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Eucteniza

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Eucteniza ( /jktəˈnzə/ yook-tə-NY-zə[1]) is a genus of trapdoor spiders in the family Euctenizidae containing at least 14 species occurring in Mexico and the southern United States. Species are distinguished by a softened rear portion of the carapace, and males possess large spines on the first two pairs of walking legs that are used to hold females during mating. Like other trapdoor spiders they create burrows with a hinged lid, from which they await passing insects and other arthropods to prey upon. Many species are known from only one or two localities, or from only male specimens. More species are expected to be discovered. Eucteniza is closely related to spiders of the genera Entychides and Neoapachella.

Description

Members of Eucteniza reach up to 27.5 mm (1.08 in) in body length (not including the legs), which is relatively large to very large among spiders. The cephalothorax (the anterior body segment bearing the eyes, mouthparts, and legs) is oval, slightly longer than wide, and slopes considerably towards the rear when viewed from the side. The rear third of the carapace (the upper surface of the cephalothorax) is relatively soft, and is distinctly lighter in coloration. The carapace is often devoid of hairs, and in some species fringed with black bristles. The eight eyes are not elevated and are arranged in two rows in a rectangular area: the hind row is recurved, or slightly curved upwards (as in a bowl or shallow U), while the anterior row is slightly procurved (slightly curved downwards, as in an inverted bowl). Eucteniza spiders are colored in various shades of brown, with males generally appearing a dark reddish brown. The jaws (chelicerae) are dark brown. The abdomen sometimes has a dark brown blotch on the upper surface.[2][3] Measurements of most species only refer to the cephalothorax, which is less variable than the abdomen: recorded cephalothorax lengths range from 3.5 mm (0.14 in) in E. huasteca to 11.5 mm (0.45 in) in E. relata.[2]

In males, the tibiae (5th leg segments) of the first and second walking legs are swollen or enlarged in many species. Additionally, males possess one or two prominent spines, known as "mating claspers" on the first and second tibiae; related genera possess claspers on the first pair of legs only. The size, number, and arrangement of these spines varies between species. Females have a double-toothed groove that the fangs recede into, unlike that of any other euctenizid genus. Both sexes also possess "preening combs", rows of stiff bristles on the rear legs.[2][3]

Behavior

Eucteniza relata builds un-branched wafer-lid burrows.

Like other trapdoor spiders, Eucteniza species construct burrows in the ground with a hinged lid, from which they wait to ambush passing prey. Burrows and prey have been studied in E. relata, and other species are believed to have similar behavior. The burrow consists of an un-branched tube, lined with silk and soil, ranging from 7 to 25 cm (2.8 to 9.8 in) in depth. The burrow is covered with a cork-like lid composed of silk and soil and hinged with silk. The bottom of the burrow accumulates molted exoskeletons and remains of prey, which include beetles, ants, and millipedes. Burrows of adults and juveniles have been found in close proximity, suggesting individuals do not disperse very far after hatching. Individuals in Texas disperse from their burrows between August and January, and hence are most often collected during this time. In Mexico, the dispersal period extends from June to January.[3]

Habitat

The predominant habitat of Eucteniza is desert and tropical dry forest of Mexico and Texas. Specimens have been collected from elevations as low as 8 to 12 m (26 to 39 ft) above sea level (e.g. the Baja California species E. cabowabo, E. diablo, and E. rosalia), to around 3,300 m (10,800 ft) (E. relata in northeastern Mexico).[2] Burrows are located on flat ground or slight inclines.[3] In southern Texas, E. relata may burrow in residential lawns, and may be particularly likely to be encountered after rains.[4] Eucteniza spiders are generally difficult to find in the wild, and are rare in museum collections.[2]

Predators

E. relata in Texas is thought to be preyed upon by tarantula hawk wasps in the genus Pepsis, which are known to paralyze tarantulas and other spiders to provide food for their young. Observations suggest female Pepsis seek out Eucteniza hosts and sting them in their burrows, leaving them paralyzed within.[4]

Taxonomy

E. cabowabo, male, native to Baja California Sur

The taxonomic history of Eucteniza spiders includes nearly 20 named species and four genera; the three additional generic names are now considered taxonomic synonyms of Eucteniza, and several named species have similarly been determined to be synonyms of previously described species.

The genus Eucteniza was established by Austrian naturalist Anton Ausserer in 1875 with the description of Eucteniza mexicana.[5] In 1895, the English zoologist and reverend Octavius Pickard-Cambridge described the new genera Favila[a] and Enrico[b] for the new species relatus and mexicanus, respectively.[6] Shortly thereafter, the reverend's nephew Frederick O. Pickard-Cambridge transferred F. relatus to Eucteniza, rendering Favila a junior synonym of Eucteniza.[7] Similarly, the elder Pickard-Cambridge's Enrico mexicanus was synonymized with Eucteniza in 2002, which required a name change to avoid confusion with Ausserer's Eucteniza mexicana. Thus Enrico mexicanus was given the replacement name Eucteniza atoyacensis.[3]

In 1940, the American biologist Ralph V. Chamberlin described Astrosoga rex, a new genus and species from Texas,[8] while Willis J. Gertsch and Stanley Mulaik described A. stolida, also from Texas.[9] Both species of Astrosoga were transferred to Eucteniza in 2002.[3]

Many taxonomic changes occurred in a 2013 revision by American biologists Jason Bond and Rebecca Godwin. Twelve new species were described, many named after nearby localities, indigenous peoples, or Mexican historic figures. Namesakes of other species include the nightclub Cabo Wabo of Cabo San Lucas, and a character from Battlestar Galactica.[2] Both E. rex and E. stolida were synonymized with E. relata, and the previously described E. atoyacensis (Pickard-Cambridge's Enrico mexicanus) was declared a dubious name: since it was originally described based on a juvenile specimen, it is unclear whether it represents a distinct species. Thus, as of 2013 a total of 14 valid species are recognized.[2]

Species and distribution

The different species of Eucteniza are distinguished by differences in body size, limb proportions, number of tibial spines, and shape of female sperm receptacles, as well as geographic location. Most species are known from male specimens only, but E. rosalia and E. panchovillai are known only from female specimens. Eucteniza species are distributed throughout central and northern Mexico, much of Texas, and the lower part of the Baja Peninsula. Many species are only known from the type locality, the site where the type specimen or specimens were originally collected. The species with the largest range is E. relata, although Bond and Godwin assert the species is actually composed of multiple as-yet-unnamed cryptic species (distinct species erroneously grouped into one) that will require additional morphological or genetic research to distinguish.[2]

Species Geographic range
T denotes only known from type locality.
Source: Bond & Godwin 2013[2] unless otherwise noted
Etymology
Source: Bond & Godwin 2013[2] unless otherwise noted. Eucteniza cabowabo Bond & Godwin, 2013 La Paz and Los Cabos municipalities, Baja California Sur Named for the restaurant Cabo Wabo in Cabo San Lucas, Mexico. Eucteniza caprica Bond & Godwin, 2013 Tamaulipas, MexicoT Named "in reference to the humanoid cylon model Caprica 6", a character from the 2004 Battlestar Galactica television series. Eucteniza chichimeca Bond & Godwin, 2013 Querétaro, MexicoT Named for the indigenous Chichimeca Jonaz people. Eucteniza coylei Bond & Godwin, 2013 Morelos, MexicoT Named for arachnologist Fred Coyle who collected the type specimen. Eucteniza diablo Bond & Godwin, 2013 La Paz Municipality, Baja California Sur Named for Picacho del Diablo, the highest peak in Baja. Eucteniza golondrina Bond & Godwin, 2013 Sótano de las Golondrinas, San Luis Potosí, MexicoT Named for the type locality. Eucteniza hidalgo Bond & Godwin, 2013 Hidalgo, MexicoT Named for the type locality, and for the Spanish word for noble or nobleman. Eucteniza huasteca Bond & Godwin, 2013 La Huasteca Canyon, Nueva Leon, MexicoT Named for the type locality. Eucteniza mexicana Ausserer, 1875 Known only from a specimen from Mexico City and the holotype, whose locality is simply "Mexico".[2][5] Not originally specified.[5] Mexicanus is Latin, meaning Mexican. Eucteniza panchovillai Bond & Godwin, 2013 San Juan del Rio, Durango, MexicoT Named for Mexican revolutionary Pancho Villa, who was born in San Juan del Rio. Eucteniza relata (O. Pickard-Cambridge, 1895) Throughout northern and central Mexico, well into central Texas. Not originally specified.[6] Relatus is Latin, meaning returned or reported. Eucteniza ronnewtoni Bond & Godwin, 2013 Val Verde and Brewster counties, Texas Named for Texas biologist Ronald Newton. Eucteniza rosalia Bond & Godwin, 2013 Mulegé, Baja California Sur, MexicoT Named for the Río de Santa Rosalía (Santa Rosalia River) Eucteniza zapatista Bond & Godwin, 2013 Paso de Cortes, Puebla, MexicoT Named for the Zapatistas, members of Mexico's Liberation Army of the South.

Classification

Eucteniza is the type genus of the family Euctenizidae (the "wafer trapdoor spiders"), a mygalomorph family formerly considered a subfamily of the Cyrtaucheniidae.[10] Within Euctenizidae, Eucteniza is classified in the subfamily Euctenizinae. Earlier comparisons of morphological and behavioral traits suggested the closest relative of Eucteniza is Neoapachella rothi, a forest dwelling spider found in Arizona and New Mexico,[3] while more recent studies, including analyses of DNA similarities, suggest a closer relationship to Entychides, a genus with several species ranging from Arizona to Texas into Mexico.[10][11][12]

Notes

  1. ^ Favila was named after a Gothic duke. The original spelling by O. P.-Cambridge is "Favila",[6][7] but was spelled "Flavila" in Bond & Opell, 2002,[3] and Bond & Godwin, 2013;[2] and "Favilla" in the online World Spider Catalog, version 16, accessed on 19 April 2015
  2. ^ Named after a Gothic ruler

References

  1. ^ Comstock, John Henry (1965). Gertsch, Willis J. (ed.). The Spider Book: A Manual for the Study of the Spiders and Their Near Relatives (Revised ed.). Ithaca, N.Y.: Comstock Publishing Associates. p. 235. ISBN 0-8014-0084-8.
  2. ^ a b c d e f g h i j k l Bond, Jason E.; Godwin, Rachel L. (2013). "Taxonomic revision of the Trapdoor spider genus Eucteniza Ausserer (Araneae, Mygalomorphae, Euctenizidae)". ZooKeys (356): 31–67. doi:10.3897/zookeys.356.6227. PMC 3867109. PMID 24363573.
  3. ^ a b c d e f g h Bond, Jason E.; Opell, Brent D. (2002). "Phylogeny and taxonomy of the genera of south-western North American Euctenizinae trapdoor spiders and their relatives (Araneae: Mygalomorphae, Cyrtaucheniidae)". Zoological Journal of the Linnean Society. 136 (3): 487–534. doi:10.1046/j.1096-3642.2002.00035.x.
  4. ^ a b Gillaspy, James E. (1990). "Pepsis novitia Banks in Texas" (PDF). Sphecos. 19: 21.
  5. ^ a b c Ausserer, Anton (1875). "Zweiter Beitrag zur Kenntniss der Arachniden-Familie der Territelariae Thorell (Mygalidae Autor)". Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien (in German). 25: 125–206.
  6. ^ a b c Pickard-Cambridge, Octavius (1895). Arachnida. Araneida. Biologia Centrali-Americana, Zoology. Vol. 1. London. pp. 145–160.
  7. ^ a b Pickard-Cambridge, Frederick O. (1897). Arachnida - Araneida and Opiliones. Biologia Centrali-Americana, Zoology. Vol. 2. London. pp. 1–40.
  8. ^ Chamberlin, Ralph V. (1940). "A new trap-door spider from Texas". Proceedings of the Biological Society of Washington. 53: 5–6.
  9. ^ Gertsch, Willis J.; Mulaik, Stanley (1940). "The spiders of Texas. I." Bulletin of the American Museum of Natural History. 77: 307–340. hdl:2246/875.
  10. ^ a b Bond, Jason E.; Hendrixson, Brent E.; Hamilton, Chris A.; Hedin, Marshal (2012). "A Reconsideration of the Classification of the Spider Infraorder Mygalomorphae (Arachnida: Araneae) Based on Three Nuclear Genes and Morphology". PLOS ONE. 7 (6): e38753. Bibcode:2012PLoSO...738753B. doi:10.1371/journal.pone.0038753. PMC 3378619. PMID 22723885.
  11. ^ Bond, Jason E.; Hedin, Marshal (2006). "A total evidence assessment of the phylogeny of North American euctenizine trapdoor spiders (Araneae, Mygalomorphae, Cyrtaucheniidae) using Bayesian inference" (PDF). Molecular Phylogenetics and Evolution. 41 (1): 70–85. doi:10.1016/j.ympev.2006.04.026. PMID 16857391.
  12. ^ Hedin, Marshal; Bond, Jason E. (2006). "Molecular phylogenetics of the spider infraorder Mygalomorphae using nuclear rRNA genes (18S and 28S): Conflict and agreement with the current system of classification". Molecular Phylogenetics and Evolution. 41 (2): 454–471. doi:10.1016/j.ympev.2006.05.017. PMID 16815045.
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Eucteniza: Brief Summary

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Eucteniza ( /juːktəˈnaɪzə/ yook-tə-NY-zə) is a genus of trapdoor spiders in the family Euctenizidae containing at least 14 species occurring in Mexico and the southern United States. Species are distinguished by a softened rear portion of the carapace, and males possess large spines on the first two pairs of walking legs that are used to hold females during mating. Like other trapdoor spiders they create burrows with a hinged lid, from which they await passing insects and other arthropods to prey upon. Many species are known from only one or two localities, or from only male specimens. More species are expected to be discovered. Eucteniza is closely related to spiders of the genera Entychides and Neoapachella.

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Description

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Small to large sized trapdoor spiders. Cephalothorax longer than wide, sloping posteriorly, lacking pubescence in most species (Fig. 2). Posterior third of carapace very lightly sclerotized (Figs 2, 23, 24). Thoracic groove intermediate to wide, procurved (Fig. 2) and deep. Eyes not on a tubercle (Fig. 3). AME, PME subequal diameter. Posterior eye row slightly recurved, anterior eye row slightly porcurved (Fig. 2). Caput moderately high (Fig. 3). Carapace of ethanol preserved specimens appears most often reddish-brown, sometimes lighter. The coloration of living spiders tends to be a darker brown, however there is considerable variation in the intensity of coloration. Male coloration in most specimens is dark reddish-brown. Abdominal coloration light to dark brown, sometimes with dark mid dorsal blotch. Sternum wider posteriorly, tapering anteriorly (Fig. 4). Posterior sigilla large and positioned mid-posteriorly nearly contiguous. Anterior margin of sigilla lacks rounded margin. Palpal endites longer than wide with numerous cuspules (Fig. 4). Labium wider than long, with numerous cuspules (Fig. 4). Chelicerae dark brown. Rastellum consists of numerous spines borne on a distinctive mound. Fangs of intermediate length and thickness. Cheliceral promargin with row of very large teeth; retromargin row comprises distinct row of large teeth interspersed with denticles. Apical PLS article short, digitiform. Spinnerets mostly with pumpkiniform spigots with several articulated spigots interspersed on apical and median articles of PLS and the PMS (Bond and Opell 2002, fig. 3E). Two to three large, articulated spigots on apical most aspect of the PLS. PMS article robust. See Bond and Opell (2002) for more detailed descriptions of spigot types. Anterior leg articles slender relative to posterior. Tarsi short and robust (Figs 5–6). Female scopulae long, dense, asymmetrical, extending full length of tarsus, metatarsus and half length of tibia on anterior legs; posterior legs lack distinct scopulae. Male tarsi I and II with short sparse scopulae that are restricted to the ventral surface. Basal palpal tooth and STC I – IV basal tooth elongate and bifid. STC IV with 5 or more teeth. Female anterior legs with very few ventral spines (Fig. 5). Prolateral surface of female patella III and IV covered in numerous thick short spines (Fig. 6). Preening comb on metatarsus IV absent; metatarsus, tarsus IV with ventral spines (Fig. 7). Tarsal trichobothria arranged in a wide band with interspersed setae. Spermathecae generally comprise a simple unbranched bulb that lacks an elongate base. Male mating clasper morphology is distinctive. Tibia legs I & II swollen mid-ventrally in most species, bearing 1-2 large spines; prolateral aspect with a small to large patch of smaller, thickened, short spines. Metatarsus of leg I lacks excavation and spur. Palpal bulb simple, with spherical base, planar distally near origin of embolus. Palpal cymbium lacks dorsal spines (Fig. 11).
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Jason E. Bond, Rebecca L. Godwin
bibliographic citation
Bond J, Godwin R (2013) Taxonomic revision of the Trapdoor spider genus Eucteniza Ausserer (Araneae, Mygalomorphae, Euctenizidae) ZooKeys 356: 31–67
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Rebecca L. Godwin
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Distribution

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Distributed primarily throughout central Mexico and Baja California (Fig. 1) with an extension northward into Texas (United States).
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Jason E. Bond, Rebecca L. Godwin
bibliographic citation
Bond J, Godwin R (2013) Taxonomic revision of the Trapdoor spider genus Eucteniza Ausserer (Araneae, Mygalomorphae, Euctenizidae) ZooKeys 356: 31–67
author
Jason E. Bond
author
Rebecca L. Godwin
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