Gryllus veletis (Alexander & Bigelow)

Weissman, David B. & Gray, David A., 2019, Crickets of the genus Gryllus in the United States (Orthoptera: Gryllidae: Gryllinae), Zootaxa 4705 (1), pp. 1-277 : 132-145

publication ID	https://doi.org/ 10.11646/zootaxa.4705.1.1
publication LSID	lsid:zoobank.org:pub:F534C43A-AB09-4CB3-9B08-FD5BDFD90298
persistent identifier	https://treatment.plazi.org/id/182387A8-09E2-FF02-51F6-F94F0282F8E9
treatment provided by	Plazi
scientific name	Gryllus veletis (Alexander & Bigelow)
status

Treatment

Spring Field Cricket

Figs 56 View FIGURE 56 , 57 View FIGURE 57 , 130–151 View FIGURE 130 View FIGURE 131 View FIGURE 132 View FIGURE 133 View FIGURE 134 View FIGURE 135 View FIGURE 136 View FIGURE 137 View FIGURE 138 View FIGURE 139 View FIGURE 140 View FIGURE 141 View FIGURE 142 View FIGURE 143 View FIGURE 144 View FIGURE 145 View FIGURE 146 View FIGURE 147 View FIGURE 148 View FIGURE 149 View FIGURE 150 View FIGURE 151 , 205, 206, Tables 1 View TABLE 1 , 2 View TABLE 2

1960 Acheta veletis Alexander & Bigelow. Evolution 14: 335. Type locality: Illinois, Piatt Co. Types in UMMZ (types photographed on OSF and Fig. 133 View FIGURE 133 , courtesy of Mark O’Brien).

1964 Gryllus veletis . Randell. R.L. 1964. Canadian Entomologist 96:1592.

G. veletis , ‘G. eastern veletis’, ‘G. #27’, ‘G. #30’, ‘G. #36’, ‘G. #43’ or ‘roadside G.’, ‘G. pecos’, and ‘G. utahensis’ of DBW notebooks.

Distribution. Most widespread US Gryllus species and found in every state except for those in the Southeast ( Walker 2019, Capinera et al. 2004).

Recognition characters and song. Shortest cerci (along with G. veletisoides and G. pennsylvanicus ) of any US Gryllus species. In fact, the cerci are so short that even in old adults, they are frequently intact and taxonomically useful in identification. Most variable cricket for body size (along with the almost equally widespread G. pennsylva-nicus— see Fig. 32 View FIGURE 32 , p. 49 View FIGURE 49 ) with western Texas specimens around twice the body length of those from North Dakota ( Fig. 134 View FIGURE 134 ). A small to large sized, black except for occasional red area on inside of hind femur, usually short hind winged, wide pronotum, cerci never as long as ovipositor in situ, spring and early summer species usually associated with a vegetation cover such as grass or roadside weeds. Song ( Fig. 135 View FIGURE 135 , R03-118) a slow, usually irregular chirp with 4–5 (range 3–7) p/c and PR generally between 20 to 30 at 25°C. Males can sing during morning hours, especially in spring when evening temperatures are cool, but also during hot afternoon hours (e.g. Blue Springs, Missouri, 21-vi-1993).

Separated from sympatric sister species G. planeta , in the Davis Mts. of Texas (S15-62) by habitat (oak woodland for G. planeta vs. grassy meadows for G. veletis ), and smaller, more slender size, with little or no overlap in the following 6 parameters ( Table 1 View TABLE 1 , p. 18): G. planeta has fewer file teeth, shorter length file, shorter tegmina length and width, shorter ovipositor and shorter hind femur length. Also overlaps in length of cercus with G. planeta but when body length size differences compensated for ( Fig. 136 View FIGURE 136 , all individuals from the Davis Mts., Texas), cerci relatively longer in G. planeta even if only by 1–2 mm.

G. veletis can be separated from the always allopatric, Clade 1 of G. montis from the Chiricahua Mts. in southeastern Arizona, by the longer cerci in G. montis .

Morphologically most similar to G. veletisoides but always allopatric (on different sides of the Sierra Nevada and Cascade Mt. Ranges) and the latter with fewer teeth in file ( Fig. 137 View FIGURE 137 ), but similar length ovipositors ( Fig. 138 View FIGURE 138 ), and ~3.5% divergent in DNA [510 bp of 16S, Tamura-Nei distance, mean ± SD, within N = 54 G. veletisoides : 0.001 ± 0.001; within N = 219 G. veletis : 0.014 ± 0.015; between G. veletisoides and G. veletis : 0.038 ± 0.004. 698 bp of ITS2, Tamura-Nei distance, mean ± SD, within N = 10 G. veletisoides : 0.0003 ± 0.0006; within N = 25 G. veletis : 0.0064 ± 0.0061; between G. veletisoides and G. veletis : 0.0359 ± 0.0028].

Separated from sympatric and sometimes synchronic G. pennsylvanicus where latter has a longer ovipositor ( Table 1 View TABLE 1 , p. 18), usually a higher tooth count, longer cerci ( Fig. 139 View FIGURE 139 ), overwinters in the egg stage with first adults appearing in August, and different DNA (721 bp of ITS2 ~5.8% divergent, N = 4 G. pennsylvanicus and N = 25 G. veletis ; Gray et al. 2019). More teeth in the file apparently results in a lower pulse rate in G. pennsylvanicus that is noticeable in the field when both species are chirping near each other and at similar temperatures.

Separated from faster chirping, microsympatric Clade 4 of G. montis near Reserve, NM (S07-53, 54) by longer cerci and narrower pronotum in latter, and DNA (Gray et al. 2019). Separated from G. fultoni , where G. veletis has almost no overlap with the former’s faster chirp rate and higher pulse rate, and in G. veletis , usually more p/c, lack of yellow cerci, and larger head not narrower than pronotum. Compared to G. veintinueve , G. veletis is larger in size, has a louder and faster chirp rate, longer tegmina, broader pronotum, and different DNA ( Fig. 6 View FIGURE 6 , p. 28 View FIGURE 28 ). Microsympatric with G. vulcanus at El Malpais, NM (S07-52), but only at edges of lava flows. The two species are easily separated there by the longer cerci and lower pulse rate in the latter.

Derivation of name. “ veletis ” chosen “in reference to its aggressive behavior” ( Alexander & Bigelow 1960) as individuals are more solitary than its 1960 “proposed” sister species G. pennsylvanicus . The web site Latin-dictionary.net defines “veles” or “velitis” (note spelling difference) as a masculine noun with the following definitions: 1. Guerrilla forces (pl.), irregular bands. 2. Light-armed foot-soldier. 3. Skirmishers. R. D. Alexander (pers. comm. to DBW, August, 2013) stated that he and Bigelow used this modified spelling in Acheta veletis (before their taxon was moved to Gryllus ) “because the ‘e’ sounds in the two words went together so appropriately.”

Geographic range. Fig. 140 View FIGURE 140 . See Walker (2019) for more eastern US localities. Probably into adjacent Mexico. Contrary to Alexander & Meral (1967), G. veletis does occur in many southwestern US areas without G. pennsylvanicus (compare Fig. 140 View FIGURE 140 with Fig. 32 View FIGURE 32 , p. 49 View FIGURE 49 ), but avoids the lower elevation, hotter desert regions.

Habitat. Usually in areas with increased vegetation cover, either natural or planted, around water courses, and on both sandy and loam surfaces. In western Utah, very easy to collect during the daytime, in May, 2015, since almost every area with good grass cover had individuals hiding under boards, rocks, and various items of trash. Rarely at edges of lava flows in New Mexico (S94-41). Also in Oregon at sulfur-saline hot springs (S04-44) and in cracks of saline-saltmarsh grassland (S04-54). From near sea level to over 2440 m (Otero Co., near Cloudcroft, NM). At La Junta, Colorado (S87-65), one singing male on tree trunk 3 m above ground with female nearby and a second female starting to climb up the same trunk. In another tree, one male singing 1 m above ground from under loose bark. At Sedalia, Missouri (S93-51), one male singing 1 m up in shrub. May be riparian associated in otherwise hot dry areas, e.g. G. veletis found in moist areas along Deer Creek, Garfield Co., UT (at 1740 m elevation) replaced by G. saxatilis within a few meters of transition to dry rocky slopes; the same situation applies in Lodore Canyon on the Green River, Moffat Co., CO (at 1560 m elevation).

Life cycle and seasonal occurrence. No egg diapause: Wanapum Dam Reservoir, WA (S83-42); north of Alturas, CA (S83-48); Sacramento Mts., NM (S83-80 & S85-53); Alpine, TX (S85-59); La Junta, CO (S87-65); near Farmington, NM (S87-81); west of Medicine Lodge, KN (S88-41); Tulsa , OK (S88-45); Mirror Pool, ND (S88-55); Flagstaff, AZ (S91-75); Kadoka, SD (S97-87); Tooele, UT (S04-107).

One generation/year. Overwinter as late instars with adults first appearing in mid to late spring, even as far north as Mirror Pool, North Dakota (S03-52). Present at Lakin, Kansas (S87-67) on 23-vi-1987 but adults absent here 27-viii-1989 (S89-67) when we only found late instars of G. veletis (doubt that they were G. pennsylvanicus , which should have already been adult starting in early August) in the same habitat. Alexander & Bigelow (1960) note how “a small percentage (of G. veletis ) matures without apparent delay (thus accounting for 2 nd generation adults), but most require several months.” We note possible 2 nd generation adults from Jensen, Utah (10-ix-1999, S99-127) and Tooele, Utah (collected 9-ix-2004, S04-107, with no egg diapause in collected females) but late surviving first generation individuals are also a possibility.

Variation. Body size: G. veletis displays more variation in body size (see Fig. 134 View FIGURE 134 ) than any US Gryllus , and this property has been partially responsible for our confusion as to how many populations are actually G. veletis . Individual body lengths, measured immediately upon killing, vary from 13.42/ 13.70 mm (male/female) from Mirror Pool, North Dakota (S03-52) to 29.5 mm (male from Alpine, Texas, S91-44)/ 26.78 mm (female from Columbus, Ohio, S03-65). Upon complete drying after pinning, these four individuals, respectively, measured 13.50/11.69 and 23.23/ 21.42 mm. Color: We have a few individuals that are light-colored as discussed by Alexander & Bigelow (1960, p. 337). Cercus length: As individuals get larger going north to south, within the US, cercus length also increases. Female tegmina length: Individuals, especially those from New Mexico and in populations with large sample sizes, display a range of tegmina lengths where from ½ to the entire female abdomen is covered, and not directly related to the shrinkage associated with drying. File teeth: see Fig. 141 View FIGURE 141 . Varies from 116 (Pecos, NM, S01- 45) to 175 (La Junta, CO, S87-65). Head size: Larger males, especially those from New Mexico and Colorado, occasionally have heads much wider than the pronotum. This pattern was not seen in those larger adults (our ‘G. #36’) from the Alpine-Marathon area of Texas. Hind wing length: Of 1091 collected US adults, 200 (18.3%) had long hind wings and 891(81.7%) had short hind wings. Those adults with long hind wings were approximately 60% male. Also see under Discussion. Songs: Roswell, NM (S09-58) male with 4-10 p/c (see Fig. 142 View FIGURE 142 , R09-73). Tegmina color: Guymon, Oklahoma (S88-39 vs. S09-77) and La Junta, Colorado (S87-65 vs. S09-82) show that both 2009 samples have much darker tegmina than the browner 1987 and 1988 samples.

Specimens examined. CANADA: Alberta: Lethbridge, Old Man River, 3-vi-2001. British Columbia: Osoyoos, Haynes Ecological Reserve, 18-vi-2004, 1160’ (S04-61 & 62). USA: Arizona: Apache Co., St. Johns , 20-v- 2001, 5830’ (S01-38). Coconino Co., Flagstaff, 5-viii-1991, 6810’ (S91-75); 15-vi-2007 (S07-57). California: Inyo Co., Bishop , 5-vi-1983, 4140’ (S83-56). 9 m N Bishop, 4-vi-1983, 4700’ (S83-55). Lone Pine, 5-vi-1983, 3700’ (S83-58). Modoc Co., 9.1 m N Alturas, 3-vi-1983, 4600’ (S83-48). San Bernardino Co., Barstow, 28-vi-2003, 2300’ (S03-73). Colorado: Boulder Co., Boulder, 30-vi-2007, 5800-7200’ (S07-72). Jefferson Co., 9.3 m W Golden, 26- vii-1997, 6440’ (S97-75). Larimer Co., Ft. Collins, 3-viii-1997 (S97-112). Otero Co., La Junta, 22-vi-1987, 4200’ (S87-65); 2-vii-2009 (S09-82). Idaho: Elmore Co., Mt. Home, 26-vi-2014 (S14-31). 1.5 m W Hill City, 16-v-2000, 5040’ (S00-8). 34 m E Mt. Home, 16-v-2000, 5520’ (S00-7). Owyhee Co., Mt. Home, 7-vi-1996, 3260’ (S96-41). 2 m N Bruneau, 2900’ (S04-40). 2 m S Bruneau, 7-vi-1996, 2600’ (S96-37). Valley Co., 6 m S Donnelly, 18-v-2000, 4700’ (S00-12). Washington Co., Andrus Wildlife Management Area, 11-v-1998, 3000’ (S98-41). Illinois: DeWitt Co., Clinton, 3-vi-2003, 820’ (S03-58). 3 m SE Clinton, 3-vi-2003, 850’ (S03-59). Indiana: Floyd Co., Georgetown, 4-vi-2003, 500’ (S03-63). Spencer Co., Dale, 4-vi-2003, 700’ (S03-60). Iowa: Freemont Co., Waubonsie State Park, 1-vi-2003, 1250’ (S03-55). Monona Co., Lewis and Clark State Park, 1-vi-2003, 1140’ (S03-54). Kansas: Barber Co., Medicine Lodge, 23-vi-1987, 1550’ (S87-68). 27 m W Medicine Lodge on Hwy 160, 14-vi-1988, 1750’ (S88-41). Comanche Co., at junction of Highways 160 and 183, 14-vi-1988 (S88-40). Kearny Co., Lakin, 23-vi-1987, 2950’ (S87-67). Norton Co., near Edmond 23-vi-1993, 2900’ (93-55). Sherman Co., Sherman Co. State Park, 24-vi-1987, 3500’ (S87-73). Goodland, 24-vi-1987, 3500’ (S87-74). Maryland: Prince George Co., College Park, 30-v-2004, 500’ (S04-35). Massachusetts: Berkshire Co., 29-vii-2006, 1700’ (S06-67). Missouri: Hickory Co., Hermitage, 20-vi-1993, 1000’ (S93-50). Pomme de Terre State Park, 20-vi-1993, 1000’ (S93-49). Jackson Co., Kansas City, 8-viii-2002, 860’ (S02-54). Pettis Co., Sedalia, 20-vi-1993, 950’ (S93-51). Montana: Gallatin Co., Bozeman, 15-vii-2004, 4450’ (S04-66). Nebraska: Dawes Co., 3 m E Crawford, 3-vii-2009, 3680’ (S09-88). Fillmore Co., Geneva, 7-viii-2002, 1420’ (S02-51). Lancaster Co., Lincoln, 7-viii-2002, 940’ (S02-52). Nevada: Humboldt Co., Paradise Valley, 8-vi-1996, 4600’ (S06-42). Elko Co., Angel Creek Camp, Ruby Mts., 6600’, 16-vi-2002, (DAG 2002-011). White Pine Co., Baker, 5300’, 9-vi-1996 (S96-49); 23-v-2015 (S15-16). 4.7-5.0 m W Baker, 9-vi- 1996, 6520’ (S96-46 & 47a). New Mexico: Apache Co., Springerville , 20-v-2001 (S01-42). St. Johns, 20-v-2001, 5830’ (S01-38). Bernalillo Co., Tijeras Canyon ~ 2 m W Tijeras, 6600-6900’, 3-5-vii-2019, S19-37 & S19-38, DC & K Lightfoot. UNM Campus, 13-vi-1994 (S94-33). 6 m NE Cedar Crest, 7035’, 12-vii-2015 (S15-78) DC Lightfoot. Manzanito Mts., Carolino Canyon, 7500’, 19-vii-2008 (S08-53) DC Lightfoot. Catron Co., Apache Creek , 3- vii-1994, 6420’ (S94-49). Reserve, 3-vii-1994, 5770’ (S94-50); 2.1-2.2 m SW Reserve, 3-vii-1994, 5920-6120’ (S94-51 & 52); 20-v-2001 (S01-40); 14-vi-2007 (S07-53). Chaves Co., Roswell, 28-vi-2009, 3800’ (S09-58). Hwy 82 near Elk, 28-vi-2009, 5840’ (S09-57). Cibola Co., El Malpais National Monument, Hwy 117 at 31 m S I40, 18- viii-1993, 7200’ (S93-71 & 73); 1-vii-1994 (S94-40 & 41); 14-vi-1996, (S96-68); 14-vi-2007 (S07-51 & 52). Grants, 18-viii-1993, 6340’ (S93-74); 1-vii-1994 (S94-43); 13-vi-1996 (S96-67). Hwy 117 at 35 m S I40, 14-vi-2007, 6870’ (S07-51). Hwy 53 at mile-post 61, 13-vi-1996, 7800’ (S96-66). Eddy Co., Guadalupe Mts., Dog Canyon 30-vii- 2003, 6289’. Lincoln Co., Carrizozo Rec Area, 2-vii-1994, 5560’ (S94-47). Valle del Sol Municipal Golf Course, 13-vi-2007, 5320’ (07-45). 12 m W Capitan, 2-vii-1994, 7060’ (S94-45). Otero Co., 1.8 m W Cloudcroft, 29-vi- 1983, 8300’ (S83-80); 7-vi-1985 (S85-53). 5 m E Cloudcroft, 30-v-2011, 8200’ (S11-24). San Juan Co., Farmington, 19-vi-1987 (S87-81). 12 m W Farmington, 19-vi-1987, S87-81, DC Lightfoot. San Miguel Co., Pecos, 21-v- 2001, 6944’ (S01-45). Pecos National Historic Park, 21-v-2001, 7280’ (S01-44). Pecos National Monument, 6-x-1994 (S94-106). Hwy 63 at mile-post 7.3, 21-v-2001, 7440’ (S01-46). Hwy 25 at Exit 307, 21-v-2001, 7270’ (S01-43). Socorro Co., Los Pinos Mts., Goat Draw, 16-viii-1993, 6440’ (S93-69). Sevilleta National Wildlife Refuge, 21-vi-1995, 5400’ (S95-50), DC Lightfoot. Valencia Co., 1 m S Los Lunas Bridge, 1-vii-1994 (S94-44). North Dakota: Richland Co., Mirror Pool, 30-vi-1988, 1030’ (S88-55); 31-v-2003, 46° 32.963’ -97° 19.310’ (S03-52). Ohio: Franklin Co., Columbus, 5-vi-2003, 800’ (S03-65). Oklahoma, Texas Co., Guymon, 13-vi-1988, 3380’ (S88- 39); 1-vii-2009 (S09-77). Optima Wildlife Refuge 15 m E Guymon, 1-vii-2009, 3120’, 36° 41’ 49.8” -101° 9’ 36.3” (S09-76). Tulsa Co., Tulsa , near intersection Hwy 244 and I44 around 5828 W. Skelly Dr., 232m, 36° 04’ 56.7” -96° 02’ 58.0”, 15-vi-1988 (S88-45); 23-vi-1993 (S93-57); 15-vii-2013 (S13-67); Near Exit 238 off I44, 2 m E Tulsa city limit, 15-vii-2014 (S13-68), 796’, 36° 09’ 37.4” -95° 47’ 33.1”. Oregon: Harney Co., Alvord Hot Springs, 13-vi- 2004, 4000’ (S04-44). Burns, 2-vi-1997, 4020’ (S97-51). 23.5 m S Burns, 14-vi-2004, 4000’ (S04-48). Jackson Co., Phoenix, 1500’, 24-ix-2004 (S04-132). Lake Co., 16-vi-2004, 4500’ (S04-54). Sherman Co., 0.7 m SE Biggs, 31-v- 1997, 220’ (S97-47). South Dakota: Jackson Co., Badlands National Park, Cedar Pass Overlook, 22-vi-1993, 3100’ (S93-53); 3-vii-2009 (S09-89). Kadoka, 30-vii-1997, 2200’ (S97-87). Jones Co., Murdo , 22-vi-1993, 2600’ (S93- 54). Minnehaha Co., Sioux Falls, 1-vi-2003, 1450’ (S03-53). Yankton Co., Lewis and Clark Lake, 21-vi-1993, 1500’ (S93-52). Texas: Brewster Co., Alpine, 9-vi-1985, 4200’ (S85-59); 5-vi-1991 (S91-44); 12-vi-2007 (S07-41); 2-vii- 2015 (S15-73). Marathon, 12-vi-2007, 3840’ (S07-40). 22 m E Marathon, 12-vi-2007, 3850’ (S07-35). Culberson Co., Hwy 10 25 m E Van Horn, 29-vi-2009, 4780’, 31° 4’ 20.4” -104° 23’ 21.8” (S09-63). Jeff Davis Co., Davis Mts., 1.1 m below Observatory, 6246’, 1-vii-2015 (S15-62). Fort Davis, Ft. Davis National Historic Site, 4852’, 1- vii-2015 (S15-67). 1 m SW Ft. Davis, 6-vi-1991, 4800’ (S91-46). Utah: Beaver Co., Beaver, 5878’, 23-v-2015 (S15-20). Milford, 4990’, 24-v-2015 (S15-19). Minersville, 19-iv-1999, 5140’ (S99-11). Garfield Co., Boulder, 24- iv-2006, 6580’ (S06-18). Panguitch, 19-v-2001, 6560’ (S01-35); 23-iv, 2006 (S06-17). Juab Co., Eureka, 5800’, 19-iv-1999 (S99-8); 23-v-2015 (S15-13). Nephi, 5200’, 10-ix-2004 (S04-115); 24-v-2015 (S15-23). Kane Co., Hwy 89 9.3 m SSW Long Valley Junction, 19-v-2001, 6740’ (S01-34). Road to Zion Narrows, 18 m NE Springdale, 10- vi-1996, 6740’ (S96-51). Millard Co., Delta, 4670’, 23-v-2015 (S15-14). Fillmore, 4920’, 31-vii-1992 (S92-102). Fillmore, Canyon Rd. 0.9 m E Business I15, 4920’ 18-v-2001, 38° 57.728’ -112° 18.439’ (S01-30). 8.5 m NW Fillmore, 18-v-2001, 4450’ (S01-28). Lava fields near Flowell, 19-iv-1999, 4520’ (S99-10); 18-v-2001 (S01-29). Salt Lake Co., Riverton, 4500’, 24-v-2015 (S15-24). Salt Lake City, near Airport, 24-v-2015 (S15-25); Salt Lake City, Avenues residential area, 19-v-2009. Sanpete Co., Manti, 5515’, 24-v-2015 (S15-23). 1.4 m SW Manti, 21-iv-1999, 5660’ (S99-17). 11 m N Manti, 21-iv-1999, 5700’ (S99-18). Sevier Co., Richfield, 5300’, 24-v-2015 (S15-21). Salina , 17-viii-2009, 5300’ (S09-122). Tooele Co., James Walter Fitzgerald Wildlife Management Area, 5200’, 23-v- 2015 (S15-12). Stockton, 5200’, 30-vii-1992 (S92-93); 23-v-2015 (S15-11). Tooele, 5200’, 9-ix-2004 (S04-107); 23-v-2015 (S15-10). Uintah Co., Jensen, 10-ix-1999, 4740’ (S99-127). Wayne Co., Hanksville, 11-vi-1996, 4500’ (S96-60). Vermont: Addison Co., Middlebury, 23-v-2009 (S09-16). Washington: Grant Co., Wanapum Dam Reservoir, 1400’, 1-vi-1983 (S83-42). Ephrata, 30-v-1997, 1240’ (S97-46). Grand Coulee, 30-v-1997, 1400’ (S97-43). Lake Lenore, 19-vi-2004, 1020’ (S04-60). Moses Lake, 21-vi-2014, S14-18). Lincoln Co., 2.5 m SE Grand Coulee, 30-v-1997, 1640’ (S97-42). Wyoming: Crook Co., Aladdin, 4-vii-2009, 3780’ (S09-90). Fremont Co., Shoshoni, 18-vii-2011, 4849’ (S11-72). 11 m N Shoshoni, 18-vii-2011, 4788’ (S11-71). Park Co., Cody, 16-vii-2004, 5140’ (S04-73). Platte Co., Guernsey, 27-vii-1997, 4000’ (S97-80). Near Guernsey, Road 270 3.9 m N Hwy. 26, 12-ix- 1999, 4300’ (S99-139).

DNA. Multilocus G30, from near type locality of G. veletis, Spencer Co. , Indiana (S03-60); and multilocus G2958, ‘ Gryllus utahensis’, Nephi, UT (S15-23) comprise one close pair within G. veletis . Multilocus G1345, our ‘ Gryllus #27’, Guymon, OK (S09-77); and multilocus G3075, our ‘ Gryllus #36’, Alpine, TX (S15-73) comprise a second close pair within G. veletis . All are sister species (Gray et al. 2019) to G. planeta . See Discussion below for consideration of whether or not DNA supports further sub-division of this widespread species.

Discussion. Our initial sorting of this species resulted in six different, geographically separate “species.” Further efforts to delineate these “species” boundaries, using morphological and genetic properties, were unsuccessful probably, in large part, due to the vast geographical area and ecological diversity covered and the fact that just one (or perhaps two) species seems to be involved. As an example, adult females of G. veletis from near the most northern limits in North Dakota are small and very agile while their western Texas conspecifics are large and so bulky (including males) that they jump poorly, if at all.

Nevertheless, it seems prudent to briefly discuss those characters that in certain populations gave us pause, as to species identification, so that further investigations can be directed here:

1. We initially (1987) considered a “Great Plains” grouping (our ‘ Gryllus #27’) from south-eastern Colorado, western Oklahoma, and Kansas to be distinct from G. veletis based upon the usual presence of long hind wings. For example, we found that 10 of 12 from La Junta, CO (S87-65); all 20 from Lakin, KS (S87-67); 3 of 4 from Medicine Lodge, KS (S87-68); all 11 from Goodland, KS (S87-74); and 21 of 23 from the town of Guymon, OK (S88-39 and S09-77), had long hind wings, loud songs, were very colonial, and at high, outbreak densities in these towns. More widespread collecting in nearby central Missouri in 1993, showed that this pattern was geographically limited since all 22 Missouri adults, about half from within towns, had short hind wings. The final tally for this 3-state region, within towns only, was that 65 out of 70 (93%) individuals had long hind wings. This pattern contrasts with ~12.5% long hind wings for G. veletis in the rest of its US distribution, both within and outside of towns. For ‘natural’ Optima Wildlife Refuge, Oklahoma (S09-76), 15 m E Guymon, 7 of 8 adults had short hind wings.

2. We initially considered individuals from around Tulsa , Oklahoma (our ‘G. #30’) to be distinct because males generally have fewer p/c and a more regular, faster chirp rate. But in comparison with our other US G. veletis , we find no consistent characters, including DNA, to separate them.

3. We initially considered individuals from western Texas (including Alpine, Marfa, Marathon, Davis Mts. and Amarillo—our ‘ Gryllus #36’) to be distinct because adult females are so large and bulky, that they cannot hop. Additionally, when the males in these western Texas populations are compared with those male G. veletis from throughout New Mexico (our ‘ Gryllus #43’), we find the heads in New Mexico to be wider, relative to pronotum width, than those of the larger males from Texas. It is doubtful that the few Texas adults that have long hind wings can fly because of their weight.

4. Sequencing of the 16S gene in individuals from western Utah revealed a second, distinctive clade (see Fig. 143 View FIGURE 143 ), which we initially called ‘G. utahensis’. Subsequent ITS2 and multilocus sequencing revealed that all of these individuals were consistent with G. veletis from Illinois and Indiana (near type locality), and that individuals in two Utah populations (Tooele, S15-10 and Richfield, S15-21) contained both haplotypes.

Despite our unwillingness now to further subdivide G. veletis , we do note strong subdivision within the nuclear ITS2 gene (see Fig. 132 View FIGURE 132 ) consistent with the two-pairs of the four samples in the multilocus tree. ITS2 similarity based on Tamura-Nei distances was 99.1 ± 0.76 within individuals from NV, NM, TX, and OK (n=10), 99.4 ± 0.45 within individuals from IN, IL, and UT (n=15), but 94.09 ± 0.26 between these groups.

G. veletis and G. pennsylvanicus are sympatric over much of their northern US distribution. Because the former overwinters as a late instar and the latter as an egg in obligate diapause, they mature and sing at different times in the subsequent spring and summer. They may have a brief period of synchronicity but such has only been studied in detail in Michigan ( Alexander & Meral 1967), and even then, complete resolution was difficult because of morphological similarity between the two taxa and almost identical songs, although Alexander & Bigelow (1960) noted that G. pennsylvanicus females tend to have longer ovipositors than G. veletis . We were consequently surprised to find the two species synchronic and easy to distinguish, based on pulse rate, number of file teeth, and length of ovipositor at localities in South Dakota, Nebraska, Missouri, and Utah ( Table 2 View TABLE 2 ). For instance, at Kadoka, SD (S97-87), both G. veletis and G. pennsylvanicus adults were singing on 30-vii-1997 although most individuals of the latter were several molts to adult. Our 2 collected adult male G. veletis had 134 and 138 teeth with laboratory recorded pulse rates of 25-35, at 23°C and 26.5°C, respectively. These rates were faster than the 4 male G. pennsylvanicus collected as late instars (we were unable to collect any adult males) and that molted to adults in mid-late August: they had 149-171 teeth and laboratory recorded pulse rates between 17.9-21.7, at 22.5-25°C. Interestingly, teeth/mm were essentially the same: 40.0- 40.6 in the 2 G. veletis males and 39.8-44.4 in the 4 G. pennsylvanicus males. But these pulse rate differences were easily appreciated in the field when the two taxa were singing together at similar temperatures. We also collected 1 adult female G. veletis with a short ovipositor, long tegmina, and no egg diapause when compared to those female G. pennsylvanicus collected as nymphs and that molted to adult 2 weeks later in mid-late August. A similar situation existed at the 5 other Nebraska, Missouri, and Utah localities listed in Table 2 View TABLE 2 , although we were never at Hanksville at the right time of year to document potential synchronicity. Likewise, the 5 localities from Oklahoma, Kansas, Missouri, and Colorado, where G. pennsylvanicus also has increased numbers of file teeth when compared with surrounding areas, do not have documented populations of G. veletis , but we suspect they do exist if we had been there earlier in the season. We wonder if these increased number of file teeth in G. pennsylvanicus are a reflection of character displacement in areas where the 2 taxa have overlap in singing adult males? In any case, these localities should be compared, for synchronicity between these 2 taxa, with the areas around Chadron, NB (S97-83 and 97-84), where, on 28-vii-1997, we collected 2 adult males and many late instars that yielded 9 G. pennsylvanicus males with 139-166 teeth, 39.7-48.8 teeth/mm, and pulse rates of 19.2-25, and 1 adult female and several late instars that yielded 5 G. pennsylvanicus females with ovipositor lengths of 14.0-16.0 mm. While we didn’t collect or hear any G. veletis there, we did see many small Gryllus nymphs that could be recently hatched G. veletis . Perhaps the 2 species around Chadron are not synchronic, or minimally so, so that character displacement doesn’t occur there?

Where it occurs, G. veletis is usually the most common field cricket, with apparent outbreaks, in towns, of mostly long hind winged individuals, occurring in certain Colorado, Kansas, and Oklahoma localities in 1987, 1988 and 2009. This finding is in contrast to reports of typically solitary individuals for more eastern US populations (summarized in Alexander & Bigelow 1960). We frequently find nymphs and adults of G. veletis , in normal population-size years, congregating together under objects in spring and early summer and infrequently associated with specific burrows. Because most of the western US dries out in the summertime, resulting in deep substrate cracks and suitable refuges, burrows may be less necessary for shelter. Eastern populations, where summer rains can be more significant and substrate cracks rare, could find burrows more important.

Although males in different populations can vary by over 100% in body length ( Figs 144 View FIGURE 144 , 145 View FIGURE 145 , 146 View FIGURE 146 ), showing a distinct north to south size cline, we find no similar differences in tooth file architecture with reference to number of teeth ( Figs 141 View FIGURE 141 , 147 View FIGURE 147 ). Because southern males are much larger than northern males, they also have correspondingly longer files ( Fig. 148 View FIGURE 148 ). If tooth density remained the same going south, then files would contain many more teeth, a situation not seen ( Fig. 147 View FIGURE 147 ). We do find that the density of file teeth increases ( Fig. 149 View FIGURE 149 ) going northward. Given that the PR ( Fig. 150 View FIGURE 150 ) and dominant frequency remain essentially the same throughout G. veletis’ range, this suggests that the larger southern males apparently close their wings faster (over a longer file with fewer teeth/mm) in order to have a constant pulse duration and a constant dominant frequency.

Preliminary hybridization studies conducted in 2003 between California virgin G. veletisoides and virgin G. veletis from the Midwestern US produced the following results: 3 females from Indiana crossed with Fresno, CA, males—two females had documented matings (with passed spermatophores) but only one of the two females laid eggs. The female in the third pairing laid eggs but no eggs from any female hatched. One female, from Illinois, mated with a Fresno, CA, male and laid eggs but none hatched. One female from Iowa matched with a male from Los Gatos, CA, laid eggs (never saw a passed spermatophore) but they never hatched. As controls, an Indiana female mated with a North Dakota male produced lots of eggs that hatched while a North Dakota female that mated with an Illinois male produced eggs that didn’t hatch.

G. veletis appears in the literature as an example of allochronic speciation ( Alexander & Bigelow 1960), studies on calling, movement, parasites, age, and mating ( Alexander & Meral 1967; French & Cade 1987; Zuk 1987, 1988; Thomson & Bertram 2014), genetics of wing morphology and diapause ( Bégin & Roff 2002), changes in calling songs as males age ( Fitzsimmons & Bertram 2011), and freeze tolerance adaptations ( Toxopeus et al. 2018).

Tachinid fly Exoristoides johnsoni View in CoL emerged from adult males from Utah (S96-50 and S96-51), Nebraska (S97- 83), and South Dakota (S03-53), the latter from a penultimate instar male.