Creaserinus clausus, Johnson & Stern & Crandall, 2021

Johnson, Daniel P., Stern, David B. & Crandall, Keith A., 2021, The genus Creaserinus Hobbs, 1973 (Decapoda: Cambaridae) in Texas, Zootaxa 5017 (1), pp. 1-84 : 42-52

Creaserinus clausus n. sp.

Figs. 3–5, 6b, h, 7a, f–i, 9–12, 18–24, Tables 1 , 2, 7–10, 22–24

Cambarus hedgpethi .— Penn & Hobbs, 1958:454, 462, 465, 467, 471, 473, 476–478, Figs. 11, 28, 42, 55 [in part]. Fallicambarus hedgpethi .— Hobbs, 1969:111, Fig. 20f [in part; by implication].

Fallicambarus (Creaserinus) fodiens .— Hobbs & Robison, 1989:672, Figs. 9h, i, k, s [in part].— Hobbs, 1990:575 [in part]. Fallicambarus (Creaserinus) fodiens “TX-2”.— Ainscough et al., 2013:8, Figs. 3, 4.

Creaserinus fodiens .— Crandall & De Grave, 2017:627 [in part].

Diagnosis. Adults with rostrum devoid of marginal spines. Areola obliterated along part of its length. Antennal scale more than twice as long as broad. Cheliped with sufflamen. Lateral margin of chela costate, dorsal surface lacking tubercles on lateral half, ventrolateral surface lacking arched row of prominent setiferous punctations; opposable margin of dactyl with distinct excision in basal half; mesial margin bearing at least 2 rows of tubercles, those of main row extending from base at least two-thirds length of finger. Length of carpus distinctly less than width of palm of chela. Ventral surface of merus with mesial and lateral rows of tubercles. Mesial surface of chela of 2 nd pereiopod bearing prominent tufts of plumose setae. Hooks on ischia of 3 rd pereiopod only. Boss on coxa of 4 th pereiopod, rounded, slightly compressed, and scarcely protruding ventrally. Gonopod of form I male lacking proximomesial spur and terminating in two distinct parts (mesial process and central projection). Central projection corneous, bladelike, lacking subapical notch, recurved 160–190° to axis of shaft, with distal part directed caudoproximally to cephaloproximally with tip never crossing central projection of corresponding gonopod, and base not inclined laterally, length 37.0–45.3% (x = 40.1, s = 1.7, n = 49) of total gonopod length. Mesial process noncorneous, recurved 105–185° to axis of shaft, proximal half inflated. Proximal gap between processes absent or very reduced. Uropodal endopod with distolateral spine; distomedian spine premarginal. Telson divided with spine on anterolateral flank of suture. Function 6 positive when b refers to Creaserinus clausus n. sp. and a refers to any of other five Texas species (refer to Tables 2, 7–10). Sex ratio of juvenile population (<16 mm CL) not strongly skewed female.

Holotypic male, form I. Eyes pigmented and with faceted cornea, but small. Body subcylindrical, very weakly depressed (Figs. 18a, b). Abdomen slightly narrower that cephalothorax (9.8 and 10.9 mm, respectively). Greatest width of carapace near one-third length of areola from cervical groove, where subequal to height (10.9 and 10.6 mm, respectively). Areola obliterated over most of its length and comprising 37.0% of CL and 44.4% of POCL. Rostrum with slender, slightly concave margins converging from base to modestly delimited acumen; apex corneous, distinctly upturned, and reaching midlength of ultimate podomere of antennular peduncle; dorsal surface concave with submarginal rows of punctations and others widely scattered in between. Subrostral ridge weak but visible in dorsal aspect to base of acumen. Postorbital ridge slender, well defined, and relatively abruptly terminating anteriorly just posterior to margin of orbit. Cervical spine absent. Weak branchiostegal tubercle present. Suborbital angle obtuse and very weak. Carapace punctate dorsally and weakly granulate-punctate laterally.

Abdomen subequal in length to carapace (22.9 and 22.7 mm, respectively); pleura small with subrounded margins; pleuron of 2 nd segment clearly overlapping that of 1 st. Telson divided and deeply incised laterally; caudolateral angle of cephalic section with pair of spines, more mesial one movable.

Cephalic lobe of epistome subtriangular, bearing a few punctations, margins slightly elevated and smooth ventrally; main body with weak fovea. Ventral surface of proximal podomere of antennule with spine slightly distal to midlength. Antennal peduncle without spines, flagellum reaching cephalic margin of 3 rd tergum. Antennal scale (Fig. 19f) reaching tip of acumen and ultimate podomere of antennular peduncle; lamella broadly rounded distomesially, broadest distal to midlength and broader than thickened lateral part. Mandible with cephalic noncorneous, subtuberculiform molar process and caudal corneous, irregular molar process. Ventral surface of ischium of 3 rd maxilliped with medium-length setae laterally and long setae mesially; ventral surface of basis with long setae somewhat obscuring proximal part of ischium.

FIG. 18. Creaserinus clausus n. sp.: (a) lateral and (b) dorsal views of holotype. Specimen exhibits broad-striped phase.

Right chela of cheliped (Figs. 19a–d) 2.2 times as long as broad, strongly depressed; width of palm 1.8 times length of mesial margin. Latter bearing main row of 7 tubercles flanked dorsolaterally by three rows of 6, 2, and 1 (left with 6, 1, and 0) slightly smaller tubercles and ventrolaterally by row of 4 (left with 4) distinctly smaller tubercles. Dorsal surface of palm and fingers bearing setiferous punctations, those of dactyl and fixed finger much more distinct. Lateral margin of propodus costate along distal 85%, but broadly rounded proximally. Ventral surface punctate, with punctations more prominent on fingers, single tubercle opposite base of dactyl. Long conspicuous setae present on proximal half of opposable margin of fixed finger. Dorsal and ventral surfaces of both fingers with well-defined ridges flanked by punctations. Opposable margin of fixed finger with row of 3 tubercles (3 rd from base largest) along proximal third and 2 more projecting from lower level in distal half; single row of minute denticles extending from 3 rd tubercle to corneous tip of finger. Opposable margin of dactyl with prominent excision in proximal third; 2 tubercles borne within excision, a larger one at its distal extremity, and 4 distal to excision; single row of minute denticles extending from 2 nd distalmost tubercle to corneous tip of finger. Mesial margin of dactyl with main row of 9 tubercles bordered dorsolaterally by row of 5 (left with 6; left bordered by additional row of 2 very small tubercles).

FIG. 19. Creaserinus clausus n. sp.: (a) dorsal, (b) ventral, (c) lateral, and (d) mesial views of chela; (e) tail; (f) antennal scale; (g) ventral view of merus, (h) proximal podomeres of pereiopods showing hook; and (i) annulus ventralis. a–h from holotype, i from allotype.

Carpus of cheliped 1.5 times as long as broad and 1.4 times as long as palm mesial margin; dorsal surface with longitudinal median trough, flanked with punctations; mesial surface minutely tuberculate with large spine on distal margin; lateral and ventral surfaces punctate; ventral surface with large distomedian spine and small tubercle at ventrolateral condyle. Merus with 2 squamous tubercles near dorsodistal extremity; mesial and lateral surfaces sparsely punctate; ventral surface (Fig. 19g) with mesial row of 13 tubercles (left with 12) and lateral row of 4 (left with 2). Ischium punctate and lacking tubercles ventromesially (left with 1).

Second pereiopod bearing conspicuous long setae on dorsal and ventral margins of chela and carpus, and proximal half of ventral margin and dorsodistal extremity of merus, and mats of well-developed plumose setae on mesial faces of palm and carpus.

Hook (Fig. 19h) on ischium of 3 rd pereiopod simple, reaching level of basioischial articulation, and not opposed by tubercle on corresponding basis. Coxa of 4 th pereiopod as in “Diagnosis.” Coxa of 5 th pereiopod lacking boss.

Gonopod (Figs. 20a, c, e) reaching coxa of 3 rd pereiopod when abdomen flexed and substantially obscured by setae extending from surrounding sternum; shaft of appendage straight, otherwise as in “Diagnosis.” Proximal podomere of uropod with both lobes bearing single small spines; endopod with distolateral spine and distinctly premarginal distomedian spine.

Allotypic female. Differing from holotype in nonsecondary sexual characters as follows: abdomen width equal to that of carapace; branchiostegal spine absent; cephalic lobe of epistome subpentagonal and lacking punctations; flagellum reaching cephalic margin of 1 st tergum; mesial margin of palm with 9 tubercles dorsolateral to and 3 ventrolateral to main row of 7; single tubercle on ventral surface of palm opposite dactyl absent (present on left chela); opposable margin of fixed finger with row of 2 tubercles (2 nd from base largest) along proximal third and 3 more projecting from distal half; opposable margin of dactyl with row of 5 tubercles distal to excision; mesial margin of dactyl with main row of 13 tubercles bordered dorsolaterally by row of 7; carpus of cheliped 1.4 times as long as broad and 1.5 times as long as palm mesial margin; merus of cheliped with 2 squamous tubercles near dorsodistal extremity, ventral surface with mesial row of 15 tubercles (left with 15) and lateral row of 5 (left with 5); ischium of cheliped lacking tubercles ventromesially; second pereiopod lacking conspicuous plumose setae on mesial faces of palm and carpus.

Annulus ventralis (Fig. 19i) 1.4 times as broad as long, cephalically fused to sternum. Caudal and lateral margins elevated (ventrally) and surrounding shallow, cephalic, caudodextrally-directed trough. Short sinus originating at caudomedian border, curving sinistrally 30°, then dextrally 180°, before terminating in small caudomedian fossa. Postannular sclerite subtriangular and less than half as long and less than half as wide as annulus. First pleopod present.

FIG. 20. Creaserinus clausus n. sp.: (a) mesial, (c) ventral, and (e) lateral views of gonopod of holotype; and (b) mesial and (d) lateral views of gonopod of morphotype.

Morphotypic male, form II. Differing from holotype in following respects: areola comprising 39.4% of CL and 46.5% of POCL; cephalic lobe of epistome subsemicircular, lacking punctations; flagellum reaching cephalic margin of 6 th tergum; right chela of cheliped 2.1 times as long as broad; width of palm 1.7 times length of mesial margin; latter bearing main row of 7 tubercles flanked dorsolaterally by row of 6 slightly smaller tubercles and ventrolaterally by irregular group of 7 distinctly smaller tubercles; ventral surface of propodus with a few weak tubercles near dactylar articulation; opposable margin of fixed finger with row of 3 tubercles along proximal third, 2 nd and 3 rd from base much larger than 1 st; opposable margin of dactyl with 3 tubercles distal to excision; single row of minute denticles extending from 3 rd distalmost tubercle to corneous tip of finger; mesial margin of dactyl with main row of 11 tubercles bordered dorsolaterally by row of 7 tubercles; carpus of cheliped 1.3 times as long as broad, mesial surface with small spine proximal to one on distal margin; merus of cheliped with 3 small squamous tubercles near dorsodistal extremity, ventral surface with mesial row of 13 tubercles (left with 12) and lateral row of 5 (left with 7); ischium of cheliped with ventromesial row of 3 tubercles (left with 4); hook on ischium of 3 rd pereiopod and coxa of 4 th pereiopod both reduced; gonopod (Figs. 20b, d) with noncorneous central projection, both processes more inflated.

FIG. 21. Creaserinus clausus n. sp.: mesial views of gonopods of form I male paratypes. Each photo annotated with name of county where collected. Part 1 of 2.

Type locality. Roadside ditch 0.5 km (0.3 mi) south of junction with Farm to Market Road 529 on Farm to Market Road 2855, Waller County, Texas (29.87033, -95.89055). Water presence appears to be very temporary. Vascular plant species noted at the site include Eleocharis sp. , Paspalum urvillei , Setaria geniculata , Acmella oppositifolia , Polygonum hydropiperoides , Mimosa strigillosa , and Ludwigia sp. See Fig. 23.

Disposition of types. The holotype, allotype, and morphotype (nos. 1640952, 1640953, and 1640954, respectively) are deposited in the National Museum of Natural History , Smithsonian Institution. All paratypes remain in DPJ’s collection .

FIG. 22. Creaserinus clausus n. sp.: mesial views of gonopods of form I male paratypes. Each photo annotated with name of county where collected. Those annotated with an asterisk paratopotypes. Part 2 of 2.

Size. The largest specimen examined is a female from Fort Bend County with a CL of 32.5 mm and a POCL of 28.0 mm. Corresponding lengths for the smallest ovigerous female are 22.1 and 18.5 mm. Carapace lengths for form I males range from 19.5 to 32.0 (x = 24.1) mm (n = 345).

Range and specimens examined. Creaserinus clausus n. sp. has been found at 382 sites in 36 counties in Texas (Fig. 10), of which 312 (81.7%) are represented by at least one form I male and 35 (9.1%) are represented in the molecular phylogeny. Based on Fig. 9s of Hobbs & Robison (1989:675), this species is also present in Louisiana.

Anderson: 32.0415, 95.4962, E, I, 3/30/07; 31.8915, 95.4329, E, I; 31.8719, 95.6102, I, 5/9/10; 31.6489, 95.4137, I, 5/3/14; 31.8193, 95.4810, I; 31.8233, 95.5524, I; 31.9453, 95.5346, I, 5/4/14; 31.9743, 95.5735; 31.8882, 95.8749, I; 31.8559, 95.8907, I, DJ474. Angelina : 31.2418, 94.7933, I, USNM no. 219180; 31.2536, 94.8080, I, USNM no. 219182; 31.2640, 94.8276, I, USNM no. 219188; 31.1801, 94.5166, I, 3/2/08, 4/26/15; 31.3723, 94.8126, I, 3/2/08, 4/25/15; 31.1633, 94.7327, I, 2/15/13; 31.1695, 94.4315, I, 3/27/15; 31.1794, 94.3772, I, 3/29/15; 31.2179, 94.4119, I; 31.2618, 94.4272, I; 31.2800, 94.4946; 31.2816, 94.5541, I; 31.4276, 94.9039, I, 4/25/15; 31.4094, 94.6190, I, 4/26/15; 31.3843, 94.5490, I; 31.3393, 94.5337, I; 31.0992, 94.4769, I; 31.1206, 94.5493, I; 31.1779, 94.5780; 31.1666, 94.6336, I; 31.2506, 94.7014, I, DJ672; 31.3948, 94.9153; 31.2276, 94.7690, I, 5/17/15; 31.1709, 94.7887, I. Cass: 33.0008, 94.5180, I, 4/6/14; 32.9209, 94.2614, I, 4/10/15; 33.0501, 94.6021, I, 4/6/14, DJ386. Chambers : 29.6804, 94.6747, I, 9/22/13; 29.6616, 94.6256, I. Cherokee: 32.0345, 95.4237, E, 4/27/07; 31.6566, 95.0309, I, 3/30/13; 31.4803, 95.0121, I; 31.6331, 95.0945, I; 31.7666, 95.1505, I, 3/31/13; 31.8501, 95.0113, I; 31.8886, 95.0860, I; 31.9102, 95.1271, I; 31.9758, 95.1691, I; 32.0264, 95.1725, I; 32.1042, 95.1669, I; 31.6362, 95.2752, I, 5/3/14, DJ412; 31.6369, 95.2364, I; 31.6724, 95.2345, I; 31.7196, 95.2363, I; 31.7064, 95.3042, I; 31.7839, 95.2174, I. Fort Bend : 29.6912, 95.8883, I, 4/13/09; 29.6917, 95.8820, I; 29.7689, 95.8399, I; 29.6956, 95.8351, I, 4/13/09, 5/29/15; 29.6998, 95.7873, I, 5/29/15; 29.7510, 95.7754, I. Freestone: 31.6129, 95.9711, I, 5/31/14, DJ448. Gregg : 32.4811, 94.9532, I, 4/12/15. Grimes : 30.3540, 95.8091, I, 5/18/14, DJ488; 30.3243, 95.9145; 30.2387, 95.9029, I; 30.5428, 95.8346, DJ398. Hardin : 30.4642, 94.6721, E, 3/1/07; 30.1268, 94.4041, I, 12/8/07; 30.3217, 94.5622, I, 2/2/08; 30.4005, 94.4727; 30.4805, 94.4628; 30.4659, 94.1808; 30.5099, 94.1783, I; 30.2173, 94.3293; 30.2974, 94.4592, I, 2/13/10, DJ130; 30.2170, 94.2941, I, 2/2/13; 30.2163, 94.2710, I; 30.2274, 94.1942, I, 8/18/16; 30.2266, 94.1941, I, 12/4/16. Harris : 29.8958, 95.2019, I, 12/28/08; 29.9731, 95.1717, I; 30.0845, 95.1110, I; 30.1603, 95.0969, I; 29.9343, 95.3786, I, 7/14/12; 29.8215, 95.8066, I, 10/14/13; 29.8895, 95.8070, I; 29.8796, 95.7412, I; 29.8130, 95.6181, I, 3/1/14; 29.9437, 95.7129, I; 30.0694, 95.8398, I, DJ359; 29.7901, 95.7626, I, 5/29/15; 29.7737, 95.4426, 5/30/15; 29.6266, 95.2632, I, 5/31/15; 29.5899, 95.0348, I; 29.8605, 95.2616, I; 29.9055, 95.2495, I; 29.9926, 95.0921, I, 8/14/16; 29.9759, 95.0885. Harrison: 32.6716, 94.4233, I, 3/23/08; 32.7352, 94.2871, I, 4/6/14; 32.6206, 94.5065, I, 4/11/15; 32.6271, 94.5777, I; 32.6340, 94.6726, I. Henderson: 32.3130, 95.4596, I, 6/12/14; 32.3058, 95.6080, I; 32.3116, 95.6715, I, DJ477; 32.3354, 95.7453, I; 32.2171, 95.6612, I. Houston: 31.5748, 95.1695, I, 3/30/08; 31.4350, 95.3191, I; 31.1328, 95.6733, I; 31.1538, 95.5759, I; 31.1409, 95.4883, I, 3/30/08, 4/24/15; 31.2133, 95.4825, I, 5/9/09; 31.0722, 95.4260, I; 31.0042, 95.5784, I, 5/29/14, DJ485; 31.0960, 95.4954, I, 4/24/15; 31.1407, 95.3971, I; 31.2626, 95.3701, I, DJ671; 31.3587, 95.1721, I, 5/16/15; 31.2864, 95.5860, I; 31.2539, 95.6269, I; 31.3030, 95.5150, I; 31.3285, 95.3450, I; 31.3554, 95.2442, I; 31.3298, 95.1221, I; 31.3070, 95.1039, I; 31.3389, 95.2884, I, 5/20/16; 31.3234, 95.2290, I; 31.2650, 95.2532; 31.2602, 95.2077; 31.2929, 95.2046; 31.2956, 95.1727, I; 31.2939, 95.1372, I; 31.3271, 95.0762; 31.3535, 95.0809, I; 31.3842, 95.0742; 31.2157, 95.5447, I, 5/21/16; 31.2026, 95.5484, I; 31.1669, 95.4407, I; 31.1912, 95.4289, I; 31.1769, 95.3396, I; 31.2513, 95.1530. Jasper: 30.4700, 93.9919, I, 1/19/08; 30.5892, 93.9575; 30.6756, 93.9318; 30.6828, 94.0762; 31.0430, 94.1586; 30.2813, 93.9072, 2/23/14; 30.3359, 93.9245, I; 30.8455, 94.1089, I, 3/21/14; 30.8034, 94.1269, I; 30.7724, 94.0474, I; 30.8879, 93.9902, I, 3/22/14; 30.8726, 94.0170, I; 30.7976, 94.0071, I; 30.9007, 93.8833, I; 30.2804, 94.0417, I, 3/27/15, 8/18/16; 30.9753, 94.1293, I, 3/27/15. Jefferson : 30.0516, 94.3080, 9/23/13; 30.0596, 94.2466, I; 30.0603, 94.2411, I; 30.1329, 94.2364, I; 30.1329, 94.2409, I; 30.0371, 94.1894, I; 29.9810, 94.2053, I; 29.9308, 94.2629, I; 29.9478, 94.3344, I; 29.9503, 94.0396, I, 3/27/15; 29.8263, 94.2351, I; 29.9174, 94.1822, I. Leon: 31.5713, 95.8864, I, 5/4/14; 31.2906, 95.8042, I, 5/30/14, DJ478; 31.4016, 95.7667, I; 31.4470, 95.8702, I; 31.3096, 96.0031, I, 5/31/14; 31.1817, 95.8414, I, 4/24/15. Liberty : 30.3061, 94.7490, 2/15/08; 30.2473, 94.6940, I, 7/29/12; 29.9351, 94.5390, 2/22/14; 30.4427, 94.8428, I, 6/29/16, 6/20/18; 30.2138, 95.1048, E, I, 3/25/07; 30.3254, 95.0489, 12/23/07; 30.1884, 94.9431, 12/28/08; 30.2391, 94.9661; 30.2921, 94.9781, I; 30.1456, 94.9278, I; 30.1141, 94.9253, I, 10/20/13; 30.3648, 95.0118, I, 3/8/14; 30.4131, 94.9174, I. Marion: 32.7989, 94.3106, I, 4/6/14, DJ411; 32.8109, 94.1829, I, 4/11/08. Montgomery : 30.3005, 95.4021, 12/29/08; 30.2578, 95.1444, I; 30.2087, 95.4206, I; 30.2221, 95.6119, I; 30.2166, 95.7472, I; 30.1266, 95.6494, I, 1/3/09; 30.5275, 95.6820, I, 5/18/14; 30.4344, 95.6573, I, 5/19/14, DJ479; 30.4457, 95.7811, I; 30.0788, 95.2379, I, 6/13/15; 30.1376, 95.2176, I; 30.1281, 95.2816, I; 30.1785, 95.3203, I; 30.2145, 95.2795, I; 30.3408, 95.7688, 4/16/16; 30.2760, 95.7117, I; 30.0888, 95.2405, I, 8/14/16; 30.0962, 95.2184; 30.1271, 95.2806, I, 12/4/16, 10/22/17. Morris : 33.0084, 94.8080, I, 4/11/08. Nacogdoches : 31.4957, 94.7390, E, 3/16/07; 31.6700, 94.8812, I, 3/26/11; 31.6031, 94.5696, I, 3/29/13; 31.6502, 94.5819, I; 31.5813, 94.5986; 31.5186, 94.6545, I; 31.6150, 94.5150; 31.5665, 94.5241, I; 31.5825, 94.7261, I; 31.6701, 94.8656, I, 3/30/13; 31.6525, 94.7841, I; 31.6887, 94.7544, I; 31.8238, 94.9376; 31.7498, 94.9572, I; 31.6714, 94.9345, I. Newton: 30.2899, 93.7416, E, I, 2/3/07; 30.6662, 93.6854, 1/20/08; 30.4263, 93.8045, I, 2/13/10, DJ61; 30.2476, 93.7717, I, 2/23/14, DJ539; 30.9235, 93.8282, I, 3/22/14; 30.8474, 93.7649, I; 30.7858, 93.8017, I; 30.7357, 93.8240, I; 30.6593, 93.8106, I; 30.5662, 93.7847, I, 3/23/14; 30.6093, 93.7593, I, DJ361; 30.7218, 93.7550, I; 30.7392, 93.6977, I; 30.7750, 93.6700; 30.8813, 93.7153, I. Orange: 30.1150, 94.0079, I, 2/22/14, DJ362; 30.0701, 93.9550, I; 30.0755, 93.8875, I; 30.1611, 93.9562, I, 2/23/14; 30.1923, 93.8953; 30.1344, 93.8830, I; 30.1387, 93.8120, I; 30.2096, 93.8215, I; 30.0817, 93.8059, I; 30.0835, 93.7793, I; 30.0313, 93.8170, I; 30.1756, 93.7573, I. Panola: 32.0754, 94.1098, I, 5/2/14, DJ446; 32.2943, 94.4901, I, 4/12/15. Polk : 30.8363, 94.9506, I, 5/8/09; 30.8556, 94.8541, I, 1/4/09; 30.9046, 94.8122; 30.6576, 94.9474, I; 31.0944, 94.8034, I; 30.8569, 94.8526, I, 2/15/13; 30.5556, 94.8102, I, 3/8/14, DJ363; 30.5871, 94.7737, I; 30.5706, 94.6338, I; 30.6741, 94.6958, I; 30.7157, 94.7192, I; 30.7887, 94.7715, I; 30.7026, 94.8378, I; 30.7401, 94.6262, I, 3/9/14, DJ533; 31.0231, 94.8217, I, 4/26/15, DJ674; 30.9826, 94.8268, I; 30.8967, 94.9269, I, 5/5/16; 30.6349, 94.8624; 30.5903, 94.9165; 30.7392, 95.0244, 5/6/16; 30.7694, 95.0425; 30.8143, 95.1197; 31.0276, 94.7389, I, 7/1/16; 30.7342, 95.0749, I, 11/1/17; 30.6552, 94.9980, I, 10/16/18. Rusk : 32.0410, 94.8170, 2/16/13; 31.9807, 94.8670, I; 31.8777, 94.9221, 2/17/13; 31.9125, 94.7195; 32.0814, 94.8228, I, 5/2/14, DJ473; 31.9245, 94.9769, I, 5/3/14; 32.3626, 94.5958, I, 4/12/15, DJ673; 32.3197, 94.7338, I, 2/17/13; 32.3001, 94.7933, I, 4/12/15; 32.3262, 94.9728, I. San Jacinto: 30.4044, 95.1629, I, 3/2/14; 30.5447, 95.2929; 30.5753, 95.3431; 30.5981, 95.2077, I, DJ367; 30.5668, 95.1925; 30.4912, 95.1643; 30.4250, 95.1289, I. Smith: 32.4244, 95.5234, I, 6/13/14, DJ486; 32.2152, 95.3150, I, 4/25/15; 32.1651, 95.2688, I; 32.4339, 95.1668, I, 4/11/15. Trinity : 30.8703, 95.2047, 1/6/08; 30.9825, 95.1442; 30.9386, 95.1393, I, 5/8/09; 31.0063, 94.9977; 31.1921, 95.0289, I; 31.1477, 95.2964, I, 5/10/09, 4/24/15; 31.1710, 94.9233, I, 3/21/13, Dave Peterson ; 31.1388, 94.9082, I, 5/17/15; 31.1110, 94.9883, I; 31.0795, 95.0389, I; 31.0474, 95.1052, I; 31.1248, 95.0996, I; 31.1571, 95.0743, I; 31.2227, 94.9726, I; 31.2739, 94.9253, I; 31.2575, 95.0696, I; 31.1987, 95.0997, I; 31.1956, 95.1659; 31.1901, 95.2283, I; 31.0982, 95.1800, I; 31.0543, 95.2348, I; 30.8578, 95.1897, 5/6/16; 30.9184, 95.2414; 30.9549, 95.0999; 30.9436, 95.0879; 31.0023, 95.0989, I; 31.0447, 95.0133; 31.0642, 94.9215; 30.9763, 95.3621, 5/22/16; 30.9997, 95.3594; 31.0265, 95.3557; 30.9540, 95.2948; 31.0087, 95.1931, I. Tyler: 30.5634, 94.2042, E, I, 2/3/07; 30.6585, 94.3928, 2/3/08; 30.7645, 94.4805, I, 3/9/14; 30.8345, 94.5351, I; 30.7939, 94.3720, I; 30.7847, 94.2359, I, DJ529; 30.7041, 94.1775, I; 30.6847, 94.3046, I; 30.5672, 94.4001, I; 30.6033, 94.3020, I; 30.9299, 94.4518, I; 30.9304, 94.2418, I, 3/21/14; 30.9839, 94.2650, I; 31.0009, 94.3283, I; 31.0168, 94.3806, I, DJ368; 30.9580, 94.5721, I; 30.9059, 94.5802; 31.0300, 94.5309, I, 7/1/16. Upshur : 32.7732, 94.9453, I, 4/11/15; 32.7775, 94.9898, I; 32.7938, 95.0971, I; 32.7822, 95.1333, I. Van Zandt: 32.3617, 95.8041, I, 6/12/14, DJ447; 32.3843, 95.8501, I; 32.4186, 95.7625, I; 32.3747, 95.4779, I; 32.4273, 95.5310, I, 6/13/14. Walker : 30.9483, 95.5682, I, 5/9/09; 30.8957, 95.5173, I, 3/17/13; 30.6883, 95.6836, I; 30.6481, 95.6737, I; 30.5810, 95.5606, I, 5/19/14, DJ475; 30.5909, 95.6212, I, 3/7/15. Waller : 30.1940, 95.9337, I, 12/8/08; 30.1969, 96.0742, I; 29.8011, 95.8898, I, 9/30/13; 29.8703, 95.8906, I; 29.8741, 95.9592, I; 29.9502, 95.9583, I; 30.0236, 95.9429, I, 3/1/14, DJ487; 30.0697, 95.9809, I; 29.7870, 95.9569, I, 5/29/15; 29.9978, 96.0357; 30.0516, 96.0378, I. Wood: 32.7334, 95.2331, I, 4/9/11; 32.8532, 95.2375, I, 4/4/14.

Variations. Variations in a number of key morphometrics and meristics may be found in Tables 2 and 7–10. Additional variations not found in those tables are provided here. The rostral margins vary from straight to slightly convex and moderately to weakly converging. The acumen may be moderately-well delimited or not obviously delimited. The rostrum L/W ratio ranges from 1.2 to 1.7 (x = 1.4, s = 0.1, n = 49). The ratio of the rostrum L to CL ranges from 17.1 to 21.3% (x = 19.2, s = 1.0, n = 49). The cervical tubercle is always weak but shows some variation in size. The branchiostegal spine ranges from small but distinct to nearly imperceptible and is occasionally doubled. The suborbital angle ranges from very small to moderate in size but is always obtuse. The anterior lobe of the epistome varies in shape from subtriangular, to subsemicircular, to subtrapezoidal. The caudolateral angle of cephalic section of telson is occasionally missing one of the movable spines. Ranges in the numbers of tubercles on the various podomeres of the cheliped are as follows: 8–15 (x = 11.9, s = 1.6, n = 64) in the main row on the dactyl’s mesial margin, 4–11 (x = 7.9, s = 1.7, n = 64) and 0–3 (x = 1.5, s = 1.2, n = 11) in 2 rows dorsolaterally neighboring it, and 0–2 (x = 0.5, s = 0.8, n = 11) in the row ventrolaterally neighboring it; 6–9 (x = 7.1, s = 0.8, n = 63) in the main row of the palm’s mesial margin, 5–9 (x = 6.4, s = 0.9, n = 63) and 0–8 (x = 3.0, s = 2.4, n = 11) in the 2 rows dorsolaterally flanking the main row, and 1–5 (x = 3.2, s = 1.3, n = 11) in the row ventrolaterally flanking the main row; 11–17 (x = 13.2, s = 1.2, n = 63) in the ventromesial row of the merus, and 3–7 (x = 5.1, s = 0.9, n = 63) in the ventrolateral row; and 0–5 (x = 2.2, s = 1.4, n = 62) in the ventromesial row of the ischium. The distomedian spine of the uropodal endopod ranges from distinctly premarginal to scarcely premarginal. Figures 21–22 illustrate variation in the form I male gonopod. The angle of recurvature of the CP ranges from 160° (Fig. 21y) to 190° (Fig. 21k), while that of the MP ranges from 105° (Fig. 21a′) to 185° (Fig. 22d). The gap between the processes is usually minimal, but may be entirely absent (Fig. 21r), slight proximally (Fig. 21e), distinct distally (Fig. 21x), broken near midlength (Fig. 21y), or complete but narrow (Fig. 22c′). The MP distal extremity ranges from acuminate (Fig. 21b) to hair like (Fig. 22e) to blunt (Fig. 22q); and from slightly procurved (Fig. 21q) to slightly recurved (Fig. 21g′). The MP ranges from slightly shorter than (Fig. 21i′) to distinctly longer than (Fig. 22t) the CP. The gonopod shaft varies from unarched (Fig. 21h′) to moderately caudally arched (Fig. 22b′). The annulus ventralis never occurs as a mirrored image of that of the allotype.

FIG. 23. Creaserinus clausus n. sp. type locality habitat.

FIG. 24. Creaserinus clausus n. sp.: counts of eggs or young for reproducing females.

Life history notes. Seasonal collection data of Creaserinus clausus n. sp. ( Table 23 ) exhibit patterns consistent with that of the genus Creaserinus as a whole. Refer to its “Life history notes” for discussion. Data for ovigerous females and females carrying young are provided in Table 24 and illustrated in Fig. 24.

Ecological notes. This species has been found in prairie, pine forest, hardwood forest, mixed forest, and urban habitats; and has been collected from ditches, pools, tire ruts, ponds, swales, marshes, swamps, and occasionally from streams. Water may be clear, turbid, or tannin-stained, and is nearly always temporary.Among its crayfish associates, it prefers less temporary waters than the extremely temporary ones dominated by juveniles of Fallicambarus houstonensis , F. kountzeae , and F. macneesei ; but more temporary waters than all other non- Creaserinus crayfish associates occurring in open waters. Soils may be sandy, loamy, or clayey. Hydrophytic or nonhydrophytic vegetation may or may not be present. Plant associates include Polygonum , Cyperus , and Alternanthera philoxeroides . Fish associates include Gambusia . It is frequently associated with lowland habitats found near streams and rivers. Of burrows excavated for this species, most are in the configuration of a single, subvertical tunnel with depths ranging from 25 to 60 cm; but one burrow extended vertically from the surface to 30 cm, then horizontally 60 cm, then curved back downward to an ultimate depth of 60 cm.

Etymology. Clausus (L.) = close, alluding to the close proximity of the proximal halves of the gonopod processes typically seen in form I males of this species.

Crayfish associates. Collected with Creaserinus clausus n. sp. were the following32 crayfish taxa: Procambarus (Ortmannicus) acutus (n = 101, 26.4%), Procambarus (Girardiella) kensleyi (n = 62, 16.2%), Procambarus (Girardiella) curdi (n = 54, 14.1%), Lacunicambarus ludovicianus (n = 40, 10.4%), Cambarellus (Pandicambarus) puer (n =32,8.4%), Procambarus (Scapulicambarus) clarkii (n = 32,8.4%), Procambarus (Ortmannicus) zonangulus Hobbs & Hobbs, 1990 (n = 27, 7.0%), Fallicambarus houstonensis Johnson, 2008 (n = 25, 6.5%), Procambarus (Ortmannicus) nechesae Hobbs, 1990 (n = 24, 6.3%), Fallicambarus macneesei ( Black, 1967) (n = 23, 6.0%), Faxonella beyeri (n = 17, 4.4%), Procambarus (Girardiella) simulans (n = 17, 4.4%), Procambarus (Ortmannicus) albaughi Johnson, 2018 (n = 15, 3.9%), Cambarellus (Pandicambarus) shufeldtii (n = 14, 3.7%), Faxonella clypeata ( Hay, 1899) (n = 13, 3.4%), Faxonella blairi Hayes & Reimer, 1977 (n = 12, 3.1%), Procambarus (Ortmannicus) acutus X zonangulus ( Johnson, 2018) (n = 10, 2.6%), Procambarus (Capillicambarus) hinei ( Ortmann, 1905b) (n = 10, 2.6%), Fallicambarus devastator Hobbs & Whiteman, 1987 (n = 8, 2.1%), Procambarus (Ortmannicus) acutus X albaughi ( Johnson, 2018) (n = 5, 1.3%), Procambarus (Pennides) dupratzi Penn, 1953 (n = 5, 1.3%), Procambarus (Girardiella) ceruleus Fitzpatrick & Wicksten, 1998 (n = 4, 1.0%), Procambarus (Girardiella) nigrocinctus Hobbs, 1990 (n = 4, 1.0%), Creaserinus hedgpethi (n = 4, 1.0%), Creaserinus trinensis n. sp. (n = 4, 1.0%), Fallicambarus kountzeae Johnson, 2008 (n = 4, 1.0%), Procambarus (Pennides) natchitochae Penn, 1953 (n = 2, 0.5%), Creaserinus limulus n. sp. (n = 1, 0.3%), Cambarellus (Pandicambarus) texanus Albaugh & Black, 1973 (n = 1, 0.3%), Faxonius texanus ( Johnson, 2010) (n = 1, 0.3%), Faxonius maletae ( Walls, 1972) (n = 1, 0.3%), and Faxonius lancifer ( Hagen, 1870) (n = 1, 0.3%).