Gryllus lightfooti Weissman & Gray, 2019

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 : 204-213

Gryllus lightfooti Weissman & Gray , n. sp.

Arboreal Desert Field Cricket

Figs 205 , 207–212 , Table 1

‘G. #23’, ‘yucca Gryllus’ in DBW notebooks.

Distribution. Southeastern California through central and southern Arizona and New Mexico, into western Texas.

Recognition characters and song. A frequently arboreal, locally common field cricket that ranges from southeastern California through central and southern Arizona and New Mexico, into western Texas. Typically, a desert grassland species, extending into desert scrub and upper desert grassland/oak conifer zones. They can also colonize suburban neighborhoods. A small (especially when in sandy desert habitats) to large cricket separated, sometimes with difficulty, from similar looking, occasionally sympatric chirping species by a combination of habitat, behavior, female color patterns, DNA, and song. Almost always with short hind wings ( Fig. 210 ); cerci in situ nearly as long as, to just longer than tip of ovipositor; females (and some males) typically with “tegminal bars”, a pale longitudinal strip on the anterior dorsal-lateral margins of the forewings ( Fig. 210 ), especially visible when tegmina are dark. Males frequently sing off the ground, in vegetation, especially during the daytime from within Yucca elata , and also from within pack rat ( Neotoma spp.) nests. Song (see Fig. 211 , R09-69) a medium-fast chirp (frequently over 150 c/m), generally 3-5 p/c, PR 18-29 at 25°C. Distinguished from other US western slow chirping Gryllus , some that can be sympatric, as follows: From the G. montis complex: the latter occur in sky islands, under a tree canopy, where males have a slower CR and whose cerci are never longer than the ovipositor. Also, despite being at higher, cooler elevations, G. montis molts to adult several weeks before G. lightfooti . One can conduct elevational transects at several places in Arizona (e.g. Madera Canyon, Kitt Peak, Mt. Lemmon, Hualapai Mt. Park ) and hear an almost seamless transition of slow chirpers when descending from tree covered sky island habitats, where G. montis lives, to the open grassland-shrub habitats of G. lightfooti . Once below tree cover on these elevation-transects, wandering adult females of Arboreal Desert Field Crickets with tegminal bars can occasionally be seen on the roads, although at Hualapai Mountain Road, both species continue to occur sympatrically, in interior chaparral habitat, as Kingman is approached. Separated from sometimes microsympatric G. longicercus by no overlap in PR, minimal overlap in tooth number and cerci length, and association with rocky habitats in G. longicercus . In elevation-transects on Mt. Lemmon and Kitt Peak, both taxa can occur together in areas of rocky road cuts. Separated from G. transpecos , where the two taxa could overlap in western Texas, although no such locality known, by, in G. transpecos , cerci in situ almost always longer than ovipositor tip, female tegminal bars usually absent, associated with rocks, small but consistent ITS2 DNA differences, and never climb into vegetation to hide or sing. Separated from G. saxatilis , which is microsympatric with G. lightfooti in desert washes in the eastern California Mojave Desert at Mt Pass (S91-71) and Halloran (S91-70), by, in G. lightfooti , presence of tegminal bar in females, fewer teeth and teeth/mm (but still with overlap), climbing into vegetation, and DNA. Distinguished from G. veletis by the latter’s shorter cerci and association with more vegetated habitat. Additionally, nymphs and adults of G. veletis are never tan-mottled in color, while those of G. lightfooti are. Distinguished from G. sotol , which is only known from the Organ Mts. of southcentral New Mexico, by G. sotol ’s shorter cerci, non-grassland habitat, and association with sotol. Distinguished from G. planeta of the Davis Mts., in western Texas, by the latter’s shorter cerci and sky island forested habitat.

Holotype. Male ( Fig. 210 ): New Mexico, Dona Ana Co., USDA Jornada Experimental Range , ~ 5 m E Las Cruces off Highway 70. 3-vii-2015, 4300’, 32° 28’ 33.33” -106° 44’ 10.86”, DBW and DW Weissman. S 15-75, R15-139, G3061. 16 S GenBank accession # MK 446528 BL 17.98, HF 10.0, LC 10.89. Right tegmen removed: 123 teeth, file length 2.8, TL 9.5, TW 4.0. Deposited CAS Entomology #19264.

Paratypes. ( Total 259♂ 227♀) Arizona: Cochise Co., Apache Pass , 5106’, 32° 09’ 07.8” -109° 28’ 52.6”, 2- vi-2013 (S13-20) 2♂ 2♀ . Bisbee , 5500’, 31° 26’ 46.9” -109° 55’ 37.9”, 1-vi-2013 (S13-18) 4♂ 1♀ . Fort Bowie parking lot 2 m NE Apache Pass , 4723’, 32° 09’ 23.8” -109° 27’ 09.9”, 2-vi-2013 (S13-19) 13♂ 9♀ . Southwestern Research Station , 5423’, 31° 53’ 04.19” -109° 12’ 25.75, 20-viii-2012 (S12-103) 1♂ . Willcox Playa , 4155’, 32° 11’ 55.5” -109° 52’ 42.4”, 29-vii-2015 (S15-104) 1♂ 1♀ ; 4140’ 32° 11’ 57.2” -109° 52’ 32.4”, 28-vi-2009 (S09-55) 4♂ 3♀. Coconino Co., Hwy 89A 1 m S Sedona, 4420’, 30-vi-1994 (S94-36) 1♂ 1♀ . Gila Co., Globe , 3660’, 10- vi-2012 (S12-21) 1♂. Miami , 3500’, 16-vi-1990 (S90-50) 1♂ . Graham Co., Safford , 2920’, 16-vi-1990 (S90-51) 2♂ . Pinaleno Mts , Mt. Graham, Hwy 366 at mile post 116.2, 3700’, 4-vii-1994 (S94-55) 9♂ 6♀ ; Hwy 366 at mile post 120.6, 5000’, 32° 40’ 10.2” -109° 47’ 20.0”, 10-vi-2012 (S12-19) 4♂ ; Hwy 366 at mile post 116.9, 3782’, 32° 41’ 40.1” -109° 44’ 59.0”, 28-vii-2015 (S15-100), 7♂ 9♀ . Hwy 191 S Safford, around Roper Lake State Park, 4-vii-1994 (S94-56) 1♂ . La Paz Co., Alamo Lake , 1400’, 34° 14’ 05.6” -113° 33” 16.1”, 14-ix-2011 (S11-86) 1♂ 1♀. Hwy 60 9.9 m SW Wenden, 1752’, 33° 44’ 15.9” -113° 40’ 35.4, 14-ix-2011 (S11-88) 1♂ 1♀ . Maricopa Co., Painted Rock Petroglyph Site , 16-ix-2006, 2♂. Mohave Co., Kingman , 13-vi-2012 (S12-33) 1♀ . Hualapai Mt. Road SE Kingman, 5135’ 35° 07.859’ -113° 55.142’, 16-vi-2007 (S07-66) 1♂ ; 4918’, 35° 08’ 33.6” -113° 55’ 36.2”, 13- vi-2012 (S12-32) 2♂ 2♀. Pima Co., Ajo, 1760 ’, 20-viii-1998 (S98-74) 2♂ 1♀; 29-vii-2015 (S15-109) 1♂ . Hwy 85 8 m S Ajo, 20-viii-1998 (S98-70) 1♂ 1♀ . Baboquivari Mts. , Brown Canyon, 3960-4080’, 6-vii-1994 (S94-58) 13♂ 6♀ ; 3786’, 31° 45’ 37.0” -111° 31’ 58.8”, 29-vii-2015 (S15-105) 5♂ 13♀. Hwy 85 13m N Ajo, 1255’, 32° 32’ 44.1” -112° 52’ 48.4, 30-vii-2015 (S15-110) 5♂ 4♀. Hwy 86 W Sells, 2276’, 31° 57’ 25.4 -111° 56’ 46.4”, 29-vii- 2015 (S15-108) 2♂ 11♀ . Madera Canyon , 4200’, 26-vii-1990 (S90-80) 5♂ 7♀. Mt. Lemmon Recreation Area , Mile 7.5, 4600’, 27-vii-1990 (S90-82) 8♂ 7♀ ; Mile 9, 5240’, 32° 21’ 21.4” -110° 43’ 34.7”, 27-vi-2009 (S09-51) 1♂ . Tucson , 2375’, 18-vi-1990 (S90-53) 4♂ 2♀ ; 2600’, 27-vii-1990 (S90-84) 2♂; 2780’, 32° 16’ 44” -110° 46’ 18.4”, 27-vi-2009 (S09-52) 1♂. Tucson , 3.9 m N I10, 3040’, 32° 07’ 0.4” -110° 46’ 16.8”, 27-vi-2009 (S09-53) 1♀ . Pinal Co., Oracle , 4260’, 18-viii-1998 (S98-64) 1♂ . Yavapai Co., Agua Fria National Monument , 3200’, 34° 14’ 50.2” -112° 03’ 28.5”, 31-v-2013 (S13-13) 20♂ 12♀ ; 3529’, 34° 15’ 57.5” -112° 05’ 28.9”, 21-viii-2012 (S12-106A) 1♂ 3♀. Cottonwood , 3320’, 30-vi-1994 (S94-37) 1♂ 2♀ . Sedona , Sky Ranch Lodge Motel by airport, 5120’, 34° 51.146’ -111° 47.415’, 15-vi-2007 (S07-61) 1♂ . Hwy 179 eastern outskirts Sedona, 4000’, 31-v-2013 (S13-16) 4♂ . Camp Verde , 3146’, 12-vi-2012 (S12-29) 5♂ 2♀ . 7 m N Prescott , 5060’, 30-vi-1994 (S94-39) 1♂ . Yuma Co., Kofa National Wildlife Refuge , Hwy 95 at mile post 76.8, 10-viii-1988 (S88-88) 1♂ ; near 33° 14’ 59.93 -114° 12’ 54.40”, 1710’, 10-viii-1988 (S88-87) 5♂ 6♀. Telegraph Pass, mile post 18.6, 676’, 32° 39’ 30.7” -114° 20’ 14.2”, 15-ix-2011 (S11-92) 1♂ . California : Imperial Co., Algodones Dunes , 240’, 33° 01’ 13.4” -115° 07’ 25.3”, 15-ix- 2011 (S11-91) 2♀ . San Bernardino Co., Kelso Dunes , 2500’, 13-iii-1985 (S85-28) 2♂ , molt to adult late June. Mt. Pass , 4600’, 4-viii-1991 (S91-71) 1♂ . I15 at Cima Road exit, 3673’, 35° 26’ 30.54” -115° 40’ 24.70”, 22-vii-2016 (S16-24) 6♂ 3♀ . 13.6 m N I40 on road to Providence Mts. State Rec. Area , 3437’, 34° 56’ 29.99” -115° 28’ 53.30, 23-vii-2016 (S16-31) 4♂ 1F3. First wash n I15 on road to Halloran Springs Microwave Station, 2900’ 4-viii-1991 (S91-70) 1♂ 2♀ . Essex , washes around town, 2000’, 22-vii-1990 (S90-69) 6♂ 1♀ . Essex Road , 17.6 m NW Essex, 2900’, 5-vi-1989 (S89-29) 4♂ 2♀ . New Mexico: Bernalillo Co., Albuquerque , Tingley Beach, 4950’, 35° 25’ 16.35 -106° 44’ 33.49”, 29-vi-2015 (S15-49) 2♂ 3♀. Chaves Co., Hwy 82 ~ 33m W Artesia, 4824’, 32° 51’ 55.02 -104° 58’ 51.76”, 30-vi-2015 (S15-56) 4♂ 8♀ . Hwy 380 7.3 m E Roswell, 3960’, 33° 23’ 46.4” -104° 22’ 32.4”, 28-vi-2009 (S09-59) 9♂ 4♀ . Dona Ana Co., Las Cruces , 16-vi-1986 (S86-40) 2♂ ; 27-vi-1988 (S88-58) 1♂. USDA Jornada Experimental Range , ~ 5 m E Las Cruces off Highway 70, 4300’, 32° 28’ 33.33” -106° 44’ 10.86”, 5-vii-1987 (S87- 82), DC Lightfoot, 2♂ 6♀ ; 10-viii-1990 (S90-90) 1♂ 4♀; 3-vii-2015 (S15-75) 6♂ (including holotype) 6♀ . Eddy Co., Hope , 4095’, 32° 48’ 37.73 -104° 44’ 15.00, 30-vi-2015 (S15-57) 3♂ 2♀ . Lincoln Co., Valle del Sol Municipal Golf Course , 5519’, 33° 38’ 18.97 -105° 51’ 37.79”, 30-vi-2015 (S15-55) 1♂ 4♀. Otero Co., Hwy 54 ~ 16 m N Texas border, 4105’, 32° 12’ 20.27” -106° 11’ 37.48”, 3-vii-2015 (S15-74) 10♂ 8♀ . Socorro Co., Acamilla rest stop on E side I25 1 m S La Joya, 4850’, 15-viii-1993 (S93-68) 2♀ . Goat Draw , Los Pinos Mts, 6440’, 34° 22’ 10” -106° 32’ 0”, 16-viii-1993 (S93-69) 14♂ 6♀ . Sevilleta National Wildlife Refuge , Site 222, 5145’, 34° 24’ 30” -106° 56’ 43”, 15-viii-1993 (S93-66) 4♂ 9♀ . Sevilleta National Wildlife Refuge , Rio Salado sand dunes, 4500’, 34° 18.515’ -106° 59.316’, 15-viii-1993 (S93-67) 3♂ 1♀ . Socorro , 4420’, 13-vi-2007 (S07-50) 4♀ ; 29-vi-2015 (S15-54) 5♂ 6♀ . Hwy 60 12 m W intersection Hwy 60 and 47, 4958’, 34° 25’ 16.35” -106° 44’ 33.49”, 29-vi-2015 (S15-50) 7♂ 5♀ . Hwy 60 at milepost 185.8, 6004’, 34° 24’ 12.07” -106° 30’ 29.16”, 29-vi-2015 (S15-53) 4♂ 13♀ . Hwy 380 6.8 m W Car- rizozo, 5160’, 33° 43.276’ -105° 57.471’, 13-vi-2007 (S07-47) 1♂ 1♀ . Hwy 380 37.5 m W Carrizozo, 5200’, 33° 53.139’ -106° 24.785’ (S07-48) 1♂ 2♀ . Texas : Culberson Co., Guadalupe Mts., 4.4 km NE Pine Springs, 5240’ 31° 54.400’ -104° 46.920’, 13-vii-2001 (S01-63) 5♂ 5♀ ; 3.2 km SW Pine Springs , 5360’, 13-vii-2001 (S01-64) 2♂ . Van Horn , 4100’, 6-vi-1991 (S91-48) 4♂ 2♀ .

Sound records only. Arizona: Maricopa Co., Aguila, 2100 ’, 23-vii-1990 (S90-71) . Texas: Hudspeth Co., Cornudas , 4420’, 13-vii-2001 (S01-65).

Derivation of name. Named in appreciation of David C. Lightfoot, who first recognized this taxon during field work at the type locality, and who has helped the senior author with companionship, counsel, and comradery for over 40 years. Plus, for all of those times at 4 AM, when asked to help collect just 2 more crickets (and for which he naively continued to believe that I was serious about only collecting just 2 more!).

Geographic range. ( Fig. 212 ). Ranges from the deserts of southeastern CA through central and southern Arizona and New Mexico, into western Texas.

Habitat. From 73m (Algodones Dunes, S11-91) to 1963m (Goat Draw, S93-69), but usually below 1680m and away from dense tree canopy. Vegetation at the Chihuahuan Desert grassland/scrubland type locality includes soaptree yucca ( Yucca elata ), honey mesquite ( Prosopis glandulosa ), tarbush ( Flourensia cernua ), and burrow grass ( Scleropogon brevifolia ). Elsewhere associated with Arizona interior chaparral, dry, rocky streambed bottoms, and at base of vegetation in California sand dune habitats such as Kelso Dunes (S85-28) and Algodones Dunes (S11-91). When driving lower from the sky island, oak covered localities in Madera Canyon (S90-80) and Hualapai Mt Park (near S07-66 and S12-32), where G. montis exists, at both sites, in good numbers, one enters more open interior chaparral/grassland habitat. Once around 1676m, faster chirping (when compared with G. montis ) G. lightfooti are first heard and females, with their distinctive tegminal bar, can be seen wandering on the road. Males frequently daytime sing from pack rat nests, where they occur with females and nymphs, and can be difficult to collect because of cholla jumping cactus sections sequestered by the pack rats.

At Willcox Playa (S15-104), we collected G. lightfooti and 2 adult G. staccato males singing during afternoon hours from the same dead skirt of a Yucca elata . The only other similarly arboreal western Gryllus taxon is G. sotol at Aguirre Springs, NM (S15-77), where singing G. sotol were collected along with one non-singing G. longicercus male, in sotol ( Dasylirion wheeleri ), during late afternoon.

Life cycle & seasonal occurrence. One generation/year. No egg diapause: Las Cruces (S86-40 & S88-58); Jornada type locality (S87-82); Mt. Graham (S15-100); and Cima, CA (S16-24). D.C. Lightfoot reports (pers. comm. to DBW, October, 2015) the following for the type locality: “They have one generation a year; that year’s eggs hatch late summer (August) with the monsoon rains, slowly grow as nymphs through the winter, probably mostly inactive with cold temps, but active with warm temps, and mature in the late spring (May) and are present as adults through most of the summer. Their life cycle is very similar to G. veletis , except they mature about a month later (May/June vs. April/May) and persist as adults through the summer for a month longer (May/June vs. August/Sept.” This information agrees with our oatmeal trail, collecting efforts for G. lightfooti : 95% of hundreds of individuals seen 31-v-2013 at Agua Fria National Monument, AZ (S13-13); all 10 individuals collected 2-vi-2013 at Apache Pass , AZ (S13-20); 21 of 22 collected 2-vi-2013 from Ft. Bowie, AZ (S13-19); and 5 of 6 collected 5-vi-1989 from Essex, CA (S89-29), were penultimate or late instars when collected.

In the laboratory, all eggs from one female hatch synchronously, when the substrate is kept moist, although nymphs from the same female grow slowly and at different rates from each other, even at temperatures> 25°C. There is no obvious nymphal diapause. On the other hand, when the egg laying substrate is initially not kept moist, and then subsequently moistened, we suspect that eggs may hatch over an extended period of time. In a regime where constant damp sand is provided and nighttime temperatures are kept above 18°C year around, D.C. Lightfoot gets continuous generations and year-round egg laying.

Variation. Body length: Adults living within Y. elata plants generally smaller than those living elsewhere. Cerci length vs. ovipositor length: In females with at least one intact cercus, some 32% have that cercus longer than the tip of the ovipositor in situ. Color: Adult specimens generally with dark bodies and lightly tinged, reddish insides of hind femur. Tegminal color varies from black to light brown, with female tegminal bars most noticeable in the former. Occasional males also with tegminal bars. Hind wing length: Out of some 485 field-collected individuals, only 6 with long hind wings, as follows: CA: Essex (S90-69, 1♂), Cima (S16-24, 1♂); AZ: Ajo (S98-74, 1♂ 1♀), Kofa (S88-87, 2♀). Laboratory-hatched nymphs raised to adult can yield 50-100% long hind winged individuals of both sexes (D.C. Lightfoot, pers. comm., December, 2015).

DNA. multilocus 2016-038, Jornada type locality. Closest relatives are G. sotol and G. transpecos (Gray et al. 2019) .

Discussion. This is one of our most difficult field crickets to identify when away from Yucca elata . G. lightfooti is most easily separated from similar sounding and appearing Gryllus species by habitat—there is little else common in open, desert Southwest grassland/scrublands. Difficulty arises where it occurs in mixed habitats with grasses, sparse tree cover and rocks. For instance, on Hualapai Mt. Road, Arizona (S90-56), 1524m, 8.4 km NW of Hualapai Park, we collected G. lightfooti microsympatric along with G. montis and G. longicercus , although we could distinguish their songs in the field, mostly by a combination of differing pulse and chirp rates. At Goat Draw, New Mexico, (S93-69), we found G. lightfooti microsympatric with G. longicercus in a rocky canyon area of pinyon pine—juniper, Opuntia cactus and some grasslands.

In such Southwest US mixed-habitats, one has to also consider the widespread slow chirpers G. veletis and G. transpecos . Using a combination of distribution maps, song analysis, especially PR and CR, file tooth number, arboreal or not, and presence or absence of a female (and sometimes male) tegminal bar should enable identification of most specimens. But probably not all specimens with certainty. DNA analysis may be required in some cases.

This is our most arboreal western field cricket, with the possible exception of the geographically restricted G. sotol . Where G. lightfooti occurs with Yucca elata , the two seem to be intimately associated with male crickets frequently heard chirping, from within plants, during the hottest part of the day when air temperatures exceed 35°C. They prefer the lower dead-portion skirts of living plants as well as those dead plants on the ground. When dead Yucca plants not present to search, we have (rarely) broken off a living Yucca stalk at ground level, moved the plant into a dirt clearing or onto a nearby road, and by pounding it on the surface, dislodged the crickets, which can then be gathered. We have collected over 15 individuals from one small plant using this technique although they are quick to hop and try to reenter the same Yucca . They obviously see well, even at night with just the light from our headlamps. After sunset, they apparently climb down from Y. elata and walk around, mostly silently on the ground, as seen on Mt. Graham (S15-100), and observed by D.C. Lightfoot in his laboratory cultures (pers. comm. to DBW, 6/2017).

We wonder if this arboreal behavior provides some or all of the following benefits: relief from hot daytime temperatures when compared with ground dwelling field crickets; protection from nocturnally-active, parasitic tachinid flies; and access to a possible food (and water?) source of the plant itself, including preying on other small arthropods living within the skirt. We suspect that females oviposit into the substrate as in other field crickets. Interestingly, G. lightfooti is found in areas of western and northwestern Arizona, southeastern California, and western New Mexico where Yucca elata doesn’t occur. Conversely, perfectly suitable-looking habitat in western Texas has Y. elata but no Arboreal Desert Field Crickets.

Where G. lightfooti occurs away from Y. elata , males can still be found singing from elevated perches at night: Alamo Lake (S11-86), 1 male 1m up in Larrea . When we attempted to capture him, he retreated to base of Larrea into a pack rat nest. Town of Van Horn (S91-48), 1 male singing 4m up in a building. Hualapai Mt. Road (S07-67), 1 male 1.5m up in Palo Verde. Open Mohave Desert near Providence Mts. State Rec. area (S16-31), 1 male 0.5m up in shrub. In towns like Las Cruces and Albuquerque, NM, they are common in suburban neighborhoods, and live in bushes and will enter houses and sing from under roof tiles. Also heard in commercial areas of Socorro, NM (S15- 54).

D.C. Lightfoot notes the following (pers. comm. to DBW, 6/2017) about his 5 th generation, type locality laboratory culture: “They live and sing and mate almost exclusively above the sandy soil bottom of the terrarium, especially during the day. Females wander on the sand floor at night and lay eggs in damp sand. They are highly gregarious, and hang out together, females, males, and nymphs; singing males fight and chase each other a lot but still hang out together. Also, they exhibit a “jerking motion walking behavior” like many camouflaged insects do when they walk (like stick insects and mantises) and they blend in very well with the wood that they live on (tan and brown color). In these aspects, they are totally different from G. veletis which I have also kept in the same terrarium. G. veletis stay on the ground, are not gregarious, and do not ‘jerk walk.’”

Lightfoot reports that juveniles, from the type locality, are distinctly pale tan with mottling. Early instars have black markings on the head, many late instars have dorsum of head black, and all nymphal stages look very similar to Acheta domesticus nymphs. They become dark brown/black as adults, but some still retain the black markings on the head adjacent to the eyes. These color patterns are also seen in G. lightfooti from Southern CA (S16-31). We have not explicitly compared nymphal color patterns between different Gryllus species (but see discussion of striped nymphs under the Longicercus Group, p. 229), although such information might prove phylogenetically relevant.

We have documented parasitoid tachinids Ormia ochracea at the following Arizona localities: near Alamo Lake (S11-86), Mt. Graham (S15-100), Willcox Playa (S15-104), Brown Canyon (S15-105), and Painted Rock Petroglyph Site. We believe that the distorted adult sex ratios (e.g. Brown Canyon, S15-105; W of Sells, S15-108) seen later in the season, where adult females greatly outnumber adult males, may be related to the high incidence of tachinid-killed males ( Sakaguchi & Gray 2011). Also, not singing at night, when gravid tachinid female flies are most active and acoustically searching for singing male Gryllus , should help protect adult males that can still find adult females due to high population densities. DC Lightfoot (pers. comm. to DBW, 6/2017) also reports laboratory males singing more in afternoon and early evening than later at night.

Figures

FIGURE 205. 16S gene tree, indicating multilocus DNA clade based on locality. G. montis Clade 1 samples (orange) mostly have 16S DNA like G. veletis, as do two G. montis Clade 3 samples (blue, both from Mt. Graham). G. montis Clade 2 samples (green) fall into two groups. G. montis Clade 3 samples mostly fall into two geographically separated groups (Hualapai Mtn. near Kingman, AZ, and Sedona, AZ area). G. montis Clade 4 samples have 16S DNA like G. lightfooti. Collection stop numbers: G. montis samples: S07-45 (G1194); S07-54 (G1123, G1154, G1226); S07-58 (G1051, G1216); S07-59 (G1095, G1126); S07-60 (G1052, G1201); S07-62 (G1102, G1151); S07-63 (G1129, G1207); S07-65 (G1103, G1132); S08-50 (G690, G691); S08-51 (G700); S12-18 (G2241, G2249); S12-30 (G2251, G2257); S12-31 (G2262); S12-103 (G2416); S13-17 (G2475, G2857, G2859); S13-18 (G2461, G 2471); S13-21 (G2462, G2464, G2466); S13-36 (G2487, G2489, G2491, G2499); Parker Canyon Lake, Cochise Co., AZ (2004-44, 45, 47, 50, 73); Madera Canyon, Bog Springs, Santa Cruz Co., AZ (2004-118, 2005-003, 012); Madera Canyon, Mt. Wrightson picnic area, Santa Cruz Co., AZ (2010-082); Cave Creek Canyon, Chiricahua Mts., Cochise Co., AZ (2005-115, 122, 127, 131, 133, 136, 137, 143); Miller Canyon, Huachuca Mts., Cochise Co., AZ (2005-169, 176, 179, 181, 192, 193, 194); Reef Camp, Huachuca Mts., Cochise Co., AZ (2012-219, 220, 226). G. veletis samples: S04-107 (G2861, G2862); S07-51 (G1155, G1159); S07-53 (G1120, G1122, G1157); S07-57 (G1056, G1093); S07-72 (G1150); S08-53 (G703, G704); S09-63 (G1355); S15-19 (G2892, G2960, G2999); S15-25 (G2955, G2963, G2968, G2983, G2992); S15-51 (G3099, G3127); S15-53 (G3343); S15-67 (G3144); S15-78 (G3145, G3147). G. lightfooti samples: S09-51 (G1368); S09-59 (G1350); S11-86 (G2205); S11-88 (G2206). G. saxatilis samples: S15-89 (G3201, G3203). G. planeta samples: S15-61 (G3088, G3094, G3114, G3135, G3347); S15-63 (G3130).

FIGURE 207. Five second waveforms of typical calling songs of (A) G. lightfooti, (B) G. sotol, and (C) G. transpecos. (A) G. lightfooti: (R09-69) Cochise Co., AZ (S09-55), at 24°C; (B) G. sotol: (R17-44) from type locality (S17-4), at 24.1°C; (C) G. transpecos: (R09-93) Culberson Co., TX (S09-64), at 25°C.

FIGURE 208. One second spectrograms of typical calling songs of (A) G. lightfooti, (B) G. sotol, and (C) G. transpecos, same males as in Fig. 207.

FIGURE 209. ITS2 gene tree. Collection stop numbers for G. lightfooti samples (orange): S07-61 (G1099, G1206); S07-66 (G1218); S07-67 (G1222); S07-68 (G1133); S09-55 (G1369); S12-19 (G2253); S12-102 (G2404); S12-103 (G2414); S13-13 (G2518, G2528); S13-16 (G2480, G2513, G2519, G2520); S13-18 (G2484, G2696); S13-19 (G2505, G2527, G2533, G2592); S15-49 (G3104); S15-75 (G3061, G3063); S15-100 (G3340); S15-104 (G3328); S15-105 (G3317, G3332); S15-109 (G3333); S15-110 (G3214, G3228, G3235, G3236, G3237); S16-24 (G3435, G3445, G3448, G3450); S16-31 (G3438, G3444); S16- 33 (G3443). Collection stop numbers for G. sotol samples (purple): S15-77 (G3295); S17-4 (G3493, G3497, G3503, G3505, G3508, G3509, G3510, G3511). Collection stop numbers for G. transpecos samples (blue): S07-36 (G1081); S09-64 (G1370, G1384, G1393); S15-61 (G3189, G3279, G3346); S15-64 (G3314, G3348, G3349); S15-65 (G3117, G3138, G3139, G3274, G3327); S15-68 (G3062); S15-69 (G3067); S15-72 (G3059, G3060, G3065, G3069, G3350); S16-2 (G3376, G3377, G3383, G3384, G3389, G3391); S16-3 (G3379, G3385, G3388, G3395); S16-7 (G3375); S16-8 (G3368, G3417); S16-9 (G3396); S16- 10 (G3369); S16-12 (G3398); S16-14 (G3380). Note the two ITS2 clades of G. sotol.

FIGURE 210. Holotype male of G. lightfooti. Female from Pima Co., AZ (S15-108). Note the presence of tegminal bars (arrow), especially on female.

FIGURE 211. Calling song (R09-69) of G. lightfooti from Cochise Co., AZ (S09-55), recorded at 24°C.

FIGURE 212. Known US distribution of G. lightfooti showing distribution of two16S DNA Clades. See discussion of DNA in Barcoding section (p. 16) of this paper.