of six eyes. Their legs are long, slender, and have small claws attached to the chelicerae.

Habitat

Scytodes species are mainly nocturnal and typically do not live in webs with the exception of some tropical species, such as Scytodes longpipes. The web, however, is not used for catching prey. Instead, Scytodes species live under rocks, in crevices and can even be found living in human infrastructures.

Niche

Similar to other arachnids, Scytodes species occupy a vital role as secondary consumers and work to control the populations of prey items in their habitats.

Distribution

Scytodes has a near-global distribution, being present on almost every continent excluding Antarctica. The majority of species within Europe and the Mediterranean are distributed around the 45th parallel. Scytodes enjoys a sizable presence in Asia, Oceania and North America but extremely limited presence in Africa and South America.

Behavior

Reproduction

Scytodes species are typically solitary until mating or hunting due to their aggressive nature. Males are cautious when trying to find a mate. Females carry their eggs until they hatch, typically under their body or in their chelicerae. This is the most vulnerable stage in life, the egg-carrying period. Upon hatching, the juvenile spiders remain in their mother's web. They cooperatively capture and feed on prey caught in the web. Upon reaching sexual maturity, the young spiders leave the web, move a short distance away and exhibit solitary behavior.
Sociality and parental care
Scytodes species are known to perform a wide range of social organization, with some species being solitary, subsocial, communal-territorial, or social. Many of the subsocial and colonial species are known to give extended parental care, such as S. fusca, S. socialis, S. longipes, and S. intricata.
Females will sparsely wrap their eggs in a sac, and hold it in its chelicerae until they hatch, sometimes dropping the egg sac to capture prey, after which they return and feed beside the eggs, picking them up again once finished. This behaviour protects the eggs from desiccation, mould, parasites, and predation.
Some species are subsocial and after hatching, the spiderlings will often cohabitate with their mother until a certain instar is reached, but some may stay longer or leave earlier. During this period the spiderling and adult will hunt prey, but not spit at or prey on each other. The mothers will sometimes take captured prey to the spiderlings. S. indet and S. indet will often share the meal with their spiderlings, but S. fusca will not, eating separate meals away from the spiderlings. Cohabitating spiderlings will work together to capture large prey, but will hunt small prey individually. Conflict between spiderlings occurs in some species at early instars, but cannibalistic behaviour only arises in later instars of those species.
S. socialis is known to be a social species, and individuals will continue cohabitation into adulthood, forming colonies, in which individuals show no aggression to each other and share prey.

Hunting

Scytodes species primarily detect prey by vibration and smell. They will spit at prey until it is subdued. Once the prey is subdued, they wrap the prey in silk, and feed. When prey flees, Scytodes will sometimes pursue it, and spit web once it gets close. Prey often gets glued to substrate by the spit, and the spider will cut it loose from the substrate to wrap it. Sometimes when prey struggles during wrapping, or immediately after spitting web, the spider will quickly lunge and stab the prey with its fangs, retracting them immediately. Many species are known to be araneophagic, with some preferring to prey on other spiders.
Hunting dangerous prey such as other spiders is thought to lead to behavioural flexibility, which can be observed in Scytodes, which will adjust their hunting methods based on prey size.
S. pallida shares an interesting relationship with the jumping spider Portia labiata, in which both are each other's primary prey. These spiders do not die post-mating; males live 1.5–2 years and females live 2 to 4 years.

Webs

Some species of Scytodes are known to build webs. Webs of many species consist of a sheet of web connected to a small tubular shelter, in which the spider tends to reside.

Spitting

The spitting from which its common name derives is used as a method of trapping prey or escaping predators. Scytodes glue is made in a large gland in the cephalothorax where venom is synthesized in the front of the gland and the glue is produced in the back of gland. Sticky glue is expelled from their fangs from a small opening in their chelicerae and can be shot up to 4 -10 body lengths from the spider. Scytodes spit their glue in a zigzag pattern to trap their prey to a substrate. Though, for many years it was thought that the glue contained venom that diffused into the prey, researchers now believe that the glue does not contain any toxic properties. Once the whole prey is trapped to the ground, the spider then bites the prey with a venomous bite to kill it.

Diet

Scytodes primarily feed on soft bodied insects such as moths, other spiders, mosquitoes, silverfish and crickets. They typically hunt their prey, however scavenging behaviour has been observed to happen rarely. When scavenging occurs a spider will not spit on the prey as the glue is energetically expensive to produce.

Fossil record

One of the oldest fossils of an extinct species of Scytodes is Scytodes weitschati found in Baltic amber dating to the Eocene 43 million years ago. Its morphology is similar to that of Scytodes seen today.

Species

, this genus includes 228 species and one subspecies.
Species with articles on Wikipedia:Scytodes arenacea Purcell, 1904 – Namibia, South AfricaScytodes armata Brescovit & Rheims, 2001 – Costa RicaScytodes broomi Pocock, 1902 – Namibia, South AfricaScytodes caffra Purcell, 1904 – DR Congo, South Africa, EswatiniScytodes cedri Purcell, 1904 – South AfricaScytodes clavata Benoit, 1965 – DR Congo, South AfricaScytodes constellata Lawrence, 1938 – South AfricaScytodes drakensbergensis Lawrence, 1947 – South AfricaScytodes elizabethae Purcell, 1904 – South AfricaScytodes flagellata Purcell, 1904 – South AfricaScytodes fusca Walckenaer, 1837 – North to South America. Introduced to St. Helena, Europe, Africa, Seychelles, India, Myanmar, Indonesia, China, Japan, HawaiiScytodes globula Nicolet, 1849 – Bolivia, Brazil, Uruguay, Chile, ArgentinaScytodes gooldi Purcell, 1904 – South AfricaScytodes karrooica Purcell, 1904 – South AfricaScytodes kumonga Zamani & Marusik, 2020 – Oman, IranScytodes lanceolata Purcell, 1904 – South AfricaScytodes lawrencei Lessert, 1939 – DR Congo, South AfricaScytodes leipoldti Purcell, 1904 – South AfricaScytodes longipes Lucas, 1844 – Central and South America. Introduced to Guinea, DR Congo, Indonesia, Japan, Australia, Pacific Is. HawaiiScytodes lycosella Purcell, 1904 – South AfricaScytodes lyriformis Purcell, 1904 – South AfricaScytodes maritima Lawrence, 1938 – South AfricaScytodes marshalli Pocock, 1902 – South AfricaScytodes montana Purcell, 1904 – South AfricaScytodes pallida Doleschall, 1859 – India, China, Philippines, Indonesia Scytodes quinqua Lawrence, 1927 – Namibia, South AfricaScytodes rubra Lawrence, 1937 – South AfricaScytodes schultzei Purcell, 1908 – South AfricaScytodes silvatica Purcell, 1904 – South AfricaScytodes subulata Purcell, 1904 – South AfricaScytodes symmetrica Lawrence, 1938 – South AfricaScytodes testudo Purcell, 1904 – South AfricaScytodes thoracica – Europe, North Africa, Turkey, Iran, temperate Asia to China, Korea, Japan. Introduced to North America, Argentina, South Africa, India, Australia, New Zealand Scytodes triangulifera Purcell, 1904 – South AfricaScytodes trifoliata Lawrence, 1938 – South AfricaScytodes univittata Simon, 1882 – Turkmenistan, Kyrgyzstan, India. Introduced to Hawaii, Mexico, Cuba, Venezuela, Brazil, Paraguay, Chile, Ascension Island, Cape Verde, Canary Islands, Morocco, Spain, Italy, Cyprus, Egypt, Iraq, IranScytodes venusta – Sri Lanka to Indonesia S. adisi Rheims & Brescovit, 2009 – BrazilS. aethiopica Simon, 1907 – EthiopiaS. affinis Kulczyński, 1901 – EthiopiaS. aharonii Strand, 1914 – IsraelS. akytaba Rheims & Brescovit, 2006 – BrazilS. alayoi Alayón, 1977 – Mexico, CubaS. albiapicalis Strand, 1907 – ChinaS. alcomitzli Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. alfredi Gajbe, 2004 – IndiaS. altamira Rheims & Brescovit, 2000 – BrazilS. amazonica Dupérré & Tapia, 2023 – EcuadorS. annulipes Simon, 1907 – Morocco, Algeria, Tunisia, LibyaS. antonina Rheims & Brescovit, 2009 – BrazilS. apuecatu Rheims & Brescovit, 2006 – BrazilS. arboricola Millot, 1946 – Ivory CoastS. arenacea Purcell, 1904 – Namibia, South AfricaS. argelia Dupérré & Tapia, 2023 – EcuadorS. armata Brescovit & Rheims, 2001 – Costa RicaS. aruensis Strand, 1911 – Indonesia S. arwa Rheims, Brescovit & van Harten, 2006 – Yemen, Oman, IranS. atlacamani Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. atlacoya Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. atlatonin Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. auricula Rheims & Brescovit, 2000 – Ecuador, BrazilS. ayampe Dupérré & Tapia, 2023 – EcuadorS. balbina Rheims & Brescovit, 2000 – BrazilS. becki Rheims & Brescovit, 2001 – BrazilS. bertheloti Lucas, 1838 – Mediterranean, TurkmenistanS. blanda Bryant, 1940 – CubaS. bocaina Rheims & Brescovit, 2009 – BrazilS. bonito Rheims & Brescovit, 2009 – BrazilS. brignolii Rheims & Brescovit, 2009 – BrazilS. broomi Pocock, 1902 – Namibia, South AfricaS. brunnea González-Sponga, 2004 – VenezuelaS. caffra Purcell, 1904 – DR Congo, South Africa, EswatiniS. caipora Rheims & Brescovit, 2004 – BrazilS. camerunensis Strand, 1906 – CameroonS. canariensis Wunderlich, 1987 – Canary IslandsS. caratinga Rheims & Brescovit, 2009 – BrazilS. caure Rheims & Brescovit, 2004 – BrazilS. cavernarum Roewer, 1962 – MalaysiaS. cedri Purcell, 1904 – South AfricaS. cellularis Simon, 1907 – DR CongoS. championi F. O. Pickard-Cambridge, 1899 – Mexico, BrazilS. chantico Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. chapeco Rheims & Brescovit, 2009 – BrazilS. chiconahui Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. chiquimula Brescovit & Rheims, 2001 – GuatemalaS. choco Dupérré & Tapia, 2023 – EcuadorS. chopim Rheims & Brescovit, 2009 – BrazilS. clavata Benoit, 1965 – DR Congo, South AfricaS. cogu Brescovit & Rheims, 2001 – Costa RicaS. congoanus Strand, 1908 – CongoS. constellata Lawrence, 1938 – South AfricaS. coronata Thorell, 1899 – West AfricaS. costa Dupérré & Tapia, 2023 – EcuadorS. cotopitoka Rheims, Barreiros, Brescovit & Bonaldo, 2005 – BrazilS. cubensis Alayón, 1977 – Cuba, TrinidadS. curimaguana González-Sponga, 2004 – VenezuelaS. curupira Rheims & Brescovit, 2004 – BrazilS. darlingtoni Alayón, 1977 – CubaS. diminuta Valerio, 1981 – Costa RicaS. dissimulans Petrunkevitch, 1929 – Puerto RicoS. dollfusi Millot, 1941 – Ivory CoastS. domhelvecio Rheims & Brescovit, 2009 – BrazilS. dorothea Gertsch, 1935 – United StatesS. drakensbergensis Lawrence, 1947 – South AfricaS. eleonorae Rheims & Brescovit, 2001 – BrazilS. elizabethae Purcell, 1904 – South AfricaS. farri Alayón, 1985 – JamaicaS. flagellata Purcell, 1904 – South AfricaS. florifera Yin & Xu, 2012 – ChinaS. fourchei Lessert, 1939 – DR CongoS. fusca Walckenaer, 1837 – North, South America. Introduced to St. Helena, Europe, Africa, Seychelles, India, Myanmar, Indonesia, China, Japan, HawaiiS. genebra Rheims & Brescovit, 2009 – BrazilS. gertschi Valerio, 1981 – PanamaS. gilva – India, MyanmarS. globula Nicolet, 1849 – Bolivia, Brazil, Uruguay, Chile, ArgentinaS. gooldi Purcell, 1904 – South AfricaS. grammocephala Simon, 1909 – VietnamS. guapiassu Rheims & Brescovit, 2009 – BrazilS. guttipes Simon, 1893 – Venezuela, TrinidadS. hahahae Rheims & Brescovit, 2001 – BrazilS. humilis L. Koch, 1875 – EthiopiaS. iabaday Rheims & Brescovit, 2001 – BrazilS. iara Rheims & Brescovit, 2004 – BrazilS. ilhota Rheims & Brescovit, 2009 – BrazilS. imbituba Rheims & Brescovit, 2009 – BrazilS. immaculata L. Koch, 1875 – Egypt, GreeceS. insperata Soares & Camargo, 1948 – BrazilS. intricata Banks, 1909 – Mexico, Costa RicaS. itabaiana Rheims & Brescovit, 2009 – BrazilS. itacuruassu Rheims & Brescovit, 2006 – BrazilS. itapecerica Rheims & Brescovit, 2009 – BrazilS. itapevi Brescovit & Rheims, 2000 – BrazilS. itzana Chamberlin & Ivie, 1938 – MexicoS. itzli Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. jaguar Dupérré & Tapia, 2023 – EcuadorS. janauari Brescovit & Höfer, 1999 – BrazilS. jousseaumei Simon, 1907 – DjiboutiS. jurubatuba Rheims & Brescovit, 2009 – BrazilS. jurupari Rheims & Brescovit, 2004 – BrazilS. jyapara Rheims & Brescovit, 2006 – BrazilS. kaokoensis Lawrence, 1928 – NamibiaS. karrooica Purcell, 1904 – South AfricaS. kinsukus Patel, 1975 – IndiaS. kinzelbachi Wunderlich, 1995 – Turkey, JordanS. kumonga Zamani & Marusik, 2020 – Oman, IranS. lanceolata Purcell, 1904 – South AfricaS. lawrencei Lessert, 1939 – DR Congo, South AfricaS. leipoldti Purcell, 1904 – South AfricaS. leprosula Strand, 1913 – Central AfricaS. lesserti Millot, 1941 – GuineaS. lewisi Alayón, 1985 – JamaicaS. lineatipes Taczanowski, 1874 – St. Vincent, Bonaire, Venezuela, French GuianaS. liui Wang, 1994 – ChinaS. loja Dupérré & Tapia, 2023 – EcuadorS. longipes Lucas, 1844 – Central America and South America. Introduced to Guinea, DR Congo, Indonesia, Japan, Australia, Pacific Islands, HawaiiS. lorenzoi Alayón, 1977 – CubaS. luteola Simon, 1893 – VenezuelaS. lycosella Purcell, 1904 – South AfricaS. lyriformis Purcell, 1904 – South AfricaS. major Simon, 1886 – AfricaS. mangabeiras Rheims & Brescovit, 2009 – BrazilS. mapia Rheims & Brescovit, 2000 – BrazilS. mapinguari Rheims & Brescovit, 2004 – BrazilS. maquine Rheims & Brescovit, 2009 – BrazilS. maresi Rheims & Brescovit, 2001 – BrazilS. maritima Lawrence, 1938 – South AfricaS. marlieria Rheims & Brescovit, 2009 – BrazilS. maromba Rheims & Brescovit, 2009 – BrazilS. marshalli Pocock, 1902 – South AfricaS. martiusi Brescovit & Höfer, 1999 – BrazilS. mawphlongensis Tikader, 1966 – India, Nepal, ThailandS. mayahuel Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. minus Dupérré & Tapia, 2023 – EcuadorS. montana Purcell, 1904 – South AfricaS. monticola González-Sponga, 2004 – VenezuelaS. multilineata Thorell, 1899 – West, Central AfricaS. nambiobyrassu Rheims & Brescovit, 2009 – BrazilS. nambiussu Rheims & Brescovit, 2006 – BrazilS. nanahuatzin Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. nigristernis Simon, 1907 – Guinea-BissauS. noeli Alayón, 1977 – CubaS. obelisci Denis, 1947 – EgyptS. opoxtli Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. orellana Dupérré & Tapia, 2023 – EcuadorS. oswaldi Lenz, 1891 – MadagascarS. paarmanni Brescovit & Höfer, 1999 – BrazilS. pallida Doleschall, 1859 – India, China, Philippines, Indonesia S. panamensis Brescovit & Rheims, 2001 – PanamaS. panguana Brescovit & Höfer, 1999 – PeruS. paramera González-Sponga, 2004 – VenezuelaS. pholcoides Simon, 1898 – SeychellesS. pintodarochai Rheims & Brescovit, 2009 – BrazilS. piroca Rheims & Brescovit, 2000 – BrazilS. piyampisi Rheims, Barreiros, Brescovit & Bonaldo, 2005 – BrazilS. propinqua Stoliczka, 1869 – Pakistan, IndiaS. pulchella Berland, 1914 – East AfricaS. punctipes Simon, 1907 – São Tomé and PríncipeS. quarta Lawrence, 1927 – NamibiaS. quattuordecemmaculata Strand, 1907 – ChinaS. quinqua Lawrence, 1927 – Namibia, South AfricaS. redempta Chamberlin, 1924 – MexicoS. reticulata Jézéquel, 1964 – Ivory CoastS. robertoi Alayón, 1977 – CubaS. romitii Caporiacco, 1947 – Guyana, BrazilS. rubra Lawrence, 1937 – South AfricaS. ruizensis Strand, 1914 – ColombiaS. rupestris González-Sponga, 2004 – VenezuelaS. saaristoi Rheims & Brescovit, 2009 – BrazilS. saci Rheims & Brescovit, 2004 – BrazilS. sansibarica Strand, 1907 – Tanzania S. schultzei Purcell, 1908 – South AfricaS. semipullata – TibetS. seppoi Bosmans & Van Keer, 2014 – Algeria, TunisiaS. sexstriata Roewer, 1960 – AfghanistanS. silvatica Purcell, 1904 – South AfricaS. sincora Rheims & Brescovit, 2009 – BrazilS. skuki Rheims & Brescovit, 2001 – BrazilS. socialis Miller, 2006 – MadagascarS. sordida Dyal, 1935 – PakistanS. stoliczkai Simon, 1897 – IndiaS. strussmannae Rheims & Brescovit, 2001 – BrazilS. subadulta Strand, 1911 – Indonesia S. subulata Purcell, 1904 – South AfricaS. symmetrica Lawrence, 1938 – South AfricaS. tabuleiro Rheims & Brescovit, 2009 – BrazilS. tacapepucu Rheims & Brescovit, 2006 – BrazilS. tapacura Rheims & Brescovit, 2009 – BrazilS. tapuia Rheims & Brescovit, 2009 – BrazilS. tardigrada Thorell, 1881 – Myanmar, New Guinea, Australia S. tayos Dupérré & Tapia, 2023 – EcuadorS. tegucigalpa Brescovit & Rheims, 2001 – HondurasS. tenerifensis Wunderlich, 1987 – Canary IslandsS. tertia Lawrence, 1927 – Angola, NamibiaS. testudo Purcell, 1904 – South AfricaS. tezcatlipoca Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. thoracica – Europe, North Africa, Turkey, Iran, temperate Asia, China, Korea, Japan. Introduced to North America, Argentina, South Africa, India, Australia, New Zealand S. tikaderi Biswas, 2023 – BangladeshS. tinkuan Rheims & Brescovit, 2004 – BrazilS. tlaloc Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. triangulifera Purcell, 1904 – South AfricaS. trifoliata Lawrence, 1938 – South AfricaS. tropofila González-Sponga, 2004 – VenezuelaS. turvo Rheims & Brescovit, 2009 – BrazilS. tuyucua Brescovit, Rheims & Raizer, 2004 – BrazilS. tyaia Rheims & Brescovit, 2009 – BrazilS. tyaiamiri Rheims & Brescovit, 2006 – BrazilS. tyaiapyssanga Rheims & Brescovit, 2006 – BrazilS. tzitzimime Rheims, Brescovit & Durán-Barrón, 2007 – MexicoS. uligocetes Valerio, 1981 – Costa RicaS. una Rheims & Brescovit, 2009 – BrazilS. univittata Simon, 1882 – Turkmenistan, Kyrgyzstan, India. Introduced to Hawaii, Mexico, Cuba, Venezuela, Brazil, Paraguay, Chile, Ascension Island, Cape Verde, Canary Islands, Morocco, Spain, Italy, Cyprus, Egypt, Iraq, Iran