Creaserinus hedgpethi ( Hobbs, 1948 ) 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 : 58-64

Creaserinus hedgpethi ( Hobbs, 1948) stat. rev., n. comb.

Figs. 3, 5, 6f, l, 7c, g, j, 8c–e, 9–12, 30–34, Tables 1 , 4, 8, 11, 14–16, 27 , 28

Cambarus argillicola Faxon, 1884:115 [in part].

Cambarus fodiens .— Huntsman, 1915:158 [in part; by implication].

Cambarus hedgpethi Hobbs, 1948:224 , Figs. 17a–f, h–j, l [in part].

Fallicambarus hedgpethi .— Hobbs, 1969:111 [in part].

Fallicambarus (Creaserinus) hedgpethi .— Hobbs, 1973:463 [in part; by implication].

Fallicambarus (Creaserinus) fodiens .— Hobbs & Robison, 1989:672, Figs. 9l–p [in part].

Fallicambarus (Creaserinus) fodiens “TX-1”.— 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 135–175° to axis of shaft, with distal part directed caudoproximally with tip never crossing central projection of corresponding gonopod, and base not inclined laterally, length 28.3– 37.6% (x = 33.8, s = 1.5, n = 184) of total gonopod length. Mesial process noncorneous, recurved 85–155° to axis of shaft, proximal half not strongly inflated. Proximal gap between processes usually present. 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 hedgpethi and a refers to any of other five Texas species (refer to Tables 4, 8, 11, 14, and 16). Distribution west of San Jacinto River or south of latitude 30.28. Sex ratio of juvenile population (<16 mm CL) not strongly skewed female.

Size. Of 197 form I males for which measurements were made, carapace lengths range from 21.3 to 38.6 (x = 29.0) mm.

Range and specimens examined. Creaserinus hedgpethi has been found at 203 sites in 28 counties in Texas (Fig. 10), of which 165 (81.3%) are represented by at least one form I male and 14 are represented in the molecular phylogeny. Based on the distributional patterns, an additional 220 sites in the southwestern coastal plain are very likely attributable to this species. These sites are represented by juveniles and females, which are unidentifiable to species. Creaserinus hedgpethi may be regarded as a Texas endemic; however, it would not be surprising to find this species in the southwestern coastal plain of Louisiana as the Texas distribution nearly reaches the Louisiana border.

Prior to its synonymization with Fallicambarus (Creaserinus) fodiens by Hobbs & Robison (1989:681) (using the accepted nomenclature at the time), F. (C.) hedgpethi was erroneously regarded to have a broad southern distribution which included Texas, Oklahoma, and Louisiana, eastward through Tennessee and Georgia ( Penn, 1959:195; Reimer, 1969:50; Hobbs, 1972:102; Hobbs, 1981:269).

Aransas: 28.2328, 96.8741, I, USNM no. 85146. Austin : 29.7561, 96.0487, E, 12/31/06; 29.6399, 96.0567, I, 12/6/08; 29.6378, 96.0797, I; 29.7372, 96.1439, I; 29.8683, 96.3538, I; 29.7766, 96.1062, I, 10/14/13; 29.7076, 96.1934, I. Brazoria: 29.1432, 95.6271, E, I, USNM no. 144454; 29.1875, 95.3905, I, USNM no. 144464; 29.0273, 95.5615, 12/7/08; 29.3592, 95.2705, 1/31/10; 29.4536, 95.4284, 2/11/11; 28.9726, 95.5305, I, 12/7/08; 29.1168, 95.3821, I; 29.2839, 95.2107, I; 29.0485, 95.4710, I; 29.3207, 95.4507, I, 1/31/10; 29.5589, 95.3526, I, 12/7/08; 29.3022, 95.7292, 10/13/13; 29.2983, 95.7327, I; 29.3976, 95.2572, I, 6/27/14; 29.3703, 95.3248, I; 29.3522, 95.4299, I; 29.3232, 95.4513; 29.2673, 95.4422, I, DJ517; 29.2490, 95.4171, I; 29.2457, 95.4031, I; 29.2767, 95.3552, I; 29.2923, 95.3212; 29.3152, 95.2503, I; 29.2643, 95.1509, I, DJ466; 29.2941, 95.1216; 29.4295, 95.2243, I, 6/14/15; 29.5180, 95.3082, I. Brazos: 30.6527, 96.4434, I, USNM no. 144458; 30.8685, 96.1932, E, I, USNM no. 144466; 30.8754, 96.3050, I, 3/15/15. Burleson : 30.4078, 96.8167, 3/15/15; 30.4523, 96.6533, I, DJ656; 30.4152, 96.5432, I. Calhoun: 28.4196, 96.6080, 2/28/10; 28.4774, 96.5653, I; 28.5833, 96.6531, I; 28.6871, 96.5183, I; 28.6064, 96.8568, I, 5/2/13. Chambers : 29.8158, 94.6569, I, 9/22/13; 29.7590, 94.6703, I; 29.6727, 94.4885, I; 29.7345, 94.3826, I; 29.7332, 94.4556, I; 29.7302, 94.6058, I; 29.7906, 94.5406, I; 29.7927, 94.4103, I; 29.8513, 94.3843, I; 29.8596, 94.4915, I; 29.8599, 94.5651, I; 29.8656, 94.6246, I; 29.7784, 94.9127, I, 10/20/13; 29.7985, 94.8504, I; 29.8692, 94.8781, I. Colorado: 29.4091, 96.5438 , 5/22/10; 29.6613, 96.5761; 29.8013, 96.3577, I, 8/12/07; 29.5816, 96.3200, I, 5/22/10; 29.7481, 96.3719, E, I, 12/31/06; 29.6447, 96.2560, I, 10/14/13; 29.5063, 96.3162, I; 29.4558, 96.3820, I. DeWitt: 29.1240, 97.0261, I, 5/19/12. Fort Bend : 29.6504, 95.6860, 4/13/09; 29.5471, 95.7452, E, I, 3/11/07; 29.6130, 95.6837, I, 4/13/09; 29.6496, 95.9765, I; 29.6274, 95.8118, I, 1/29/10; 29.6711, 95.6233, I, 2/12/10; 29.5437, 95.9581, I, 7/21/12; 29.3503, 95.5910, I, 10/13/13; 29.3516, 95.6660, I; 29.3583, 95.7885, I; 29.3464, 95.8465, I; 29.4040, 95.8432, I; 29.4467, 95.8950, I; 29.5330, 95.6674, 1/24/15; 29.5125, 95.6114, I; 29.4977, 95.5933, I; 29.4987, 95.6814; 29.4751, 95.6561, I; 29.4575, 95.6443, I; 29.4286, 95.6892, I; 29.3976, 95.6005, I; 29.6649, 95.7206, I, 5/29/15. Galveston : 29.4386, 95.1114, I, 1/29/10, DJ75; 29.3186, 95.0695, 6/27/14; 29.3395, 94.9868, I, DJ497; 29.3584, 94.9415, DJ523; 29.5061, 95.1855, I, 6/14/15; 29.5001, 95.1275, I; 29.5131, 95.0041, I; 29.4556, 94.9704, I; 29.4107, 94.9709, I; 29.3930, 95.0310, I; 29.3983, 95.0985, I; 29.3909, 95.1773, I; 29.4812, 95.2192, I. Goliad: 28.4322, 97.3232, 4/23/10; 28.5964, 97.2590, I. Grimes: 30.7195, 96.1606, E, 4/13/07. Harris : 30.0285, 95.0754, I, 2/2/13; 29.9349, 95.3616, I, 7/14/12; 29.6081, 95.4742, I, 1/30/10; 29.6799, 95.4491, I, 2/12/10; 29.7197, 95.1590, I, 2/22/09; 29.8191, 95.1429, I, 2/12/10; 29.6930, 95.3858, I, 10/13/13; 29.8029, 94.9978, I, 10/20/13; 29.7436, 95.6051, I, 1/2/15, DJ546; 29.7261, 95.6881, I, 5/29/15; 29.6358, 95.4082, I, 5/31/15; 29.6507, 95.3560, I; 29.6336, 95.3085, I; 29.6307, 95.1665, I; 29.6171, 95.0876, I; 29.5899, 95.0348, I; 29.6626, 95.0364, I; 29.6644, 95.1020, I; 29.7112, 95.1027, I; 29.7455, 95.2575, I; 29.7837, 95.2515, I; 29.8187, 95.2667, I; 29.9009, 95.0636, I, 6/13/15; 29.9555, 95.0813, I; 29.5566, 95.1794, I, 6/14/15; 29.5736, 95.0879, I; 29.9414, 95.0749, I, 8/14/16, 10/22/17, 10/22/17. Jackson : 29.0166, 96.5031, 5/20/12; 28.8940, 96.6592, I, 1/16/10; 28.9508, 96.4905, I, 5/20/12; 28.9847, 96.7336, I, 5/19/12; 29.0658, 96.7698, I, 6/19/10; 29.1224, 96.7547, I, 5/19/12. Jefferson : 30.0395, 94.4079, I, 12/8/07; 29.9536, 94.3500, 2/17/08; 29.9605, 94.4023, I, 9/23/13; 30.0516, 94.3080, I; 29.8896, 94.0035, 3/27/15; 29.8696, 94.0686, I; 29.8405, 94.1693, I. Lavaca: 29.2368, 96.7873, 7/14/07; 29.1729, 96.9365, I, 4/24/09. Lee : 30.3389, 96.9723, I, 3/25/12; 30.3885, 97.0635, I; 30.2720, 97.0169, I, 3/14/15; 30.3478, 96.8945, I; 30.3299, 96.8546, I; 30.3312, 96.7770, I; 30.4482, 96.8743, I, 3/15/15. Leon : 31.2471, 96.2163, I, 6/6/09; 31.2528, 96.3170, I, 5/31/14, DJ518; 31.1741, 96.2913, I. Liberty: 29.9323, 94.6090, I, 9/22/13; 29.9270, 94.9014, I, 10/20/13; 30.0617, 94.7712, I; 30.1243, 94.7374, I; 29.9947, 94.9591, I; 29.8982, 94.4913, I, 2/22/14, DJ397; 29.9351, 94.5390, I; 29.9866, 94.5451; 30.0038, 94.4640, I; 30.0020, 94.4709, I. Matagorda: 28.8075, 95.9505, E, I, USNM no. 144483; 29.1261, 95.9821, 5/20/10; 29.1113, 95.8767, 5/21/10; 28.9306, 96.0959; 28.8195, 96.2099; 29.0611, 95.8418, I. Refugio: 28.3021, 97.3830, I, 2/27/10; 28.3815, 97.1967, I; 28.3294, 97.2759, 3/14/15. Robertson : 31.2554, 96.3392, I, 5/31/14, DJ491; 31.1356, 96.3842, DJ516; 30.8430, 96.6077, 5/16/15. San Patricio : 28.0440, 97.3298, I, 2/26/10. Victoria : 28.6006, 97.0438, 5/19/12; 28.7995, 96.8348, I, 1/16/10; 28.9491, 96.9915, I, 5/19/12. Waller : 29.9433, 96.0542, I, 9/30/13; 29.8892, 96.0341, I; 29.8244, 95.9935, I, 5/29/15; 29.8495, 96.0261, I; 29.9264, 96.0270, I; 29.9978, 96.0357, I; 30.1178, 96.0867, I. Washington: 30.2807, 96.1109 , I, 4/16/16. Wharton : 29.5895, 96.0909, 4/13/09; 29.4061, 96.2408; 29.2231, 96.3947, 5/11/12; 29.1482, 96.3974, I; 29.0291, 96.2076; 29.0417, 96.2701; 29.0882, 96.3248, I; 29.1919, 96.3512; 29.2344, 96.3748; 29.2294, 96.0352, 5/20/12; 29.4063, 96.0820, I.

Variations. Variations in a number of key morphometrics and meristics may be found in Tables 4, 8, 11, and 14– 16. Additional variations not found in those tables are provided here.All are based on form I males, unless otherwise noted. The rostral margins vary from moderately convex to straight to slightly concave, and weakly to strongly converging with a weakly- to moderately-distinctly-delimited acumen. The rostrum L/W ratio ranges from 1.1 to 1.4 (x = 1.3, s = 0.1, n = 44). The ratio of the rostrum L to CL ranges from 15.0 to 19.4% (x = 17.7, s = 0.9, n = 44). The branchiostegal spine varies from distinct and acute to absent. The suborbital angle varies from small and obtuse to obsolete. The shape of the anterior lobe of the epistome ranges from subtriangular to semicircular to elliptical. The caudolateral angle of cephalic section of telson normally bears 2 spines, but occasionally the mesial movable spine may be absent or as many as 3 movable spines may be present on one side. The telson is normally distally rounded, but in one specimen it is strongly emarginate. Ranges in the numbers of tubercles on the various podomeres of the cheliped are as follows: 8–14 (x = 10.2, s = 1.2, n = 141) in the main row on the dactyl’s mesial margin, 2–8 (x = 5.2, s = 1.2, n = 178) and 0–1 (x = 0.2, s = 0.4, n = 10) in the 2 rows dorsolaterally bordering it, 0–4 (x = 0.6, s = 1.3, n = 10) in the row ventrolaterally bordering it; 5–9 (x = 6.6, s = 0.8, n = 74) in the main row of the mesial margin of the palm, 2–8 (x = 5.2, s = 1.3, n = 74) and 0–1 (x = 0.4, s = 0.5, n = 10) in the 2 rows dorsolaterally flanking it, and 0–4 (x = 1.8, s = 1.5, n = 10) in the row ventrolaterally flanking it; 11–16 (x = 13.7, s = 1.1, n = 118) and 3–8 (x = 5.2, s = 1.1, n = 118) in the merus’s ventromesial and ventrolateral rows, respectively; and 0–7 (x = 3.8, s = 1.1, n = 186) in the ventromesial row of the ischium. One specimen bore two well-formed uropodal exopods on one side. Figures 32 and 33 illustrate variation in the form I male gonopod. The angle of recurvature of the CP ranges from 135° (Fig. 32a) to 175° (Fig. 32h′), while that of the MP ranges from 85° (Fig. 32c) to 155° (Fig. 33z). The caudal offset of the MP varies from slightly less than (Fig. 32e′) to distinctly greater than (Fig. 32b′) that of the CP. The distal half of the MP may be procurved (Figs. 32j, t, w), recurved (Fig. 33z), or straight (Fig. 32q). The processes may be distinctly separate along their entire lengths (Fig. 33i), or the separation may be absent at midlength (Fig. 32y), or along the distal two-thirds (Fig. 32n′). The tip of the MP may be sharp (Fig. 32e′) or blunt (Fig. 33s). The distal half of the MP may be broad (Fig. 32l′) or narrow (Fig. 33y). The gonopod shaft varies from unarched (Fig. 32c) to moderately arched (Fig. 33f′). The opposable margin of the MP varies from gently rounded (Fig. 33h) to subangular (Fig. 32x). The annulus ventralis never occurs as a mirrored image of that of the allotype (see Hobbs, 1948:226).

FIG. 30. Creaserinus hedgpethi : (a) lateral and (b) dorsal views of form I male. Specimen exhibits broad-striped phase.

FIG. 31. Creaserinus hedgpethi : dorsal view of form II male. Specimen exhibits narrow-striped phase.

Life history notes. Seasonal collection data of Creaserinus hedgpethi ( Table 27 ) 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 28 and illustrated in Fig. 34.

Ecological notes. This species has been found in prairie, rangeland, cropland, brushland, hardwood woodland, and urban areas; and has been collected from ditches, pools, tire ruts, swales, flooded fields, marshes, canals, ephemeral streams, and occasionally from semipermanently flowing streams and bayous. Waters are almost always temporary, vary from clear to tannin stained and/or turbid, and are nearly always fresh, but occasionally brackish. Depths are typically shallow but can be up to 1.2 m deep after rains. Aquatic environments vary from heavily vegetated to devoid of vegetation, with vegetation varying from hydrophytic to nonhydrophytic. Notable plant associates include Sagittaria , Polygonum , Lemna , Spartina , Ludwigia , Distichlis , and Tamarix . Algae are occasionally present. Fish associates include Gambusia . Exposure is sunny to shaded. Soils vary from clayey to sandy, although where this species overlaps with Creaserinus clausus , it is typically found in more clayey soils than is the latter. Of three excavated burrows for which notes were made, one had a single, subvertical but undulating shaft extending to 6 dm deep; a second was also single shafted, but extended vertically to 2 dm, then horizontally 6 dm, before continuing downward at least another 2 dm; a third was simply noted to have four entrances.

Crayfish associates. Collected with Creaserinus hedgpethi were the following 20 crayfish taxa: Procambarus (Scapulicambarus) clarkii (n = 46, 22.4%), Procambarus (Girardiella) simulans (n = 33, 16.1%), Lacunicambarus ludovicianus (n = 25, 12.2%), Fallicambarus macneesei (n = 25, 12.2%), Cambarellus (Pandicambarus) puer (n = 20, 9.8%), Procambarus (Ortmannicus) albaughi (n = 20, 9.8%), Cambarellus (Pandicambarus) texanus (n = 12, 5.9%), Procambarus (Ortmannicus) zonangulus (n = 10, 4.9%), Procambarus (Ortmannicus) parvus Johnson, 2018 (n = 7, 3.4%), Procambarus (Girardiella) curdi (n = 7, 3.4%), Cambarellus (Pandicambarus) ninae Hobbs, 1950 (n = 5, 2.4%), Fallicambarus houstonensis (n = 4, 2.0%), Creaserinus clausus (n = 4, 2.0%), Procambarus (Capillicambarus) incilis Penn, 1962 (n = 3, 1.5%), Cambarellus (Pandicambarus) shufeldtii (n = 3, 1.5%), Procambarus (Girardiella) ceruleus (n = 2, 1.0%), Procambarus (Capillicambarus) hinei (n = 2, 1.0%), Procambarus (Ortmannicus) acutus (n = 1, 0.5%), Procambarus (Ortmannicus) acutus X zonangulus (n = 1, 0.5%), and Procambarus (Ortmannicus) luxus Johnson, 2011b (n = 1, 0.5%).

FIG. 32. Creaserinus hedgpethi : mesial views of gonopods of form I males. Each photo annotated with name of county where collected. Part 1 of 2.

FIG. 33. Creaserinus hedgpethi : mesial views of gonopods of form I males. Each photo annotated with name of county where collected. Part 2 of 2.

FIG. 34. Creaserinus hedgpethi : counts of eggs or young for reproducing females.