EUCTENIZINAE, Raven, 1985

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), pp. 487-534 : 509-525

THE EUCTENIZINAE CLADE

Euctenizinae Raven, 1985 (type genus Eucteniza Ausserer )

Diagnosis: Members of this clade can be distinguished from other rastelloid taxa by having an STC with a single claw row (in most cases the basal claw tooth is bifid or elongate), a palpal claw row that is distally offset prolaterally and medially positioned proximally, preening combs (lost in the Eucteniza ), and 2–3 enlarged spigots on the distal most aspect of the PLS.

Higher clade and generic composition: Homostola , Myrmekiaphila , Entychides , + (the California clade) Promyrmekiaphila , Apomastus , Aptostichus , + (the Euctenizoid clade) Neoapachella , Eucteniza . Appendix 2 summarizes the revised south-western Euctenizinae taxonomy.

EUCTENIZA AUSSERER, 1875

( FIGURES 3E , 4D , 7A , 8–10 )

Eucteniza Ausserer, 1875: 149 ; pl. V, figs 8 and 9 (type species by monotypy, Eucteniza mexicana Ausserer , juvenile HOLOTYPE from Mexico, deposited in BMNH, examined). – E. Simon, 1892b: 110. – F.O.P.- Cambridge, 1897: 12; pl. I, fig. 2. – Platnick 2001.

Flavila O.P.- Cambridge, 1895: 156; pl. XIX, fig. 6 (type species by monotypy, Flavila relatus O.P.- Cambridge, female HOLOTYPE from Mexico, Amula in Guerrero, deposited in BMNH, examined). First synonymized by F.O.P.- Cambridge, 1897: 13.

Enrico O.P.- Cambridge, 1895: 157; pl XIX, fig. 8 (type species by monotypy, Enrico mexicanus O.P.-Cambridge, juvenile HOLOTYPE from Mexico, Atoyac in Veracruz, deposited in BMNH, examined). – F.O.P.- Cambridge, 1897: 12; pl I, fig. 7. – E. Simon, 1903b: 899. – Platnick, 2001. – syn. nov. – Eucteniza mexicana (O.P.- Cambridge, 1895) is a junior secondary homonym and is replaced by Eucteniza atoyacensis nom. nov. (etymology: after the type locality, Atoyac, Mexico).

Astrosoga Chamberlin, 1940: 5 (type species by monotypy, Astrosoga rex Chamberlin , male HOLO- TYPE from Kingsville, Texas, deposited in AMNH, examined). – Chamberlin & Ivie, 1945: 556; figs 8–10. – Platnick, 2001. syn. nov.

TAXONOMIC KEY

Males

1. Tibia I with a large mid-ventral megaspine ( Fig. 8A )..................................................................................................2 Tibia I without a large mid-ventral megaspine............................................................................................................3

2(1). Tibia II with a large mid-ventral megaspine; Texas and Mexico.................................................................. Eucteniza Tibia II without a large mid-ventral megaspine; New Mexico and Arizona....................................... Neoapachella

3(1). Cymbium with dorsal apical spines, usually 2–4 ( Fig. 13C ); California, Nevada, Arizona, and Baja California ........................................................................................................................................ Aptostichus Cymbium without dorsal apical spines..........................................................................................................................4

4(3). Spines on mating clasper, tibia I, borne on a low retrolateral apophysis ( Fig. 12A ); Arizona, Texas, New Mexico and Mexico....................................................................................................... Entychides Spines on mating clasper, tibia I, not borne on an apophysis.....................................................................................5

5(4). Thoracic groove straight or procurved, large patch of long thin spines and setae on ventral aspect of tibia I ( Fig. 15A ); Northern California .............................................................................. Promyrmekiaphila Thoracic groove recurved, no distinct spine patch on tibia 1 ( Fig. 17A )................................................... Apomastus Females

1. Thoracic groove straight or recurved.............................................................................................................................2 Thoracic groove procurved..............................................................................................................................................3

2(1). Very distinct comb-like arrangement of setae on ventral aspect of tarsus IV, lacks preening combs on metatarsus IV, New Mexico and Arizona .............................................................................. Neoapachella No distinct comb – like arrangement of setae on ventral aspect of tarsus IV, preening combs on metatarsus III and IV, Southern California .......................................................................................... Apomastus

3(1). Spinule patch on patella IV; Texas and Mexico.............................................................................................. Eucteniza No spinule patch on patella IV.......................................................................................................................................4

4(3). Preening combs on metatarsus IV absent; Arizona, Texas, New Mexico, Mexico...................................... Entychides Preening combs on metatarsus IV present...................................................................................................................5

5(4). Cuspules on palpal endites uniformly distributed across entire face of endite, wide dark bands of coloration on abdomen; Northern California .................................................................... Promyrmekiaphila Endite cuspules concentrated posteriorly, distinct mottled band of abdominal coloration ( Fig. 13D ); California, Nevada, Arizona, and Baja California .................................................................... Aptostichus

Remarks. Based on a comparison of the types of Eucteniza mexicana and Flavila relatus, Cambridge (1897) considered these genera to be congenerics. Our comparisons of cheliceral furrow morphology and leg spination patterns indicate that Enrico and Astrosoga ( Roth (1993) considered Astrosoga to be a likely synonym of Eucteniza ) are likewise subjective synonyms of Eucteniza . Chamberlin and Gertsch probably did not examine the types of either Eucteniza or Flavila because male mating clasper morphology, particularly spination of the ventral aspect of tibia I & II, of Flavila relatus is identical to that of Astrosoga rex and A. stolida .

Diagnosis. Males of this genus can be recognized by the presence of at least one mid-ventral megaspine on the tibia of legs I and II ( Fig. 8A, B ) and the conformation of the palpal bulb ( Fig. 8D ) which has a planarform surface from which the embolus tip arises. Unlike other euctenizine genera, some Eucteniza females have what appear to be a bi-dentate cheliceral furrow and have a distinct rastellum positioned on a moderate to high rastellar mound, whereas other genera lack a distinct rastellar mound and have a single row of promarginal teeth and a small patch of denticles. Additional Eucteniza autapomorphies include an irregularly spaced row of tarsal trichobothria in larger species, a patch of spinules on the prolateral surface of patella IV, and a weakly sclerotized posterior carapace margin ( Fig. 7A ).

Description. Very large trapdoor spiders. Cephalothorax longer than wide, with slight posterior slope, lacks pubescence. Posterior 1/3 of carapace lightly sclerotized ( Fig. 7A ), appearing much lighter in colour. Thoracic groove intermediate to wide, procurved, deep. Eyes not on a tubercle. AME, PME subequal in diameter. Posterior eye row slightly procurved, anterior eye row slightly recurved. Caput moderately high. Carapace of ethanol preserved specimens appears orangered. Coloration of freshly collected female specimens usually darker brown. Male coloration in most specimens is darker reddish – brown. Female abdominal coloration is light brown sometimes with dark middorsal blotch. Male abdominal coloration similar, sometimes uniform brown.

Sternum as in most Euctenizinae , wider posteriorly and tapering anteriorly. Posterior sigilla large, mid-posteriorly positioned, almost contiguous. Anterior margin of sigilla lacks concentric margin. Palpal endites longer than wide and covered in numerous cuspules. Labium wider than long with numerous cus- median articles of PLS and the PMS. Three large articulated spigots on apical most aspect of the PLS ( Fig. 4D ). PMS article robust.

Anterior leg articles slender relative to posterior articles. Tarsi short, robust. Female scopulae long, dense, and asymmetrical, extending full length of tarsus, metatarsus, and half the length of the tibia of the anterior walking legs. Posterior legs lack distinct scopulae. All male tarsi with short dense scopulae that is restricted to ventral surface. Female basal palpal claw tooth and STC I–IV basal tooth bifid. STC IV with few teeth. Female anterior legs with very few spines. Prolateral surface of female patellae III and IV covered in numerous spinules. Metatarsus IV lacks preening comb. Distal ventral aspect of tarsus IV with patch of short spines. Tarsal trichobothrial pattern is wide band typically interspersed among setae. Spermathecae short, unbranched, lacking elongate base ( Fig. 8E ).

Male mating clasper armature distinctive ( Fig. 8A– C ). Patellae elongate, tibiae of legs I & II swollen midventrally, bearing 1–2 large megaspines. Tibia of leg III tends to be slimmer or lacks mid-ventral swollen aspect altogether. Retrolateral aspect of tibia I has a number of short, distally placed spines. Metatarsus lacks excavation and spur. Palpal cymbium lacks spines. Palpal bulb spherical basally and planar distally near origin of embolus ( Fig. 8D ). Palpal femur short with dorsal row of thin spines, tibia short, robust.

pules. Chelicerae dark brown. Rastellum of female consists of numerous spines borne on a distinctive mound. Fangs of intermediate length and thickness. Cheliceral furrow promargin with row of very large teeth. Retromarginal row consists of distinctive row of large teeth interspersed with denticles.

Apical PLS article digitiform, short. Spinnerets mostly with pumpkiniform spigots ( Fig. 3E ), with several articulated spigots interspersed on apical and Natural history. Figure 9 shows typical burrow construction in Eucteniza rex from Webb county Texas collected in 1974 by W. Icenogle. Eucteniza species appear to construct unbranched burrows that are either located on slight inclines or on flat ground. Burrow depths, of those spiders collected by W. Icenogle in 1974 and by us in 1995, ranged from 7 to 25 cm. Burrows are covered with a thin silk plus soil, wafer trapdoor attached by a thin silken hinge. Burrow lining consists of a moderate layer of silk and soil that is thinner than that reported for ctenizid species (e.g. Bond & Coyle, 1995). These spiders place molts and arthropod prey remains at the bottom of the burrow. Prey items collected from burrows at the Webb county locality in 1974 included beetle elytra, ant head capsules, and millipede remains. Many adult and juvenile burrows were found in large aggregations, suggesting dispersal abilities may be minimal. Based on collecting label data, North American (Texas) males appear to disperse during the period between early fall and early winter months (August–January). Dispersal times appear more variable in Mexico, ranging from June through early January.

Distribution. United States, south into Mexico and Baja California ( Fig. 10 ).

Additional type material examined. Astrosoga stolida Gertsch & Mulaik, 1940: 310 ; figs 1-4, 26 (female HOLOTYPE from Austin , Texas, deposited in AMNH, examined).

Material examined. MEXICO: BAJA CALIFORNIA NORTE: Nuevo Leon; La Huasteca Canyon, 3 miles south-west of Santa Catarina, 11 August 1978 (L. Malaret, AMNH), ♂; BAJA CALIFORNIA SUR: La Paz, 8 miles south-east, 1000 ft, 13 October 1968 (E. Sleeper & F. Moore, AMNH) ♀ ♂; Cabo San Lucas, 6 miles east, 10 ft, 13 January 1974 (H. Ridgway, AMNH), ♂; 59 miles north-west of La Paz, 1200 ft, 17 November 1968 (E. Sleeper & F. Moore, AMNH), ♀; La Paz, El Sombrero Trailer Park, 3 July 1968 (M. Bentzien, AMNH), 2 ♀; Mulege, 26 January 1965 (V. Roth, AMNH), ♀; La Paz, 14 July 1970 (R. Funk & C. May, AMNH), ♀; La Paz, 2 miles south, 10 August 1966 (J. Chemask, AMNH), ♀; La Paz city limit, 13 July 1968 (C. Williams et al. AMNH), ♂; 27.3 miles south of Santa Rita, 27 July 1968 (Williams et al. AMNH) ♂; Casas Viejas, 1 mile east, Sierra de la Victoria Mts., 800ft, 28 October 1968 (E. Sleeper & F. Moore, AMNH), ♀; 2 miles south-east of Santa Rita, 1000 ft, 16 November 1968 (E. Sleeper & F. Moore, AMNH), ♀; COAHUILA: Hidalgo, 2–4 August 1973 (T. Kaspar, AMNH), ♂; DURANGO: El Palmito, 10 August 1963 (D. E. Bixler, AMNH), 2 ♀; San Juan del Rio, 1 August 1947 (W. Gertsch, AMNH), 4 ♀; GUERRERO: Taxco, 29 July 1956 (Roth & Gertsch, AMNH), ♀; MORELOS: Tepoztlan, 0.5 miles west, Rt. 115D interchange on road to Ocotepec, 1800 m, 10 June 1982 (F. Coyle, AMNH), ♂; PUEBLA: Puebla, 3000 m, 18 July 1943 (C. Bolivar, AMNH), ♂; QUERÉTARO: Pinal de Amoles, 20 km North, 5–6 July 1971 (Russell & Greer, AMNH), ♂; TAMULIPAS: Antiguo Morelos, 21 June 1963 (J. Beatty, AMNH) ♂; Conrada Castillo, May–June 1980 (P. Sprouse, AMNH), ♀; Tampico 1942 (Ekhomb, AMNH), ♀. UNITED STATES: TEXAS: Atascosa County: Jourdanton, 27 November 1935 (Rutherford, AMNH), ♀; 1–2 September 1936 (C. Rutherford, AMNH), ♀; Bastrop County: Bastrop State Park, 26 March 1958 (D. Hunsacker, AMNH), ♀; Bastrop, 10 miles north-west on Little Sandy Creek, 4 October 1971 (B. Vogel, AMNH), ♂; Bexar County: San Antonio, 15 December 1939 (L. Griffith, AMNH), ♀; Hidalgo County: Edinburg, 1 May 1937 (S. Mulaik, AMNH), ♀; Edinburg, March 1938 (S. Mulaik, AMNH), 2 ♀, 1 juv; Edinburg, 27 February 1939 (S. Mulaik, AMNH), 2♀; Edinburg, 10 miles north-west, 24 December 1949 (AMNH), ♀; North of McCook, 28 November 1937 (D. Mulaik, AMNH), ♀; Kerr County: Raven Ranch, 27 June 1941 (J. McHenry, AMNH), ♀; Kleberg County: Kingsville, October 1940 (AMNH), ♂; Kingsville, 24 November 1969 (AMNH) 2 ♂; Kingsville, November 1947 (J. Cross, AMNH), ♂; Kingsville 1944 (J. Cross, AMNH), 2 ♂; Nueces County: Robstown 14 August 1968 (Richard, AMNH), ♀; San Patricio County: Sinton, ~ 8 miles north-east, 15 October 1959 (H. Laughlin, AMNH), ♂; Starr County: 25 September 1940 (V. Wilder, AMNH), ♀; Travis County: Austin, 5 miles east, 21 January 1957 (W. Blair, AMNH), 1 m; Austin 3 December 1945 (Casteel, AMNH), ♂; Austin Caverns, 3 October 1964 (B. Russel, AMNH), ♂; Val Verde County: Pecos River, on rocks at bridge, 2 September 1968 (J. Brubaker & F. Moore, AMNH), ♂; Ward County: 5 miles north of Monahans, 7 November 1993 (J. Brown, AMNH), ♂; Webb County: Near Highway 83, 1.8 miles North of Junction Highway 35 (15 miles North of Laredo), 800¢, 8 September 1974 (W. Icenogle, AMNH), ♀; Near Highway 83, 1.8 miles North of Junction Highway 35 (15 miles North of Laredo), 800¢, N 27∞46¢ 48.0≤, W 99∞26¢ 57.9≤, 7 August 1997 (J. Bond, JEB – CAS), 2 ♀.

NEOAPACHELLA GEN. NOV.

( FIGURES 3F , 4B , 10 , 11 )

Type species. Neoapachella rothi sp. nov.

Etymology. The generic name, feminine in gender, is in honour of the Apache Indian Nation that has a reservation near the type locality.

Remarks. Roth (1993) was the first to recognize individuals placed in this genus as a distinct taxon and suggested that there were two species distributed in eastern Arizona and western New Mexico. Although there is some variation in male mating clasper morphology that would be indicative of multiple species it is not possible at this time to rule out this variation as intraspecific, thus at present the genus appears to be monotypic.

Diagnosis. The male mating clasper of leg I is very similar to that of Eucteniza , tibia I swollen with a ventral megaspine ( Fig. 11A, B ); however, the tibia of leg III is unmodified and the leg I metatarsus has a slight proximal ventral excavation. In contrast, Eucteniza species have an unmodified metatarsus. The male palpus also has on its retrolateral surface a patch of spines ( Fig. 11C ). Females can be distinguished from all other genera by the presence of a wide, straight thoracic groove and a unique setal patch on the retrolateral surface of the leg IV tarsus.

Description. Small to medium sized spiders. Cephalothorax longer than wide, flat posteriorly, males and females lacking pubescence. Carapace sclerotization uniform across its length. Thoracic groove intermediate to wide, straight in males and females. Carapace of males fringed in stout black setae. Median eyes or all eyes on low tubercle. AME and PME subequal diameter. Both eye rows straight. Caput moderately high. Carapace coloration of alcohol preserved specimens orangish-brown. Female and male abdominal coloration similar, dark brown with dark medial band.

Sternum wider posteriorly, tapering slightly anteriorly. Posterior sigilla small, mid-posteriorly positioned. Anterior margins of sigilla rounded. Palpal endites of female longer than wide with many cuspules uniformly spread across endite surface. Labium wider than long with few cuspules. Labium and palpal endites of male lack cuspules. Chelicerae dark brown. Rastellum of females consists of numerous (5–10) spines in females not borne on a distinctive mound. Fangs long, slender. Cheliceral furrow promargin with row of very large teeth. Retromarginal row consists of a basal patch of few denticles.

Apical PLS article short, digitiform. Spinnerets mostly with small articulated spigots with several large articulated spigots interspersed on apical and median articles of PLS and PMS. Two to three large articulated spigots on apical most aspect of PLS ( Fig. 2B ). PMS article robust.

Anterior leg articles slender relative to posterior articles. Tarsi short, robust. Female scopulae long, dense, asymmetrical, extending full length of tarsus, but no further than metatarsus, scopulae extend no further than tarsus of pedipalp. Posterior legs lack distinct scopulae. Males with short, sparse scopulae restricted to ventral surface of legs I & II. Basal palpal claw tooth, STC I–IV basal tooth elongate and positioned on the median keel but not bifid. STC IV with few teeth. Female anterior legs with very few ventral spines. Prolateral surface of female patella III covered in numerous thick spines. Distal ventral/prolateral aspect of tarsus IV with unique comb-like spine arrangement. Preening combs absent. Spermathecae with long lateral base, does not form secondary spermathecal bulb ( Fig. 11D ).

Male mating clasper like that of Eucteniza ( Fig. 11A, B ), ventral aspect of tibia I swollen, bearing 1–2 megaspines. Metatarsus I with slight proximal ventral to retrolateral excavation. Tibia I with few, small, thick, retrolateral and prolateral spines. Palpal cymbium lacks dorsal spines ( Fig. 11C ). Palpal bulb normal, embolus without serration, tibia with distinct retrolateral distal spine patch. Palpal femur short with dorsal row of thin spines, tibia short and robust.

Natural history. All collecting records of members of this genus have been taken at altitudes above 2100 m. Little is known about the biology of this species. Until recently the only females collected and the only specimens collected without using pitfall traps were those collected by Fredrick Coyle along the banks of the West Fork of the Little Colorado River from shallow burrows he described as ‘ Actinoxia ’ like (he was probably referring to Promyrmekiaphila ). He noted that the burrows were 14–18 cm in length, lined with heavy white silk, and either sealed or covered by a thin wafer trapdoor. More recently the first author and M. Hedin have collected additional female specimens at the same locality recorded by Coyle. Females were found in 10–13 cm deep burrows lined with very heavy silk on a south-western facing slope of the river bank.

Distribution and material examined. Northern/central Arizona and New Mexico ( Fig. 10 ).

NEOAPACHELLA ROTHI SP. NOV.

( FIGURES 10 , 11 )

Types. Male holotype and female paratype from Arizona, Apache County, 1 mile south of Greer on the West Fork of the Colorado River , 8400 ft. (F. Coyle, 29 August 1967), deposited in AMNH .

Etymology. The specific epithet is a patronym in honour of the late Vincent D. Roth. In addition to being a great arachnologist Vince was always helpful and encouraging to new spider systematists. His presence at the arachnological meetings, in Portal, and elsewhere will be sorely missed.

Diagnosis. This species is distinguished in its generic diagnosis.

Male (holotype). Total length (all measurements in mm): 12.36. Cephalothorax length: 6.06, width: 5.06; with setal fringe, lacks pubescence. Carapace of alcohol preserved specimens light orangish/tan–brown, abdominal coloration dark brown, uniform coloration, slightly darker medially. Thoracic groove straight but slightly recurved at margins, width 1.60. Cephalic length 3.68, width 3.24. Ocular quadrangle borne on low tubercle, length: 0.70, width 1.16. Labium length 0.64, width 0.98. Palpal endite length 2.20, width 1.10, lacking cuspules. Sternum length 3.16, width 2.80, sigilla very light and difficult to see. Chelicerae: rastellum row of 2 large spines, promargin with 6 teeth, furrow with proximal sigmoid row of 6 denticles.

Chaetotaxy (spines): Femora: I ~9DM; II 6DM; III 6DM post. 2: 3 (count from right leg), 3PM ant. 3 RM ant.; IV 6–9DM, P/DA with patch of heavy spines, palp

5DA. Patellae: I–II, III ~26DP; IV 0, palp 0. Tibiae: I 5P ant. 1: 3, 2R ant. 2: 4, 2VM 2: 3; II 1PM ant. 1: 4, 2VA, 2VM 2: 3; III 8PM ant. 2: 3, 3VA, 1VM, 2RA; IV too many spines missing for accurate count; palp 9 RM ant. 1: 3 (patch – like). Metatarsi: I VA; II 3 V post. 1: 3, 4VA; III 2PA inf., 4PM, 6DM, 4VA, 1va, 3 V post. 1: 3, 2 V ant. 1: 3; IV 1DA, 3PA, ~7–9 VM. Tarsi: I–II 0, III 2vm ant. 1: 2, 2pm ant. 1: 2; IV large unique spine patch on V/R aspect. Leg article lengths: Femora: I 5.06; II 4.80; III 3.80; IV 4.80; palp 3.48. Patellae: I 2.76; II 2.48; III 2.20; IV 2.56; palp 1.68. Tibiae: I 3.44; II 3.00; III 2.12; IV 3.68; palp 2.36. Metatarsi: I 3.20; II 2.80; III 2.60; IV 3.60. Tarsi: I 2.20; II 2.00; III 2.00; IV 2.32; palp 0.92. Leg coloration uniform, light reddish-brown. Tarsi I–IV not pseudosegmented, straight, robust. Scopulae sparse on tarsus and mid metatarsus I and II. Prolateral surface of tibia Leg I with a few distal robust short spines. Metatarsus I with slight ventral proximal excavation. ( Fig. 11A–C )

Female (paratype). Total length: ~20.71. Cephalothorax length: 8.22, width: 6.81. Carapace dark orangishbrown in ethanol preserved specimens, abdomen dark tannish – brown, lacking distinct markings. Thoracic groove straight, width 2.68. Cephalic length 7.64, width 5.31. Ocular quadrangle length: 0.80, width 1.68. Labium length 1.00, width 1.30, with 6 cuspules. Palpal endite length 3.36, width 1.66, many cuspules spread across entire endite face with dense concentration at posterior – most inner margin. Sternum length 4.80, width 4.00. Sternal sigilla round, moderate in size, slight inward placement. Chelicerae: rastellum group of 3–5 large spines with single row of 2 long spines anterior to fang junction; promargin with 7 teeth, furrow with 5 denticles. Spermathecae moderate length with lateral base, stalk appears heavily sclerotized ( Fig. 11D ).

Chaetotaxy: Femora: I–III, palp 0; IVDA/PA dense spine patch. Patellae: I, II, IV, palp 0; III> 30 R/DA. Tibiae: I 2VM; II 3VM; III 9PM, 3VA, 2 V ant. 1: 2, 3R ant. 1: 3; palp 10VM. Metatarsi: I, II 4VA, 5VM; III 9PM SUP, 3VA. 5vm, 7 RM SUP; IV 3VA, 7 VM. Tarsi: I, II 2–3vm; III 5va IV large comb-like patch of spines on prolateral/ventral aspect; palp 2VM. Leg article lengths: Femora: I 5.73; II 4.81; III 3.82; IV 5.48; palp 4.40. Patellae: I 3.32; II 3.07; III 2.57; IV 3.32; palp 2.49. Tibiae: I 3.49; II 2.82; III 1.99; IV 4.23; palp 2.57. Metatarsi: I 2.49; II 2.66; III 2.49; IV 3.90. Tarsi: I 1.83; II 1.83; III 1.83; IV 1.99 palp 2.49. Leg coloration similar to carapace. Heavy asymmetric scopulae on palp, leg I and II tarsi, metatarsi I and II. 7 palpal claw teeth, 4 P sup. 3 1: 4 M inf. STC teeth: I inner, juxtaposed margin 4, medial face 3; IV promarginal claw 2 teeth on juxtaposed margin, 2 on medial face; retromarginal claw 3 teeth on juxtaposed face, 2 on medial face.

PLS article lengths: apical 0.54; medial 0.84; basal 1.40. Small articulated spigots predominant spigot type with large interspersed articulated spigots. Articulated spigot distributions: apical: 2A, 2M; medial 2M ant. 1: 2; basal 1 A. PMS length 0.80, no articulated spigots evident.

Material examined. UNITED STATES: ARIZONA: Apache County: 1 mile south of Greer on West Fork of the Little Colorado River , 8400¢, 29 August 1967 (F. Coyle, AMNH), 3♀; 11 juv ; NEW MEXICO: Cibola County: Mount Taylor, 11 300 ft, 6 July 1997 (W. O’Keefe, DBR), ♂; Grant County : Meadow Creek , 7000 ft, 31 May 1977 (M. Muma, AMNH), ♂; Meadow Creek , 7000¢, 31 May 1977 (G. Thompson, AMNH), 4 ♂ ♀; Meadow Creek , 7000 ft, 16 June 1977 (M. Muma, AMNH), ♂; Meadow Creek , 7000 ft, 14 July 1976 (M. Muma, AMNH) , 3 juv; San Juan County: Chuska Mountains, South of Toadlena , 8000 ft, 2 June 1997 (M. Hedin, JEB), ♂ .

ENTYCHIDES SIMON, 1888

( FIGURES 4C , 10 , 12 )

Entychides Simon, 1888: 213 (type species Entychides aurantiacus Simon by subsequent designation ( Simon, 1892b) female HOLOTYPE from Mexico, deposited in MNHP, examined). – Simon, 1892b: 109; fig. 107 (spelling emendation to Eutychides ). – F.O.P.- Cambridge, 1897: 11; pl I, fig. 3 (listed as Eutychides aurantiacus , emendation in spelling is attributed to Entychides as a misprint). – Platnick, 2001.

Eutychides Simon, 1892b: 109 . Unjustified emendation, rejected by Platnick, 1989: 62.

Remarks. Simon (1892b) emended the spelling of the genus to Eutychides . This emendation was subsequently retained by a number of authors (e.g. Smith, 1908; Gertsch & Wallace, 1936; Chamberlin, 1937). However, Platnick (1989) considered the subsequent change in spelling by Simon to be unjustified and rejected the emendation.

Smith (1908) considered Actinoxia to be a junior synonym of Entychides . His redescription of A. versicolor is most likely Promyrmekiaphila gertschi . This tentative conclusion is based on locality data (there are records of P. gertschi from Sonoma County, CA but not for Aptostichus ), descriptions of burrow architecture, which are consistent with those we have observed for Promyrmekiaphila , and his illustrations of abdominal coloration (pl. XIII, fig. 9), which indicate a Promyrmekiaphila pattern. Based on cheliceral, STC and male mating clasper differences Chamberlin (1937) revived Actinoxia , thus removing it from Entychides . However, at the same time, he transferred Eutychides arizonicus Gertsch & Wallace to Actinoxia . Diagnosis. Males of this genus can be recognized by the presence of a group of spines that are borne on an apophysis on the distal most prolateral aspect of the tibia of leg I ( Fig. 12A ). Entychides females are similar to those of Eucteniza ; however, they lack the diagnostic spination on patella IV and a short spermathecal bulb without a lateral base, as found in Eucteniza . Additional diagnostic features include very dark carapace and leg coloration, and a very dark brown abdomen without pattern.

Description. Medium sized trapdoor spiders. Cephalothorax longer than wide, sloping slightly posteriorly, lacking pubescence. Carapace sclerotization lighter posteriorly. Thoracic groove intermediate to wide, procurved and deep. Carapace of males fringed in stout black setae. Eyes not on a tubercle, in some male specimens median eyes appear to be on a very low tubercle. AME, PME subequal diameter. Posterior eye row slightly procurved or straight, anterior eye row slightly recurved. Caput moderately high. Carapace coloration dark reddish-brown with males’ coloration similar to that of females. The only exception is a lighter coloured species collected in Texas. Female and male abdominal coloration similar, dark brown without any observable pattern.

Sternum wider posteriorly, tapering anteriorly. In some male specimens sternum almost oval in shape. Posterior sigilla large, mid-posteriorly positioned. Anterior margin of sigilla rounded. Palpal endites longer than wide with many cuspules which are spread across the entire endite surface, but more strongly concentrated posteriorly. Labium subquadrate to wider than long with many cuspules. Chelicerae dark brown. Rastellum of females consists of numerous spines borne on very low, distinctive mound. Fangs long and slender. Promargin of cheliceral furrow with row of very large teeth. Retromarginal row consists of a patch of denticles.

Apical PLS article short, digitiform. Spinnerets mostly with small articulated spigots with several large articulated spigots interspersed on apical and median articles of PLS and PMS. Two to three large articulated spigots on apical-most aspect of the PLS ( Fig. 4C ). PMS article robust.

Anterior leg articles slender relative to posterior. Tarsi short, robust. Female scopulae long, dense, asymmetrical, extending full length of tarsus, no further than metatarsus, no further than tarsus of pedipalp. Posterior legs lack distinct scopulae. Males with short, sparse scopulae restricted to ventral surfaces of legs I & II. Male legs III & IV tarsi appear to have very sparse scopulae. Male tarsi straight, unsegmented. Basal palpal claw tooth and STC I–IV basal tooth elongate, bifid, and positioned on the median keel. Female STC IV with few teeth. Female legs I & II with very few ventral spines. Prolateral surface of female patella III covered in numerous thick spines. Distal ventral aspect of tarsus IV with short, sparse spine patch. Preening combs absent. Spermathecae with long lateral base, that does not form secondary spermathecal bulb ( Fig. 12B ).

Male mating clasper morphology is distinctive ( Fig. 12A ). Metatarsus I with proximal ventral to retrolateral excavation bordered distally by a prominent mound or spur. Tibia I with a few thin spines distributed retrolaterally. Palpal cymbium lacks dorsal spines. Palpal bulb normal and embolus without serration. Palpal femur short with a dorsal row of thin spines, tibia short and robust.

Natural history. There are no records of Entychides burrow construction and very few females of this genus have been collected in the United States. Attempts by the first author and others to collect these females were not productive at localities in the Chiricahua Mountains near Portal and Sabina Canyon near Tucson. Within the Madrean Evergreen Woodland community of south-western Arizona ( Brown, 1982; Bond & Opell, 1997) Entychides males are collected predominantly during the rainy season of late summer.

Distribution. From central Mexico into Texas, New Mexico and Arizona ( Fig. 10 ).

Additional type material examined. Entychides dugesi Simon, 1888: 214 (female HOLOTYPE from Mexico, deposited in MNHP, examined). – Entychides guadalupensis Simon, 1888: 214 (male HOLOTYPE from Guadalupe, Mexico, deposited in MNHP, examined).

Material examined. MEXICO: MORELOS: 1/ 2 mile west of Tepoztlan, 1800 m, 10 June 1982 (F. Coyle, AMNH), ♀, 11 juv; OAXACA: Cueva del Cenpies, Huatla de Jimenez, Rio Iglesia, Dolina, 26 March 1981 (A. Grubbs & S. Zeaman, AMNH), ♀; SAN LUIS POTOSI: Valles, July 1959 (Stuede, AMNH), ♂; SINALOA: 40 miles south of Culiacan, 6 August 1956 (V. Roth & W. Gertsch, AMNH), ♀; SONORA: 8 miles west of Yecara, 4500¢, 8 August 1986 (V. Roth, AMNH), ♀; Sierra de los Ajos, 20 July 1971 (V. Roth, AMNH), ♂; UNITED STATES: ARIZONA: Cochise County: Portal, 26 August 1964 (W. Gertsch, AMNH), ♂; Portal, 1 August 1959 (A. Klots, AMNH), ♂; Portal, 4700 ft., 4 August 1967 (D. Bixler, AMNH), ♂; Portal, 26 August 1964 (R. Hastings & W. Gertsch, AMNH), 2 ♂; Chiricahua Mts., 5400 ft., 15 July 1970 (V. Roth, AMNH), ♂; SWRS, 15 July 1964 (V. Roth, AMNH), ♂; near Portal, 10 September 1991 (J. Rozen, AMNH), 1 ♂; 5 miles south-west of Portal, 20 August 1969 (V. Roth, AMNH), ♂; Gilman Ranch, 11 August 1952 (H. Leech & W. Gertsch, AMNH), ♂; SWRS, September 1984 (AMNH), ♂; 5 miles south-west of Portal, 27 July 1963 (V. Roth, AMNH), ♂; Portal, 25 August 1966 (Rozen, AMNH), ♂; 5 miles west of Portal, 5400 ft, 3 August 1976 (G. Johnson, AMNH), ♂, 1f; Portal, 29 August 1964 (W. Gertsch, AMNH), ♂; 5 miles south-west of Portal, 26 August 1955 (W. Gertsch, AMNH), ♂; Portal, 21 August 1974 (V. Roth, AMNH), ♂; 26 July 1976 (D. Marque, AMNH), ♂; Huachuca Mtns., Carr Canyon, 19 July 1965 (C. Ross, AMNH), ♂; Huachuca Mtns., Garden Canyon, 12 July 1950 (W. Creighton, AMNH), ♂; Pima County: Tucson (O. Bryant, AMNH), ♂; Tucson, 25 November 1946 (G. Morris, AMNH), ♀; Madera Canyon, 17 July 1975 (T. Allen, AMNH), ♂; Madera Canyon, 14 July 1975 (D. Marqua, AMNH), ♂; TEXAS: Bell County: 3 miles south of Belton, 28 December 1941 (AMNH), ♀; Brewster County: Big Bend National Park Basin, 6000 ft., 20 August 1967 (W. Gertsch, AMNH), ♂; Erath County: Stephenville, 25 April 1981 (C. Agnew, AMNH), ♂; Stephenville, 7 April 1982 (C. Agnew, AMNH), ♂; San Patricio County: Sinton, 30 September 1959 (H. Laughlin, AMNH), 3 ♂; about 5 miles north-east of Sinton, 28 October 1959 (H. Laughlin, AMNH) ♂; about 8 miles north-east of Sinton, 15 October 1959 (H. Laughlin, AMNH) 3 ♂; Sinton, 11 August 1959 (H. Lauglin, AMNH), 3 juv; Travis County: Austin, 14 January 1969 (B. Vogel, AMNH), ♂; Austin, 10 December 1968 (B. Vogel, AMNH), ♂; Austin, 11 April 1969 (B. Vogel, AMNH), 1 m; Wichita County: 1 March 1973 (Hicks, AMNH), ♀.

APTOSTICHUS SIMON

( FIGURES 4A , 7C , 13 , 14 )

Aptostichus Simon, 1891a: 317 ( Aptostichus atomarius female LECTOTYPE here designated from CA, San Bernardino; specimen AR4263 deposited in MNHP, examined). – Simon, 1892b: 108. – Simon, 1903b: 901. – Smith, 1908: 220–221; pl. XIII, fig. 32; pl. XIV, figs 30–32. – Platnick, 2001.

Actinoxia Simon, 1891a: 318 (type species by monotypy Actinoxia versicolor Simon juvenile HOLOTYPE from California, deposited in MNHP, examined). – Simon, 1892b: 109. Smith, 1908: 214; pl. XIII, figs 1–19; pl. XIV, figs 1–16; pl. XVI, figs 1, 2 (Smith considered Actinoxia to be a junior synonym of Eutychides ). – Chamberlin, 1937: 9; pl. 2, figs 7–11. – Platnick, 2001. syn. nov.

Nemesoides Chamberlin, 1919: 1–2 ; pl. 1, fig. 2 (type species by monotypy Nemesoides hespera Chamberlin male HOLOTYPE from Claremont, California, deposited in MCZ, examined). – Roth, 1993: D-1. – Platnick, 2001. syn. nov.

Transferred to other genera due to synonymies. Actinoxia arizonica ( Gertsch & Wallace, 1936) is transferred back to Entychides ( Entychides arizonica Gertsch & Wallace , female HOLOTYPE from Sabino Basin, Santa Catalina Mountains, Arizona deposited in AMNH examined). Aptostichus zebra Chamberlin & Ivie, 1935 (female HOLOTYPE from Palo Alto, California deposited in AMNH, examined) is newly transferred to Promyrmekiaphila [ Promyrmekiaphila zebra (Chamberlin & Ivie) comb. nov.].

Remarks. We have designated MNHP specimen AR4263 as the Aptostichus atomarius lectotype because there are two A. atomarius syntypes in the same vial, one of which is an A. simus specimen. Simon’s (1891a) description fits the A. atomarius lectotype specimen, particularly with regards to length measurement. The A. simus specimen is much smaller.

The type specimen for Actinoxia is unequivocally an immature Aptostichus species , considered here to be A. atomarius (suspected to be the case by Simon, 1903b: 900). Actinoxia versicolor Simon, 1891a = Aptostichus atomarius Simon, 1891a syn. nov. This assessment is based primarily on abdominal colour pattern, palpal endite cuspule pattern, and comparisons to juveniles from the broods of Aptostichus atomarius females.

Aptostichus is the most speciose genus of spiders in the Euctenizinae : at present there appears to be at least 30 species. Of the five nominal species only three will likely be retained ( A. atomarius , A. hesperus , and A. simus ). Aptostichus flavipes Petrunkevitch, 1925 will be placed elsewhere (Platnick pers. comm.). Aptostichus stanfordianus Smith, 1908 will likely be considered a junior synonym of A. atomarius in the revision of Aptostichus (see Bond, 1999).

Diagnosis. Males of this genus can be recognized by the presence of three or more spines on the distalmost surface of the palpal cymbium and a number of large, very thick spines on the distal- prolateral aspect of tibia I ( Fig. 13A, E, F ). Entychides males have similar spination; however, their spines are borne on a low apophysis whereas those of Aptostichus are not. Aptostichus females have cuspules on both the labium and palpal endites; labial cuspules are few and restricted to the inner margin. This condition is similar to that for Apomastus , although the latter lacks labial cuspules altogether and also lacks the distinctive Aptostichus abdominal mottled chevron pattern. Additional Aptostichus autapomorphies are spermathecae with the extended lateral base forming what sometimes appears as a secondary bulb ( Fig. 13B, H ) and a distinctive mottled abdominal chevron-like pattern ( Figs 7C , 13D ).

Description. Small to medium sized trapdoor spiders. Cephalothorax longer than wide, sloping posteriorly, moderate pubescence in most species. Carapace sclerotization equal across its length. Thoracic groove intermediate to wide, procurved, deep. In some males thoracic groove only a pit. Carapace of males fringed in stout black setae. Eyes on low tubercle. AME, PME subequal in diameter. Posterior eye row slightly procurved or straight, anterior eye row slightly recurved. Caput moderately high. Carapace of ethanol preserved specimens appears orangish-yellow. Freshly collected coloration tends to be darker brown, however, there is considerable variation coloration intensity. Male coloration in most specimens is darker reddish-brown. Female and male abdominal coloration very distinctive, consisting of light brown or grey background with dark mottled chevron-like pattern ( Figs 9C , 13D ). This pattern is less distinctive in A. simus and other psammophilic species.

Sternum wider posteriorly, sometimes wider than in other euctenizines, tapering anteriorly. Posterior sigilla large, positioned mid-posteriorly, in some species contiguous (e.g. Aptostichus hesperus ). Anterior aspect of sigilla has rounded margin. Female palpal endites longer than wide, with very few cuspules which are restricted to posterior margin. Labium wider than long, with few to moderate number of cuspules. Chelicerae dark brown. Rastellum consists of numerous spines not borne on distinctive mound. Fangs long, slender. Cheliceral furrow promargin with row of very large teeth. Retromarginal row consists of a patch of denticles.

Apical PLS article short, digitiform. Spinnerets mostly with pumpkiniform spigots with several articulated spigots interspersed on apical and median articles of PLS, PMS. Two to three large, articulated spigots on apical most aspect of PLS ( Fig. 4A ). PMS article robust.

Anterior leg articles slender relative to posterior. Tarsi short, robust. Scopulae on females long, dense, asymmetrical, extending full length of tarsus, no further than the metatarsus. Scopulae extend no further than tarsus of pedipalp. Posterior legs lack distinct scopulae. Male tarsi I, II with short sparse scopulae restricted to ventral surface. In some species, male tarsi are slightly bent, elongate and pseudosegmented (e.g. A. simus : Fig. 13E, F ). Female basal palpal claw tooth and STC I–IV basal tooth elongate and positioned on the median keel not bifid. STC IV with 5 or more teeth. Female anterior legs with very few ventral spines. Prolateral surface of female patella III covered in numerous thick spines. Distal ventral aspect of tarsus IV with short, sparse spine patch. Preening combs on distal most retrolateral surface of metatarsus IV. Tarsal trichobothria arranged in zigzag pattern. Spermathecae with elongate base which appears to forms a secondary spermathecal bulb ( Fig. 13B, H ).

Articles of male leg I bear a number of large, thickened spines positioned retrolaterally on distal aspect of tibia. Metatarsus I with proximal ventral to prolateral excavation bordered distally with a low mound. Tibia I with 3–5 elongate spines distributed retrolaterally except in some species which have denser spine patches. Palpal cymbium with four or more dorsal spines. Palpal bulb normal, embolus in some species with serrations. Palpal femur short with dorsal row of thin spines, tibia short and robust in some species (e.g. A. simus ) there is a distinctive prolateral spine patch. ( Fig. 13C, G )

Natural history. More extensive details regarding Aptostichus biology and natural history will be published elsewhere (Bond & Icenogle, in prep.). Burrows ( Fig. 14 ) are lined with a moderate amount of silk and tend to be covered with a very cryptic thin silk-soil trapdoor. Although most species of this genus build branched burrows, some construct burrows without branches. Branches are typically blind tunnels of a slightly smaller diameter that angle towards the surface. All Aptostichus species appear to place prey items and molts in the posteriormost chamber of their burrow. Male dispersal times seem to be correlated with the winter rains, which in California occur late November through January

Distribution. Greatest area of diversification is in Southern California (Los Angeles County southward) extending into Baja California. There are at least two species in Nevada and one in Arizona and Utah. Complete distribution maps are presented in the detailed revision of this genus ( Bond, 1999).

Additional type material examined. Aptostichus simus Chamberlin, 1917 (female HOLOTYPE from San Diego , California, deposited in MCZ, examined).

Material examined. Over 300 specimens of Aptostichus from the AMNH and CAS collections have been examined. Additionally, we have collected and studied over 200 specimens in the field and the lab. Detailed lists of material examined are provided in the revision of this genus (see Bond, 1999).

PROMYRMEKIAPHILA SCHENKEL

( FIGURES 3C , 7B , 15 , 16)

Promyrmekiaphila Schenkel, 1950: 28–32 ; fig. 1 (type species by monotypy, Promyrmekiaphila gertschi Schenkel , female HOLOTYPE from Berkeley , California, deposited in NMB, examined).

Remarks. Spiders placed in this genus have long been informally considered Actinoxia species , a mixed group consisting of Aptostichus , Entychides , and Promyrmekiaphila . However, with the synonymy of Actinoxia with Aptostichus (above), Promyrmekiaphila is unfortunately the only valid name for this group. The Californian species Aptostichus clathratus has the diagnostic features of Promyrmekiaphila females and is thus placed in this genus [ Aptostichus clathratus Simon, 1891a: 318 (female HOLOTYPE from Santa Rosa, California, deposited in USMN, examined) = Promyrmekiaphila clathratus comb. nov.] Subsequent studies of Promyrmekiaphila may find this species to be the senior synonym of P. zebra or P. gertschi .

Diagnosis. Males of this genus can be recognized by the presence of a large patch of spines and long thin setae on the distalmost prolateral and ventral aspect of the tibia of leg I ( Fig. 15B ). In contrast, other euctenizine genera have shorter setae and more definable patches of spines. Promyrmekiaphila females are similar to those of Aptostichus ; however, the cuspule patch on the palpal endites is distributed across the entire endite surface. Additional diagnostic features are a spermatheca with an extended lateral base that does not form a pseudo-secondary bulb as in Aptostichus ( Fig. 15C ) and a distinctive abdominal coloration pattern that consists of wide dark uniform bands that are not mottled ( Fig. 7B ).

Description. Small to medium sized trapdoor spiders. Cephalothorax longer than wide, sloping posteriorly, with moderate pubescence in most species. Carapace equally sclerotized across its length, females lacking pubescence, light pubescence on some males. Thoracic groove intermediate to wide, procurved, deep. Carapace of males fringed in stout black setae. Eyes usually not on tubercle; in some specimens median eyes appear to be on very low tubercle. AME and PME subequal in diameter. Posterior eye row slightly procurved or straight, anterior eye row slightly recurved. Caput moderately high. Carapace of ethanol preserved specimens appears orangish-yellow. Living specimens much darker brown. Coloration of males darker reddish-brown. Female and male abdominal coloration very distinctive in most species, consisting of light brown or grey background with solid dark chevron pattern ( Fig. 7B ).

Sternum wider posteriorly, tapering anteriorly. Posterior sigilla large, mid-posteriorly positioned. Anterior margin of sigilla with rounded margin. Palpal endites longer than wide with very many cuspules which are uniformly spread across the entire endite surface. Labium of females subquadrate to wider than long with no or very few cuspules. Chelicerae dark brown. Rastellum of females consists of numerous spines not borne on a distinctive mound. Fangs long, slender. Cheliceral furrow promargin with row of very large teeth. Retromarginal furrow bears a mesal patch of denticles.

Apical PLS article short, digitiform. Spinnerets mostly with pumpkiniform spigots with several articulated spigots interspersed on apical and median articles of PLS, PMS. Two to three large, articulated spigots on apical most aspect of PLS. PMS article robust.

Anterior leg articles slender relative to posterior. Tarsi short, robust. Female scopulae long, dense to slightly less dense than in other euctenizines, asymmetrical, extending full length of tarsus, no further than metatarsus, pedipalp scopulae extend no further than tarsus. Posterior legs of female lack distinct scopulae. All males with short, sparse scopulae that are restricted to ventral surface of tarsi. Male tarsi straight, not pseudosegmented. Basal palpal claw tooth of female and STC I–IV basal tooth elongate, positioned on median keel. STC IV reduced in size with few teeth. Female anterior legs with very few ventral spines. Prolateral surface of female patella III covered in numerous thick spines. Distal ventral aspect of female tarsus IV with short, sparse spine patch. Rudimentary preening combs on distal most retrolateral surface of female metatarsus IV. Spermathecae with a short lateral base that does not form a secondary spermathecal bulb ( Fig. 15C ).

Male metatarsus I with proximal ventral to prolateral excavation bordered distally by a low mound. Tibia I with a few thin spines distributed retrolaterally. Palpal cymbium lacks dorsal spines. Palpal bulb normal, embolus without serration. Palpal femur short with a dorsal row of thin spines, tibia short and robust. ( Fig. 15A, B )

Natural history. Promyrmekiaphila constructs branched burrows that tend to be located on slight inclines, hillsides, and ravine sides. Burrows reach depths of over 30 cm and are covered with a thin silksoil wafer trapdoor attached with a thin silken hinge. The lining consists of a moderate layer of silk and soil. Branches consist of blind tunnels that angle towards the surface and are slightly smaller in diameter than the main burrow. Side branches tend to have a more constricted opening than those observed for Aptostichus side branches. These spiders place molts and arthropod prey remains in the burrow bottom. Unlike its sister genus Aptostichus , Promyrmekiaphila appears to be restricted to the more mesic climates of central/northern California. Collecting label data show considerable variability in male wandering times, however, the preponderance of males are taken in the early fall through early winter, times consistent with the occurrence of the winter rains in northern California.

Distribution. Central-western and north-western California ( Fig. 16).

Material examined. CALIFORNIA: Alameda County: UC Berkeley Campus, 1 November 1974 (R. Kawin, AMNH), ♂ ; Alameda County, 27 September 1987 (S. Beebe, SCW), ♂ ; Berkeley , 26 March 1946 (J. MacSwain, AMNH), ♀ ; Berkeley , November 1906 ( AMNH) ♂ ; Berkeley , 30 August 1979 (J. Fraser, AMNH), ♂ ; Contra Costa County: Orinda , 12 July 1970 (E. Schlinger, AMNH), ♂ ; Mougan Territory Road, 8.5 miles from Marsh Creek Road, 4 November 1969 (W. Azevedo, SCW), 2 ♂ ; North-west entrance to Briones Regional Park , 7 February 1972 (M. Bentzien, SCW); Contra Costa County , 25 March 1977 (I. Bussey & L. Vincent, AMNH), ♀ ; Alhambea Valley , December 1929 ( AMNH), ♂ , 1 juv; Mt. Diablo , 26 May 1959 (L. Smith & Roschuster, AMNH), 1 juv; Orinda Village, San Pablo Ridge , 15 July 1969 (E. Schlinger, AMNH), ♀ ; with brood of 37 juv; ♂; Marin County: Mill Valley , 20 May 1969 (P. Enconomon, AMNH), ♀ ; Mendocino County: Hopland Field Station , 26 September 1972 (M. Bentzien, AMNH), ♂ ; San Francisco County: San Francisco ( AMNH), ♂ ; San Mateo County: 4 miles west of San Mateo on Highway 5, 18 April 1954 (E. Gilbert & R. Schuster, AMNH), 1 juv; La Honda , Sam MacDonald Park , 17 April 1971 (M. Bentzien, AMNH), ♀ ; San Bruno Mt. , 17 January 1971 (M. Bentzien, AMNH), ♀ ; Highway 84 on way to La Honda, 2nd growth redwood forest, N 37∞23¢ 56.8≤, W 122∞15¢ 34.3≤, 580 ft, 5 May 1997 (J. Bond, JEB- CAS), 3 ♀ ; 1 mile west of Woodside City Limit on Moore Road , N 37∞26¢ 30.8≤, W 122∞14¢ 30.1≤, 380 ft, 4 May 1997 (J. Bond, JEB- CAS), 6 ♀ ; Santa Clara County: Palo Alto , 18 November 1922 (J. Chamberlin, AMNH), ♂ ; Palo Alto , 30 June 1946 (E. Ross, AMNH), ♂ , 1 juv; Palo Alto , August 1931 ( AMNH), 2 ♂ ; San Jose, Alum Rock Park , 23 October 1970 (E. Schlinger et al. AMNH), 3 ♀ , 2 juv; San Jose, Alum Rock Park , 23 October 1970 (E. Schlinger et al. AMNH), ♀ , 6 juv; 5 miles south-west of Cupertino on Monte Bello Road 4 miles west of intersection with Stevens Canyon Road 2000 ft, 10 October 1971 (W. Icenogle, CAS), ♀ ; Santa Cruz County: Ben Lomond , 1600 ft, 2 June 1945 (L. Saylor, AMNH), ♀ ; Ben Lomond , 6 July 1956 (V. Roth & W. Gertsch, AMNH), ♀ , 1 juv; 3 miles north of Soquel , 24 April 1970 (E. Schlinger, SCW), ♀ ; Big Basin Redwood Park , 9 September 1969 (S. & J. Peck, AMNH), ♂ ; Big Basin State Park , 23 December 1953 (V. Roth, AMNH), ♀ ; Shasta County: 3.5 miles south-west of town of Ono on Platina Road , 1000¢, 17 July 1974 (W. Icenogle, CAS), ♀ ; with brood of 25 juv; 1 mile east of South Cow Creek Road on Highway 44 outside of Redding, N 40∞31¢ 52.5≤, W 122∞06¢ 34.4≤, 745 ft, 12 May 1997 (J. Bond, JEB- CAS), 3 ♀ ; Sonoma County: near Santa Rosa , 26 August 1931 (W. Ivie, AMNH), 2 ♀ ; Glen Ellen , 17 August 1959 (W. Gertsch & V. Roth, AMNH), ♀ ♂ ; 1 mile south of Trenton , 15 May 1957 (R. Schuster, AMNH), ♀ , 1 juv; Armstrong Redwoods State Park , 10 August 1967 (F. Coyle, AMNH), ♀ ; Stanislaus County : Del Puerto Canyon , 9 April 1971 (R. Coville, SCW), ♂ .

APOMASTUS GEN. NOV.

( FIGURES 17–19 )

Type species. Apomastus schlingeri sp. nov.

Etymology. The generic name is the latinized form of the Greek apomastos meaning ‘not provided with a lid’. This refers to the absence of a trapdoor on burrows constructed by members of this genus.

Remarks. Previously members of this new genus were informally considered to be members of Aptostichus (W. Gertsch in litt.). We describe in this paper only the type species, Apomastus schlingeri ; however, at least one other species is known.

Diagnosis. Males of this genus can be recognized by the presence of a recurved thoracic groove and the absence of a proximal–ventral metatarsal excavation and distinctive spination on leg I ( Fig. 17B, C ). Females are similar to those of Aptostichus , although they have a straight thoracic groove and uniform, dark brown abdominal coloration. In contrast, the thoracic groove of most Aptostichus species is recurved and the abdominal coloration is lighter with a distinctive mottled chevron colour pattern. All known species of this genus do not cover their burrows with trapdoors, whereas it appears that all other euctenizines do.

Description. Medium sized spiders. Cephalothorax longer than wide, sloping slightly posteriorly, females lacking pubescence, males with light to moderate pubescence. Carapace sclerotization equal across its length. Thoracic groove intermediate to wide, straight in females, recurved in males. Carapace of males fringed in stout black setae. Median eyes or all eyes on a tubercle. AME, PME subequal in diameter. Posterior eye row slightly procurved or straight, anterior eye row slightly recurved. Caput moderately high. Carapace coloration of both sexes brown, orangish-brown in alcohol preserved specimens. Female and male abdominal coloration similar - dark brown lacking any observable pattern.

Sternum wider posteriorly, tapering anteriorly. Posterior sigilla small, mid-posteriorly positioned. Anterior margin of sigilla with rounded margin. Palpal endites longer than wide, appearing almost subquadrate in A. schlingeri , with many cuspules which are concentrated in a tight group posteriorly as in Aptostichus . Labium wider than long, lacking cuspules. Chelicerae dark brown. Rastellum of female consists of numerous spines not borne on a distinctive mound. Fangs long and slender. Cheliceral furrow promargin with row of very large teeth. Retromarginal row bears a patch of denticles.

Apical PLS article short, digitiform. Spinnerets mostly with small pumpkiniform spigots with several large articulated spigots interspersed on apical and median articles of PLS and the PMS. Two to three large articulated spigots on apical most aspect of PLS. PMS article robust.

Anterior leg articles slender relative to posterior. Tarsi short, robust. Female scopulae long, dense, asymmetrical, extending full length of tarsus, no further than the metatarsus. Scopulae extend no further than the tarsus of pedipalp. Posterior legs lack distinct scopulae. Males with short, sparse scopulae restricted to ventral surface of legs I & II. Male tarsi long, slen- der, slightly curved and pseudosegmented. Female basal palpal claw tooth and STC I–IV basal tooth elongate, bifid, and positioned on median keel, STC IV with few teeth. Female anterior legs with very few ventral spines. Prolateral surface of female patella III covered in numerous thick spines. Distal ventral aspect of tarsus IV with short, sparse spine patch. Preening combs on female metatarsus III, IV, sometimes II. Spermathecae with long lateral base, does not form a secondary spermathecal bulb ( Fig. 17D ).

Male metatarsus I without proximal ventral to retrolateral excavation. Tibia I with few to many thin prolateral spines. Palpal cymbium lacks dorsal spines. Palpal bulb normal, embolus without serration. Palpal femur short with dorsal row of thin spines, tibia of moderate length and robust. ( Fig. 17A )

Natural history: Figure 18 shows burrow entrance construction in this enigmatic group of spiders. Apomastus species are unusual because they do not cover their burrow with a trapdoor. Their retreats consist of a burrow lined with heavy silk that extends 10–20 cm back into the substrate. The burrow opening consists of a silken tube that extends a few centimeters from the substrate to form a short to very long collar. Individuals often incorporate soil and vegetative material into the burrow extension, effectively extending the prey detection radius of the burrow. These spiders appear to prefer the north facing slopes of stream fed ravines along the coastal ranges of southern California. The exception is a Riverside County population of Apomastus sp. that is found in a more arid chaparral habitat. Apomastus species place prey remains and moults in their burrow bottoms. These spiders are also unusual because they retain overlapping brood generations in the burrow. Wendell Icenogle and the first author on numerous occasions have collected A. schlingeri females with broods that comprised a full brood from the present year and two or three larger juveniles presumably held over from the previous year.

Distribution and material examined. California counties of Los Angeles, Orange, San Bernardino and Riverside ( Fig. 19). Material examined is listed below.

Figures

Figure 2. Reanalysis of Goloboff’s (1993) data set of 71 morphological characters scored for 42 mygalomorph taxa using implied weights (k = 7; note that the topology is quite sensitive to concavity function constant). This is a strict consensus of 27 most parsimonious trees (236 steps, CI = 0.39, RI = 0.73). Filled circles indicate nodes that differ from Goloboff ’s original analysis, cyrtaucheniid taxa are in grey shaded box. A summary and a brief explanation of the characters and their scorings are presented in Appendix 1.

Figure 3. Posterior lateral spinneret spigot types for select rastelloid taxa (scale bars = 0.1 mm). A, Kiama lachrymoides. B, Cyrtauchenius luridus Simon, 1881. C, Promyrmekiaphila gertschi. D, Acontius sp. E, Eucteniza rex. F, Neoapachella rothi. G, Rhytidicolus sp. H, Myrmekiaphila torreya. I, Ummidia sp.

Figure 4. Silk spigots on apicalmost article of posterior lateral spinnerets for select euctenizine taxa (scale bars = 0.1 mm). A, Aptostichus hesperus. B, Neoapachella rothi. C, Entychides arizonica. D, Eucteniza rex.

Figure 7. Colour photographs of live Euctenizinae species. A, Eucteniza rex from Laredo, Texas. B, Aptostichus atomarius from Riverside County, California. C, Promyrmekiaphila gertschi from San Mateo County, California.

Figure 8. Eucteniza rex Chamberlin male holotype (A–D) and female paratypes (E). A, retrolateral aspect of leg I. B, prolateral aspect of leg I. C, retrolateral aspect of leg II. D, retrolateral aspect of pedipalp. E, spermathecal receptula.

Figure 9. Eucteniza rex burrow from Laredo, Texas excavated by W. Icenogle and bisected. A, lateral view with trapdoor open. B, top view with trapdoor closed.

Figure 10. North American known locality map for Neoapachella, Entychides, and Eucteniza.

Figure 11. Neoapachella rothi sp. nov. male HOLOTYPE (A–C) female paratype (D). A, leg I, retrolateral aspect. B, leg I, prolateral aspect. C, pedipalp, retrolateral aspect. D, spermathecal receptula.

Figure 12. Entychides arizonica from Cochise County, Arizona, 5 miles south-west of Portal (SWRS) in AMNH. A, male leg I, retrolateral aspect. B, spermathecal receptula.

Figure 13. Aptostichus atomarius from Los Angeles County, California, Chatsworth in CAS (A–D) and Aptostichus simus, male from Los Angeles County, California, Playa del Rey State Beach and female from the type locality San Diego County, Silverstrand State Beach (E–I). A, male leg I, retrolateral aspect. B, spermathecal receptula. C, cymbium and palpal bulb, retrolateral aspect. D, abdomen. E, male leg I, retrolateral aspect. F, male leg I, prolateral aspect. G, pedipalp, retrolateral aspect. H, spermathecal receptula. I, sternum and palpal endites.

Figure 14. Aptostichus sp. burrow from Riverside County, Winchester, California. A, burrow closed with arrow indicating its location. B, same burrow opened.

Figure 15. Promyrmekiaphila gertschi from Ben Lomond, Santa Cruz County, California. A, pedipalp, retrolateral aspect. B, male leg I, retrolateral aspect. C, spermathecal receptula.

Figure 17. Apomastus schlingeri sp. nov. HOLOTYPE (A–C) and female paratype (D). A, pedipalp, retrolateral aspect. B, leg I, retrolateral aspect. C, leg I, prolateral aspect. D, spermathecal receptula.

Figure 18. Apomastus sp. burrow opening, from Orange County, California.