Gryllus longicercus Weissman & Gray, n. sp.

Long Cercus Field Cricket

Figs 206, 222–229, 232, 233, Table 1

‘ Gryllus #13’ of DBW notebooks.

‘G. longicercus’ of Gray et al. (2016a).

Distribution. Southwestern states of Arizona, New Mexico, and Texas, with minor extensions into California, Utah, and Colorado.

Recognition characters and song. A medium to large, usually short hind winged cricket with the longest known cerci in the genus, sometimes exceeding the body length of pinned specimens (male from Kofa [S88-88, R88-172]). Intact cerci always considerably longer than ovipositor in situ (except for individuals from Tijeras Canyon, NM, S19-35). File teeth from ~170–220 (range 155–231).Almost always associated with rocks. Song (Fig. 225, R 15-159) a slow to medium fast chirp with 4–5 (range 3–6) p/c, PR usually from 9–12 with individual pulses distinguishable by ear and almost countable at 25°C. While G. longicercus can occur microsympatrically with several other slow chirping, rock loving, medium to long length cerci, western US Gryllus, its low PR makes them immediately identifiable in the field. Morphologically most similar to G. chisosensis but the latter, with its unique DNA, restricted to the Chisos Mountains in Big Bend, TX, where G. longicercus doesn’t occur (nearest population of G. longicercus is in the Chinati Mountains, just south Shafter [S16-4] and ~ 115 km from nearest population of G. chisosensis). Microsympatric with the Organ Mountains’ endemic G. sotol at Aguirre Springs, New Mexico, but separated by non-overlapping pulse rate, DNA, and fewer file teeth in the latter. Morphologically similar to G. leei but the latter, only on lava flows in Utah, separated by non-overlapping PR (lower in G. longicercus), almost non-overlapping tooth count (higher in G. longicercus) and DNA (Gray et al. 2019). Also, morphologically similar to G. vulcanus but the latter, only on lava flows in New Mexico, is a smaller cricket with fewer teeth, shorter tegmen, shorter cerci, and shorter ovipositor (Table 1, p. 18). Microsympatric with G. transpecos but separated by non-overlapping PR (lower in G. longicercus) and almost non-overlapping tooth count (higher in G. longicercus). Comparing females of G. transpecos from Van Horn, TX (S85-65, S09-64) with microsympatric G. longicercus, ovipositor in latter shorter than in former despite larger body size (can also separate female G. transpecos from female G. longicercus because of cerci length in G. transpecos are as long as or just longer than the ovipositor in situ). Frequently microsympatric, in road cuts, with G. lightfooti, but separated by non-overlapping PR (lower in G. longicercus) and almost nonoverlapping tooth count (higher in G. longicercus). Also, females of G. lightfooti difficult to distinguish although the latter usually have a tegminal bar vs. solid black tegmina in G. longicercus . Only known to be microsympatric with the more western, slow chirping, rock loving G. saxatilis in the Hualapai Mts., Mohave Co., AZ (S90-56), but the two species both occur in Washington Co., UT and could also be sympatric in south-eastern CA around Anza Borrego, San Diego Co. In such localities, the 2 taxa would still separate by pulse rate and DNA.

Holotype. Male (Fig. 226): Arizona, Yuma Co., Kofa National Wildlife Refuge, Palm Canyon, 16-ix-2011; molt to adult 25-x-2011. 2100’. 33° 21’ 37.8” -114° 06’ 22.2”. D.B. Weissman, D.W. Weissman. S11-93, R11-185, BL 19.06, HF 11.86, LC 16.48. Right tegmen removed 217 teeth, file length 4.3, TL 13.7, TW 5.4. Type deposited in CAS, Entomology Type #19267.

Paratypes. (Total 217♂ 158♀). Arizona. Cochise Co., Bisbee, 5500’, 1-vi-2013 (S13-18) 2♂ . Portal, 29-vii- 1981 4770’ (S81-37) 2♂. Chiricahua Mts., Fort Bowie, 4723’, 2-vi-2013 (S13-19) 13♂ 6♀ . Coconino Co., Sedona, 15-vi-1990, 4400’ (S90-49) 3♂ 2♀ ; 12-vi-1996, (S96-62) 5♂ 1♀. Hwy 89A 1 m S intersection with Hwy 179, 30- vi-1994, 4420’ (S94-36) 4♂ 2♀ . 2.6 m N Sedona, 25-vi-1980, 4600’ (S80-43) 2♂ 2♀ . Gila Co., Coolidge Dam, 30-vii-1981, 2400’ (S81-43) 1♂. Graham Co., 7-10 m NW Bylas, 30-vii-1981, 2800’ (S81-42) 1♂ ; 25-viii-1982 (S82-102) 8♂. Pinaleno Mts., Mt. Graham, 4-vii-1994, 5000’ (S94-54) 5♂ ; 10-vi-2012, 5000’ (S12-19) 2♂ 8♀. La Paz Co., 18-35 m N Wenden on road to Alamo Dam, 23-vii-1990, 1200’ (S90-70) 3♂ 1♀ . Mohave Co., Burro Creek Campground, 1960’, 3-iv-2004 (2004-002) 1♂ . Quartzsite, 26-vi-1980, 1000’ (S80-46) 2♂. Kingman, 3-viii-1991 (S91-68) 1♂; 2-viii-1992, 3700’ (S92-113) 1♂ 1♀. Road to Hualapai Mtn. Park SE Kingman, 19-vi-1990, 5000’ (S90-56) 3♂ 2♀. Pima Co., Ajo, 20-viii-1998, 1720’ (S98-72 & S98-74) 9♂ 6♀; 17-ix-2011 (S11-99) 1♂. Hwy 85 13m N Ajo, 1255’, 30-vii-2015 (S15-110) 1♂ 1♀ . Baboquivari Mts., Brown Canyon, 31° 45’ 37.0” -111° 31’ 58.8”, 3786’, 29-vii-2015 (S15-105) 1♂ 5♀ . Kitt Peak, Hwy 386 5.1 m from Hwy 86, 17-ix-2011, 4500’ (S11-98) 2♂ ; Kitt Peak picnic area, 10.5 m from Hwy 86, 8-vi-2013, 6277’ (S13-36) 3♂ 1♀ . Near Sonora Desert Museum, 28-vii- 1981, 2900’ (S81-35) 5♂ 2♀ . Organ Pipe Cactus National Monument Campground, 8-iv-2004, 1670’ (2004-086) 1♂. Santa Catalina Mts., Mt. Lemmon Rec. Area, 27-vii-1990, 2900’ (S90-83) 3♂ 2♀ ; 7-vii-1994, 3500’ (S94-61) 1♂; 27-vi-2009, 4820’ (S09-51) 1♂. 11-vi-2012, 4675’ (S12-23) 10♂ 9♀. Tucson, 27-vii-1990, 2600’ (S90-84) 1♂. Why, 20-viii-1998, 1740’ (S98-71) 9♂ 5♀. Santa Cruz Co., Madera Canyon, Bog Springs campground, 5010’ 19-viii-2004, 6♀ . Yavapai Co., Hwy 179 on west side Sedona, 4000’, 30-vi-2013 (S13-16) 4♂ 1♀ . Yuma Co., Telegraph Pass, 10-viii-1988, 600’ (S88-90) 1♂; 17-viii-1998 (S98-62) 4♂ 2♀. Kofa National Wildlife Refuge, Palm Canyon, 10-viii-1988, 2050’ (S88-88) 3♂ ; 4-vi-1989 (S89-26) 1♂ 6♀; 16-ix-2011 (S11-93) 10♂ 35♀. California. San Bernardino Co., Providence Mts. State Recreational Area, 5-vi-1989, 4100’ (S89-28) 2♂ 1♀ . Havasu City, 6- vi-1983 (S83-62) 1♂. 14 m N Vidal, 6-vi-1983, 1200’ (S83-63) 1♂ . San Diego Co., Road S 22 from 2.7 to 10 m SW Borrego Springs, 8-viii-1988, 1800’ (S88-81) 2♂ . Colorado. Las Animas Co., Trinidad Lake State Park, Carpios Ridge Campground, 6734’, 29-vi- 200, 1♂ , JA Cole. New Mexico. Bernalillo Co., Tijeras Canyon ~ 2 m W Tijeras, 6000-6600’ (S19-35, S19-37, S19-38), 3-5-vii-2019, 9♂ 8♀ , D.C. & K. Lightfoot. Chaves Co., Hwy 380 7.3 m E Roswell at mile post 163.4, 28-vi-2009, 3960’ (S09-59) 3♂ 5♀ . Hwy 82 ~ 33 m W Artesia, 30-vi-2015, 4824’ (S15- 56) 7♂ 9♀ . Dona Ana Co., Organ Mts., Aguirre Springs Rec. Area, 24-vi-1985, 6500’ (S85-93) 2♂ ; 17-viii-1993 (S93-70) 1♂; 3-vii-2015 (S15-77) 1♂; 19-v-2017 (S17-4) 1♂ 2♀. Tortugas Mts., near UNM, Las Cruces, 23-viii- 1982, 4000’ (S82-95 & S82-96) 2♂ . Sandoval Co., Jemez Pueblo, 22-viii-1982 (S82-90) 4♂ 2♀ ; 6-v-1985, 5600’ (S85-50) 3♂. Socorro Co., Los Pinos Mts., Goat Draw 4.5 m N Hwy 60, 16-viii-1993, 6440’, 34° 22’ 10” -106° 32’ 0” (S93-69) 1♂. Hwy 60 34° 24’ 12.07” -106° 30’ 29.16”, 29-vi-2015, 6004’ (S15-53) 4♂ 4♀ . Sevilleta National Wildlife Refuge, 15-viii-1993, 5145’ 34° 24’ 30’ -106° 56’ 43’ (S93-66) 4♂ 4♀. Torrance Co., Hwy 60 rocky road cut SW Mountainair, 29-vi-2015, 6037’, (S15-52) 5♂ 2♀ . Utah. Washington Co., Springdale, 9-viii-1991, 4000’ (S91-94) 1♂ . 2 m E of west entrance to Zion National Park, 10-vi-1996, 4220’ (S96-54) 4♂ 3♀ . Just west of Zion National Park west entrance, 10-vi-1996, 4060’ (S96-55) 1♂ 1♀ . Texas. Culberson Co., Guadalupe Mts., 4.4 km NE Pine Springs, 13-vii-2001, 5240’ (S01-63) 9♂ 4♀ . I10 8 m W Van Horn, mile post 133, 12-vi-1985, 4300’ (S85- 65) 3♂ 1♀ ; 29-vi-2009 (S09-64) 4♂. Jeff Davis Co., Davis Mts., 30° 40’ 8.73” -104° 02’ 14.72”, 6073’, 1-vii-2015 (S15-63) 2♂ 3♀ . Presidio Co., Chianti Mts., Hwy 67 0.5 m N Shafter, 3856’, 27-v-2015 (S15-2) 1♂ 1♀ . Hwy 67 just S Shafter, 3920’, 27-v-2016 (S16-4) 5♂ 1♀ . Randall Co., Palo Duro Canyon State Park, 12-vi-1988, 3600’ (S88-37) 4♂ 1♀ .

Derivation of name. In recognition of their long cerci, which sometimes exceed 19 mm (Ajo [S98-72] and Why [S98-71], Kofa [S11-93], AZ; Shafter, [S16-4], TX) in length and can be longer than body length of a pinned adult male (e.g. Kofa [S88-88]). Geographic range. (Fig 227). Populations extend into Mexico.

Habitat. From 183m (Telegraph Pass, AZ, S88-90) to 1980m (Aguirre Springs, NM). Rocky washes, rocky canyons with Joshua tree, Larrea, cactus, and Eriogonum, oak-juniper grassland with rocks, rocky slopes with pinyonjuniper, rocky road cuts, on ground at base of walls and in cracks on walls and rocky talus slopes. Never found with solid overhead tree cover. Occasionally in towns like Kingman, AZ (S91-68). Also in town in Ajo in cracks at a gas station (S98-72). One male collected from sotol at Aguirre Springs, NM (S15-77), while searching for G. sotol .

G. longicercus appears to ecologically replace G. saxatilis as one moves east and south into the Sonoran and Chihuahuan Deserts of the Southwest US. Once into western Texas, G. transpecos, the ecological equivalent of G. saxatilis, can be sympatric with G. longicercus

Life cycle and seasonal occurrence. No egg diapause (S88-37). Probably two generations/year, at least at the type locality (Palm Canyon, Kofa National Wildlife Refuge). At the type locality, DAG collected a single adult female 18-iii-2005 (and heard calling males); 12 adult males and 2 adult females 15-iv-2010; a single adult male 5-iv-2013; a single adult male 7-ix-2013 (parasitized by the tachinid Ormia ochracea); 6 adult females and 2 late instar females 6-viii-2016 (and heard calling males). Similarly, DBW collected a few singing males there on 4-vi- 1989 and saw many more late instars, which started molting to adults in early to mid-July. DBW collected apparent second-generation individuals (7 adult males, many adult females, and many late instars) on 16-ix-2011. Those late instars started molting to adult some 2 weeks later.

We have visited the Ajo/Why, Arizona, area four times after a collecting trip on 20-viii-1998, at which time we collected 15 individuals of G. longicercus and 5 other species of Gryllus . Our subsequent visits there were on 16-v-1999 (S99-26), 1-viii-2009 (S09-102), 17-ix-2011 (S11-99), and 29-vii-2015 (S15-109) and we only found G. longicercus singing during the last 2 visits, and then only one male each time. Apparently, rainfall has a significant effect on when certain species of Sonoran Desert field crickets appear.

During June, Arizona field trips, we collected many more late instar nymphs than adults at the following localities: Mt. Graham (S12-19), Mt. Lemmon (S12-23), and Fort Bowie (S13-19).

Variation. Female tegmina length: The large series of 35 females from the type locality, where G. longicercus is usually the only field cricket present [highly flight-capable G. staccato also found there once, 15-iv-2010 (DAG 2010-017 female, 2010-018 male)], demonstrates a range from long (most specimens) tegmina almost reaching the tip of the abdomen to very short ones covering about ½ of the abdomen. Both conditions are seen in other populations where several females have been collected. Hind leg color: In most specimens, entire hind leg is dark except for reddish area on inside of hind femur. All individuals from Washington Co., Utah, and near Roswell, New Mexico (S09-59), have the entire hind leg reddish colored. Hind wing length: Most field collected adults with short hind wings although many adults in towns of Ajo/Why, Arizona, with long hind wings. Additionally, over 70% of type locality late instar nymphs collected on 16-ix-2011 (S11-93) had long hind wings after molting to adult in the subsequent 2-4 weeks while all, simultaneously, field-collected adults from there were short winged. Length of cerci in situ: In G. longicercus, cerci usually considerably longer than ovipositor except for the high elevation (1830m) population from Tijeras Canyon, NM (S19-35), where the intact cerci in 4 females did not exceed the tip of the ovipositor. In fact, the two longest ovipositors in any females, from any population of this species, were from this site, perhaps indicating that the winters there are extremely cold. Two other females from 300m higher (S19-37) in the same area both had cerci longer than the ovipositor. Number of file teeth: Varies from 155-231 over its US range (Fig. 228), which is considerably more than the tooth range of 30-40 seen in most other Gryllus species. Significantly, a male from Mountainair, New Mexico (S15-52) with the fewest number of teeth (155) known for this species, sang (R15-341) with a pulse rate of 10.8 at 26°C. Both his 16S and ITS2 DNA mapped with other G. longicercus . Three other males from Mountainair had 162, 164, and 171 file teeth. Song: Male (Fig. 229A, R 93-153) from Sevilleta Refuge, NM (S93-66) with 7-9 p/c and PR 8.6. R09-112 (Fig. 229B) from Van Horn, TX (S09-64) with 9-30 p/c (most similar to a trill) and PR 11.8. Overall population characteristics: The population from 11.7 km E Roswell (S09-59) was unusual in several ways: (1) adults with brown-reddish heads compared with usual black heads. (2) Singing males were very wary and readily stopped singing when approached. They resumed singing only after much delay, even though air temperatures ~22°C. (3) Males made no chirping noises as we turned over rocks trying to collect. In most cases, in most other populations, males continue to sing after they have climbed down into their rock pile while we are moving rocks aside to locate them. (4) These males jumped well and readily while we were trying to collect them. Multilocus analysis maps a male from there (S09-59) with individuals from the type locality of Palm Canyon, AZ (2016-039), and Shafter, TX (S16-4). Tegminal markings: rare males and females have a light tegminal bar.

DNA. Multilocus 2016-039 (Kofa, type locality of Palm Canyon); G1433, 7.3 m E Roswell, NM (S09-59); and G3386, Shafter, TX (S16-4) are sister species (Gray et al. 2019) to New Mexico lava restricted G. vulcanus 2016 - 025 from El Malpais; and G1030, from Valley of Fires State Park (S07-46).

Discussion. Behaviorally, most male G. longicercus continue chirping while turning over rocks attempting to capture them, a characteristic fairly unique to this taxon but also sporadically seen in other rock chirping species. Oatmeal trails do well to attract adult females and even some adult males. Adult males sometimes climb and sing: 1 male singing 1.5m up in shrub at Palo Duro Canyon State Park, TX (S88-37).

Of the 7 adult males (probably only two or three were actually singing) collected at the type locality of Palm Canyon (S11-93) on 16-ix-2011, all were parasitized by the tachinid Ormia ochracea . Five males had 1 fly larva, one male had 2 larvae, and one male had 4 larvae. Of the many adult females and late instars of both sexes collected concurrently, none were parasitized. One of two males from Kitt Peak (S11-98) parasitized with one Ormia ochracea larva.

Late instars of G. longicercus have contrasting, banded abdomens, a common juvenile character of many species in the two rock-chirping groups: Longicercus Group and Saxatilis Group, and also seen in G. lightfooti .