Larval Eleodes nigropilosus specimens were reared from adults with the following collecting information: “USA: CA: San Diego Co. / Oceanside beach / 33.1865, -117.3778 / 14.May.2011, ADSmith”. A total of 29 eggs and larvae were reared and examined for this study, of which 34 survived to the 2nd instar or beyond. The following description is based on a detailed examination of three 8-11th instar specimens.

TL: 12–15.9 mm, HW: 1.0–1.1 mm, PL: 1.3–1.5 mm, PW: 1.0–1.2 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color light to dark tan, same or nearly the same as body segments; punctation minute, moderately dense, separated by 2–4 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons faintly rugose. Epicranial plates weakly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of four to five long setae along anterior margin near buccal cavity, not confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, not swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2–4 puncture diameters. Labrum not swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae subapical setal row with seven to eight erect setae, anterior margin straight to weakly emarginate. Epipharynx anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular diagonal rows; four subanterior sensory papillae present, arranged as transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetric, left torma smaller. Ligula apex and subapical dorsal surface densely micro-setose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length subequal or greater than maximum width. Antenna three segmented, cylindrical, length of first segment subequal to second.

Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs medium brown, thoracic sternites posterior to prolegs light brown. Prothoracic tergum subquadrate, 1.5× length of meso- or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, forming a gradient from darker brown anteriorly to lighter brown along posterior border. Meso- and metathoracic tergites wider than long, each with a faintly indicated sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout spines ventromedially; prothoracic femur with ventromedial row of three spines, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of three to four spines, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with two ventromedial spines.

Abdomen. Abdominal tergites and sternites light tan with darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous band around segments, bands dark along anterior edge, fading to segment color posteriorly. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex not forming a distinct tooth, moderately clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 14–18 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I–VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 9–12 erect setae.

Eleodes nigropilosus larvae can be separated from the other currently known Eleodes species by having the posterior pigmented band around the abdominal segments forming a color gradient from dark along anterior edge and fading to the color of the rest of the segment posteriorly.

Larval E. wheeleri specimens were reared from adults with the following collecting information: “USA: Arizona: Gila Co. / Tonto Natural Bridge SP / N34.3214, W111.4569 / 11.IX.2010, ADSmith”. A total of 15 eggs and larvae were reared and examined for this study, with all surviving until the 2nd instar or beyond. The following description is based on a detailed examination of five 8-11th instar specimens.

Measurements: TL: 18.0–23.9 mm, PL: 1.6–2.1 mm, PW: 2.1–2.7 mm, HW: 1.8–2.3 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color light tan, same or nearly the same as body segments; punctation minute, dense, separated by 1–2 puncture diameters. Epicranial suture stem length approximately onethird head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons weakly rugose. Epicranial plates weakly rugose dorsally; lateral portions sparsely setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2–4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with seven to eight erect setae, anterior margin straight to weakly emarginate. Epipharynx anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; four anterior sensory papillae present, arranged in two irregular longitudinal rows; four subanterior sensory papillae present arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetric, left torma smaller. Ligula apex lacking microsetae, two long subapical setae present ventrally, eight or more subapical setae present dorsally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, weakly trapezoidal, nearly rectangular. Antenna three segmented, cylindrical, first segment length subequal to second.

Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs light brown, thoracic sternites posterior to prolegs light tan to brown. Prothoracic tergum wider than long, 1.2× or more length of meso- or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with very faint granulated band, nearly concolorous with protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, each with a faintly indicated sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized and sickle-shaped; prothoracic trochanter with two stout spines ventromedially; prothoracic femur with ventromedial row of four spines, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of five to six spines, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with four to five ventromedial spines.

Abdomen. Abdominal tergites and sternites light tan with slightly darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I sparsely clothed in long erect setae along anterior margin. Abdominal laterotergites concolorous with tergites, lacking distinct pigmented margins. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex forming a small tooth, sparsely clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 14–18 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I–VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 11–15 erect setae.

Eleodes wheeleri larvae can be separated from the other currently known Eleodes species by the pentagonal hypopharyngeal sclerome, the lack of a distinct apical tooth on the pygidium, the presence of two long subapical ventral setae on the ligula with eight or more setae present dorsally, and the lateral margins of the protergum with a very faint granulated band, nearly concolorous with protergal disc.

Eleodes wheeleri was recently described (Aalbu et al. 2012) from Tonto Natural Bridge Caverns in Arizona and is known only from the type locality.

Larval Eleodes armatus specimens were reared from adults with the following collecting information: “USA: CA: Riverside Co. / Palm Desert, 38th Ave / off Washington St. / N33.7721, W116.3071 / 10.X.2010, ADSmith”; “USA: AZ: Maricopa Co. / Phoenix, E. Eugie Ave / & 7th St. N33°36.665’ / W112°03.849’, 418 m., / 25 May 2011, R.Dornburg.” A total of 1805 eggs and larvae were reared and examined for this study, with 128 persisting to the 2nd instar or later. The following description is based on a detailed examination of fifteen 8-11th instar specimens

TL: 21.0–35.0 mm, HW: 2.4–3.8 mm, PL: 2.4–3.4 mm, PW: 2.9–4.6 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color ferruginous, more heavily pigmented than body segments; punctation minute, dense, separated by 1–2 puncture diameters. Epicranial suture stem length approximately one-fourth head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons weakly rugose. Epicranial plates weakly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2–4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with seven to eight erect setae, subapical setal row with seven to eight erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 9B) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular rows, each with two posterior papillae and one near the anterior margin; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetric, left torma smaller. Ligula apex lacking microsetae, two long subapical setae present ventrally, eight or more subapical setae present dorsally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length less than maximum width. Antenna three segmented, cylindrical, first segment longer than second.

Thorax. Thoracic tergites light tan to ferruginous, prothoracic sternite anterior to legs ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum wider than long, 1.2× or more length of meso-, metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout spines ventromedially; prothoracic femur with ventromedial row of six to ten spines, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of eight to eleven spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with five to seven ventromedial spines.

Abdomen. Abdominal tergites and sternites light tan to ferruginous, with slightly darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I moderately clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex forming a distinct tooth, sparsely clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 22–24 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I–VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 11–15 erect setae.

Eleodes armatus larvae can be separated from the other currently known Eleodes species by presence of an apical tooth on the pygidium and the absence of stout spiniform setae on the anterolateral margins of the epipharnyx.

Larval Eleodes caudiferus specimens were reared from adults with the following collecting information: “USA: Arizona: Navajo Co. / dunes ~4mi N Chilchinbito / off route 59, el. 1738m / N36.58143, W110.06973 / 26.August.2010, ADSmith”. A total of 85 eggs and larvae were reared and examined for this study, of which 53 survived untill the 2nd instar or later. The following description is based on a detailed examination of eleven 3-5th instar specimens.

TL: 7.8–12.8 mm, HW: 1.0–1.4 mm, PL: 1.0–1.8 mm, PW: 1.3–1.7 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width narrower than prothorax; sides rounded; strongly constricted before occipital foramen; color dark tan, same or nearly the same as on body segments; punctation minute, moderately dense, separated by 2–4 puncture diameters. Epicranial suture stem length approximately one-fourth to one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons rugose. Epicranial plates rugose dorsally; lateral portions densely setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2–4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with 10–14 erect setae, subapical setal row with 10–14 erect setae, anterior margin straight to weakly emarginate. Epipharynx anterior setal row with eight or more stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular rows; four subanterior sensory papillae present arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae symetrical or weakly asymmetric. Ligula apex densely microsetose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length less than maximum width. Antenna three segmented, cylindrical, first segment subequal to second.

Thorax. Thoracic tergites ferruginous, prothoracic sternite anterior to legs ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum subquadrate, 1.5× length of meso- or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, with sclerotized transverse line on anterior fifth absent, dense transverse band of short setae present near anterior margins of both tergites. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout spines ventromedially; prothoracic femur with ventromedial row of five to six spines, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of five to six spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with row of three ventromedial spines.

Abdomen. Abdominal tergites and sternites light tan to ferruginous, with slightly darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I tomentose in anterior third, setae denser along near lateral margins. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex attenuated and sclerotized, rarely forming a small tooth, sparsely clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 28–38 socketed spines present, forming two or three irregular rows around posterior two-thirds to one half of segment, narrowing to single row around apex. Abdominal sternites I–VIII with longitudinal tomentose bands present along lateral margins. Pygopods short, subconical, each with 17–24 erect setae.

Eleodes caudiferus larvae can be separated from the other currently known Eleodes species by the presence of longitudinal tomentose bands along the lateral margins of abdominal sternites I–VIII.

Larval Eleodes hispilabris specimens were reared from adults with the following collecting information: “USA: TX: El Paso County / El Paso, sand dunes off / Hwy 180/Montana Ave. / N31.82327, W106.13234 / 21-22.VIII.2010, ADSmith”. A total of 46 eggs and larvae were reared and examined for this study, with 36 surviving until the 2nd instar or beyond. The following description is based on a detailed examination of five 8–11th instar specimens.

TL: 21.0–32.0 mm, PL: 2.6–3.2 mm, PW: 3.0–3.7 mm, HW: 2.4–3.1 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width narrower than prothorax; sides rounded; strongly constricted before occipital foramen; color ferruginous, more heavily pigmented than body segments; punctation minute, dense, separated by 1–2 puncture diameters. Epicranial suture stem length approximately one-fourth head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons rugose. Epicranial plates rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of four to five long setae along anterior margin near buccal cavity, not confluent with setae on lateral portions of plates, with a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation dense, separated by 1–2 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with 10–14 erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 9C) anterior setal row with eight or more stout spiniform setae, anterolateral margins with stout spinose setae; six anterior sensory papillae present, arranged in two irregular rows, each with two posterior papillae and one near the anterior margin; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; seven to eight posterior sensory papillae present, arranged in an irregular cluster. Tormae strongly asymmetric, left torma larger. Ligula apex lacking microsetae, two long subapical setae present ventrally, eight or more subapical setae present dorsally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length less than maximum width. Antenna three segmented, cylindrical, first segment longer than second.

Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs light tan to ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum wider than long, 1.2× or more length of meso- or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with one or two stout ventromedially spines; prothoracic femur with ventromedial row of six to ten spines, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of eight to eleven spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with four to five ventromedial spines.

Abdomen. Abdominal tergites and sternites light tan, with slightly darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex forming a distinct tooth, sparsely clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 17–23 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I–VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 9–12 erect setae.

Eleodes hispilabris larvae can be separated from the other currently known Eleodes species by the presence of an apical tooth on the pygidium, stout spiniform setae on the anterolateral margins of the epipharnyx, and a row of 6–10 ventromedial spines on the prothoracic femur.

Larval Eleodes tenuipes specimens were reared from adults with the following collecting information: “USA: TX: El Paso County / El Paso, sand dunes off / Hwy 180/Montana Ave. / N31.82327, W106.13234 / 21-22.VIII.2010, ADSmith”. Atotal of five eggs and larvae were reared and examined for this study. The following description is based on a detailed examination of one late instar specimen.

Measurements: TL: 39.0 mm, HW: 4.1 mm, PL: 4.0 mm, PW: 4.8 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color ferruginous, more heavily pigmented than body segments; punctation minute, dense, separated by 1–2 puncture diameters. Epicranial suture stem length approximately one-fourth head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons rugose. Epicranial plates rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation dense, separated by 1–2 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with 10–14 erect setae, anterior margin straight to weakly emarginate. Epipharynx anterior setal row with eight or more stout spiniform setae, anterolateral margins with stout spinose setae; six anterior sensory papillae present, arranged in two irregular rows, each with two posterior papillae and one near the anterior margin; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae strongly asymmetric, left torma smaller. Ligula apex lacking microsetae, two long subapical setae present ventrally, eight or more subapical setae present dorsally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length less than maximum width. Antenna three segmented, cylindrical, first segment longer than second.

Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum wider than long, 1.2× or more length of meso- or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with one stout ventromedially spine; prothoracic femur with ventromedial row of 13–14 spines, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of eight to eleven spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with five to seven ventromedial spines.

Abdomen. Abdominal tergites and sternites light tan, with slightly darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex forming a distinct tooth, sparsely clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 27 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I–VIII lacking longitudinal tomentose bands along lateral margins.

Eleodes tenuipes larvae can be separated from the other currently known Eleodes species by the presence of an apical tooth on the pygidium, stout spiniform setae on the anterolateral margins of the epipharnyx, and a row of 13–14 ventromedial spines on the prothoracic femur. It is further differentiated from Eleodes hispilabris by having a row of three ventromedial spines on the mesotarsus and having the ventral portion of the epicranial plates with a row of six or more long setae along anterior margin near buccal cavity, confluent with setae on lateral portions of plates.

Five eggs or early instar larvae were initially placed in a rearing chamber on 25 September 2010, though by the first sifting only one specimen was found. The last specimen thrived until 27 January 2011 when it was preserved for this study.

Larval Eleodes tribulus specimens were reared from adults with the following collecting information: “USA: AZ: Pinal Co. / I-10W Rest Area, mm183 / 33.029288, -111.771716 / 02 May 2011, ADSmith”. A total of 824 eggs and larvae were reared and examined for this study, of which 134 survived until the 2nd instar or later. The following description is based on a detailed examination of ten 8-11th instar specimens.

TL: 13.0–19.0 mm, HW: 1.5–2.2 mm, PL: 1.2–2.7 mm, PW: 1.3–2.7 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides angular; strongly constricted before occipital foramen; color light tan to medium brown, more heavily pigmented than body segments; punctation minute, moderately dense, separated by 2–4 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons rugose. Epicranial plates weakly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2–4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae subapical setal row with six to seven erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 9D) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; five to six anterior sensory papillae present, arranged in two irregular longitudinal rows or an irregular cluster; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; seven to eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetric, left torma larger. Ligula apex and subapical dorsal surface densely micro-setose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length subequal or greater than maximum width. Antenna three segmented, cylindrical, first segment length subequal to second.

Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs medium brown, thoracic sternites posterior to prolegs light brown. Prothoracic tergum subquadrate, 1.5× length of meso- or metaterga; lateral margins with distinct granulated band, darker than protergal disc; anterior transverse striated band present, darker than tergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, each with a faintly indicated sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout spines ventromedially; prothoracic femur with ventromedial row of two spines and three to five longer setae, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of three to four spines, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with three ventromedial spines.

Abdomen. Abdominal tergites and sternites light tan with darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I moderately clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex not forming a distinct tooth, moderately clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 8–14 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I–VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 11–15 erect setae.

Eleodes tribulus larvae can be separated from the other currently known Eleodes species based on the pentagonal hypopharyngeal sclerome, lack of a caudal tooth on the pygidium, presence of 8–14 marginal spines on the pygidium, and the angular, nearly straight sides of the head capsule.

Larval Eleodes extricatus specimens were reared from adults with the following collecting information: “USA: TX: El Paso County / El Paso, sand dunes off / Hwy 180/Montana Ave. / N31.82327, W106.13234 / 21–22.VIII.2010, ADSmith”, “USA: Arizona: Navajo Co. / dunes ~4mi N Chilchinbito / off route 59, el. 1738m / N36.58143, W110.06973 / 26.August.2010, ADSmith”, “USA: AZ: Graham Co. / Pinaleño Mnts, Hospital Flat Camp / N32°39’58.0”, W109°52’30.9” / el.9000’ 22.Aug.2010 / ADSmith”, “USA: Arizona: Gila County / E. Verde River off NF-272 / N34.303, W111.3496 / 27.August.2010, ADSmith”. Approximately 219 eggs and larvae were reared and examined for this study, with 150 surviving until the second instar or beyond. The following description is based on a detailed examination of thirteen 8–11th instar specimens.

Measurements: TL: 15.4–33.3 mm, PL: 2.4–3.8 mm, PW: 2.2–3.8 mm, HW: 2.0–3.0 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color light tan, same or nearly the same as body segments; punctation minute, dense, separated by 1–2 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons faintly rugose. Epicranial plates faintly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen or not, unicolorous, minutely punctate, punctation dense, separated by 1–2 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with six to seven erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 10A) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae symmetrical or weakly asymmetrical with left torma smaller. Ligula apex densely microsetose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length less than maximum width. Antenna three segmented, cylindrical, first segment longer than second.

Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum subquadrate, 1.5× length of meso- or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout ventromedially spines; prothoracic femur with ventromedial row of two spines and three to five longer setae, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of three to four spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with four to five ventromedial spines.

Abdomen. Abdominal tergites and sternites light tan, with slightly darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex lacking a distinct tooth, sparsely clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 17–23 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I–VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 11–15 erect setae.

Eleodes extricatus larvae can be separated from the other currently known Eleodes species based on the pentagonal hypopharyngeal sclerome, small or absent apical tooth on the pygidium, lateral margins of prothoracic tergum with a distinct granulated band, and having antennal segment I longer than antennal segment II.

Eleodes extricatus is a widespread species found on dunes and at high elevations. Specimens from Arizona and Texas showed no population differences in the larval stage. Adults varied in the presence or prominence of muricate tubercles on the elytra.

Larval Eleodes anthracinus specimens were reared from adults with the following collecting information: “USA: AZ: Maricopa Co. / Eugie Ave & 7th St. / 25 Oct. 2011, R. Dornburg.” A total of 28 eggs and larvae were reared and examined for this study, of which all survived until the 3rd instar or later. The following description is based on a detailed examination of four 8–11th instar specimens.

TL: 23.8–28.1 mm, HW: 2.3–2.4 mm, PL: 2.0–2.4 mm, PW: 2.5–2.8 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color medium brown to brown-grey, nearly as on body segments; minute punctation moderately dense dorsally. Epicranial stem approximately one-third head capsule length; frontal arms U-shaped, not obscured by sculpturing. Frons and dorsal portion of epicranial plates faintly rugose; lacking non-primary setae. Lateral portions of epicranial plates moderately setose; setae golden, erect, length equal to or longer than antennal segment 2; ventral portions of epicranial plates with a row of four long setae along anterior margin near buccal cavity with a patch of short setae medially forming a triangular pattern with its base near the anterior margin; two stemmata present on each plate, pigmented spots often faded. Clypeus trapezoidal; not swollen, moderately punctate, darker medially in basal half. Labrum not swollen, basal half more darkly pigmented; sides rounded; two transverse rows of seven to eight erect setae present medially and subapically; anterior margin straight. Epipharynx (Fig. 3) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular diagonal rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in two irregular rows.Tormae asymmetrical, left torma smaller. Ligula with four long setae near apex. Hypopharyngeal sclerome trapezoidal. Gula distinct, trapezoidal, widest in basal half. Antenna three segmented, cylindrical; first segment longer than second.

Thorax. Grey-brown to medium brown dorsally and anterior to legs on prothoracic sternite, tan on rest of sternites; lighter transverse striated band present along anterior fourth of prothoracic tergum; thin darkly sclerotized transverse line present on anterior fifth of meso- and metathoracic tergites; striated bands present along posterior 5th of all thoracic tergites, color forming a gradient from darker brown anteriorly to lighter brown along posterior border. Eight evenly arranged setae present on dorsal surface of each thoracic terga, lateral margins more densely setose. Prothoracic tergum subquadrate, 1.5× length of meso- or metaterga; lateral margins lacking pigmented band. Meso- and metaterga wider than long, lacking pigmented bands along lateral margins; mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracle; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; trochanter with row of two stout spines and two longer setae ventromedially, tibia with ventromedial row of two spines and four to five longer setae, tarsus with ventromedial row of four spines. Dorsal surface of protibia (at rest) with faintly indicated basal sclerotized band; dorsal surface of protarsus slightly more sclerotized than ventral surface.

Abdomen. Tergites grey-brown to medium brown dorsally, lightening towards lateral margins, sternites light to dark tan; transverse striated bands not visible on abdominal sternites, barely visible on posterior 5th of terga I–VIII, nearly concolorous with rest of tergites. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline, abdominal segments II–VIII each with two sparse transverse bands of long erect setae, posterior margin of segment 8 denser setal band. Abdominal laterotergites concolorous with tergites, lacking distinct pigmented margins. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, sparsely clothed in short and mid length erect setae, dorsally more sclerotized in apical two-thirds with faint maculations; marginal row of 14–18 socketed spines present apical half, apex not forming distinct sclerotized projection. Pygopods short, subconical, each with 11–15 erect spines.

Variation. Little variation was observed between specimens beyond the number of spines on the legs and pygidium, and the overall degree of sclerotization.

Eleodes anthracinus larvae can be separated from most currently known Eleodes species based on their darker dorsal coloration on all segments, the absence of pigmented bands along the lateral margins of the thoracic terga, and the lack of a distinct sclerotized tooth at the apex of the pygidium. They can be distinguished from Eleodes carbonarius larvae by their lighter ventral segments and lack of distinct posterior pigmented bands on the abdominal terga. Larvae of Eleodes tricostatus (Say), another species in the subgenus Melaneleodes, are mentioned as being “nearly black” by McColloch (1918). However, no other diagnostic characters are mentioned that would separate them from the other Eleodes anthracinus or Eleodes carbonarius.

Larval Eleodes carbonarius specimens were reared from adults with the following collecting information: “USA: CO: Montezuma Co. / Ute RA off Hwy 160 / 37.3535, -108.44385 / 05 Jun 2011, ADSmith”. A total of 129 eggs and larvae were reared and examined for this study, with 45 surviving until the 2nd instar or later. The following description is based on a detailed examination of five 8–11th instar specimens.

TL: 15.5–26 mm, HW: 2.3–3.0 mm, PL: 1.9–2.5 mm, PW: 3.0–3.5 mm.

Head. Prognathous, weakly flattened, narrower than prothorax; sides rounded, strongly constricted before occipital foramen; color ferruginous to dark brown, nearly as on body segments; minute punctation moderately dense dorsally. Epicranial stem approximately one-third head capsule length; frontal arms U-shaped, not obscured by sculpturing. Frons and dorsal portion of epicranial plates faintly rugose; lacking non-primary setae. Lateral portions of epicranial plates moderately setose; setae golden, erect, length equal to or longer than antennal segment 2; ventral portions of epicranial plates with a row of four to five long setae along anterior margin near buccal cavity and a patch of short setae medially forming a triangular pattern with its base near the anterior margin; two stemmata present on each plate, pigmented spots often faded. Clypeus trapezoidal; not swollen, moderately punctate, darker medially in basal half. Labrum not swollen, basal half more darkly pigmented; sides rounded; two transverse rows of six to seven erect setae present medially and subapically; anterior margin straight to weakly emarginate. Epipharynx (Fig. 9A) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular diagonal rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetrical, left torma larger. Hypopharyngeal sclerome trapezoidal. Ligula with four long setae near apex. Gula distinct, trapezoidal, widest in basal half. Antenna three segmented, cylindrical; first segment longer than second.

Thorax. Dark brown to ferruginous dorsally and anterior to legs on prothoracic sternite, lighter brown on rest of sternites; distinct longitudinally striated band present along anterior fourth of prothoracic tergum; thin darkly sclerotized transverse line present on anterior fifth of meso- and metathoracic tergites; striated bands present along posterior 6th of all thoracic tergites, darker than rest of surface. Eight evenly arranged setae present on dorsal surface of each thoracic terga, lateral margins more densely setose. Prothoracic tergum wider than long, 1.5× length of meso- or metaterga; lateral margins lacking pigmented band. Meso- and metaterga wider than long, lacking pigmented bands along lateral margins; mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracle; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; trochanter with two stout spines ventromedially, tibia with ventromedial row of three to four spines and four to five longer setae, tarsus with ventromedial row of five spines. Dorsal surface of protibia (at rest) with basal sclerotized band; dorsal surface of protarsus more sclerotized than ventral surface.

Abdomen. Tergites dark brown to ferruginous, concolorous or lightly lighter than tergites; longitudinally striated bands not visible on abdominal sternites, distinct on posterior 5th of terga 1–8. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline, abdominal segments 2–8 each with two sparse transverse bands of long erect setae, posterior margin of segment 8 denser setal band. Abdominal laterotergites concolorous with tergites, lacking distinct pigmented margins. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, sparsely clothed in short and mid length erect setae, apical two-thirds with faint maculations; marginal row of 18–20 socketed spines present apical half, apex not forming distinct sclerotized projection. Pygopods short, subconical, each with 9–12 erect spines.

Variation. Little variation was observed between specimens beyond the number of spines on the legs and pygidium, and the overall degree of sclerotization.

Eleodes carbonarius larvae can be separated from most currently known Eleodes species their darker dorsal coloration on all segments, the absence of pigmented bands along the lateral margins of the thoracic terga, and the lack of a distinct sclerotized tooth at the apex of the pygidium. They can be further distinguished from Eleodes anthracinus larvae as outlined in that species diagnosis.

Eleodes carbonarius adult morphology is notoriously variable across the species range and even within populations. Nine subspecies are currently recognized (Triplehorn and Thomas 2011). The specimens examined were all reared from a few females of Eleodes carbonarius knausi Blaisdell collected at a single locality. Larval characters showed little variation; however, this may change as more specimens are reared from other localities and subspecies.

Larval Eleodes goryi specimens were reared from adults with the following collecting information: “USA: TX: Hidalgo County / Bentsen-Rio Grande Valley / State Park, fm2062 Mission / N26°10.37’, W098°22.93’ / 02.Sept.2011, Aaron Smith”. A total of 460 eggs and larvae were reared and examined for this study, with 25 surviving until the 2nd instar or beyond. The following description is based on a detailed examination of three 8–11th instar specimens.

TL: 25.0–25.4 mm, HW: 2.0–2.1 mm, PL: 2.0–2.1 mm, PW: 2.2–2.4 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color ferruginous to dark brown, more heavily pigmented than body segments; punctation minute, moderately dense, separated by 2–4 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms U-shaped, not obscured by sculpturing. Frons faintly rugose. Epicranial plates faintly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2–4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with six to seven erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 10B) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae strongly asymetrical with left torma larger. Ligula apex densely microsetose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length subequal or greater than maximum width. Antenna three segmented, cylindrical, first segment subequal to second.

Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs ferruginous to medium brown, thoracic sternites posterior to prolegs medium brown. Prothoracic tergum wider than long, 1.2× or more length of meso- or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout ventromedially spines; prothoracic femur with ventromedial row of two spines and three to five longer setae, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of three to four spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with three ventromedial spines.

Abdomen. Abdominal tergites and sternites 1–7 light tan, with slightly darker transverse striated bands present along posterior margins of segments I–VIII, forming near contiguous unicolorous band around segments. Abdominal tergite 8 more darkly pigmented than preceding segments. Abdominal sternite I moderately clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex lacking a distinct tooth, moderately clothed in short and mid length erect setae, dorsally more sclerotized in apical two-thirds with faint maculations; marginal row of 18–20 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I–VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 11–15 erect setae.

Eleodes goryi larvae can be separated from the other currently known Eleodes species based on the darkly pigmented eighth and ninth abdominal tergites. It is further distinguished by the pentagonal hypopharyngeal sclerome, lack of a caudal tooth on the pygidium, and the presence of 3–4 ventromedial spines on the protibia.

Larval Eleodes subnitens specimens were reared from adults with the following collecting information: “USA: Arizona: Gila Co. / Tonto Natural Bridge SP / N34.3214, W111.4569 / 11.IX.2010, ADSmith”. A total of 7 eggs and larvae were reared and examined for this study, of which four survived until the 2nd instar or later. The following description is based on a detailed examination of two 8–11th instar specimens.

TL: 23.1–30.8 mm, HW: 2.0–3.0 mm, PL: 2.0–2.9 mm, PW: 2.2–3.1 mm.

Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color ferruginous, more heavily pigmented than body segments; punctation minute, moderately dense, separated by 2–4 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons faintly rugose. Epicranial plates faintly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker in apical half, minutely punctate, punctation moderately dense, separated by 2–4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with seven to eight erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 10C) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymetrical with left torma smaller. Ligula apex densely microsetose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length subequal or greater than maximum width. Antenna three segmented, cylindrical, first segment subequal to second.

Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum wider than long, 1.2× or more length of meso- or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso- and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout ventromedially spines; prothoracic femur with ventromedial row of two spines and three to five longer setae, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of five to six spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with four to five ventromedial spines.

Abdomen. Abdominal tergites and sternites I–VIII light tan, with slightly darker transverse striated bands present along posterior margins8, forming near contiguous unicolorous band around segments. Abdominal sternite I moderately clothed in long erect setae to posterior pigmented band. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex lacking a distinct tooth, moderately clothed in short and mid length erect setae, dorsally sclerotization uniform throughout, lacking maculations; marginal row of 18–20 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites 1–8 lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 17–24 erect setae.

Eleodes subnitens larvae can be separated from the other currently known Eleodes species by the pentagonal hypopharyngeal sclerome, prothoracic tergum wider than long, 8th and 9th abdominal tergites not darker than proceeding segments, lack of a caudal tooth on the pygidium, and the presence of 5–6 ventromedial spines on the protibia.

Larval Eleodes pilosus specimens were reared from adults with the following collecting information: “NEVADA: Washoe Co. / N39°16.427’, W119°47.070’ / November 14, 2011 / P.Skelley, sifting lakeside dunes”. A total of 208 eggs and larvae were reared and examined for this study, of which 94 survived until the 2nd instar or later. The following description is based on a detailed examination of nine 8–11th instar specimens.

TL: 14.2–26.0 mm, PW: 1.7–3.3 mm, PL: 1.4–3.4 mm, HW: 1.6–2.6 mm.

Head. Prognathous, weakly flattened, narrower than prothorax; sides rounded, strongly constricted before occipital foramen; color nearly as in body segments. Epicranial stem short, one-fourth head capsule length; frontal arms U-shaped, partially obscured by sculpturing. Frons and dorsal portion of epicranial plates rugose; sparsely setose; densely punctate, punctures minute, lacking setae. Ventrolateral portions of epicranial plates densely setose; setae golden, erect, most longer than antennal segment 2, interspersed with shorter setae; two stemmata present on each plate, pigmented spots often faded. Clypeus trapezoidal; swollen, weakly transversely raised medially; moderately punctate, rugose in basal half. Labrum slightly swollen, basal fourth darkly pigmented; sides rounded, minutely tomentose; two transverse rows of six to eight erect setae present medially and subapically; anterior margin straight. Epipharynx (Fig. 10D) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; seven anterior sensory papillae present, arranged in two irregular longitudinal rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; seven posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetrical, left side torma smaller with or without a small spine near emergent edge. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half. Antenna three segmented, cylindrical; first segment longer than second.

Thorax. Light to dark tan, darker longitudinally striated bands present on anterior margin of prothoracic tergum and posterior 5th of all thoracic tergites. Sparsely setose along dorsal margins of terga near striated bands, lateral margins more densely setose. Prothoracic tergum subquadrate, 1.5× length of meso- or metaterga; lateral margins with pigmented band along entire length. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracle; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Prothoracic leg slightly longer, much thicker than meso- and metathoracic legs; tarsungulus strongly sclerotized, sickle-shaped; trochanter with two stout spines ventromedially, tibia and tarsus each with a ventromedial row of four to seven spines, number of spines often differing between prolegs. Dorsal surface of protibia (at rest) with basal sclerotized band; dorsal surface of protarsus sclerotized.

Abdomen. Light to dark tan, darker longitudinally striated bands present on posterior 5th of segments I–VIII. Abdominal sternite I moderately clothed in long erect setae, sparser medially, extending to posterior pigmented band, abdominal tergite I and segments II–VIII sparsely clothed in short to mid length setae. Lateral margins of abdominal laterotergites I–VIII darkly pigmented, ventral margin with two pigmented bands. Abdominal segment IX (pygidium) triangular in dorsal view, moderately clothed in long erect setae; marginal row of 10–20 socketed spines present apical half, apex not forming distinct sclerotized projection. Pygopods short, subconical, each with 16–20 erect spines.

Eleodes pilosus larvae can be separated from the other currently known Eleodes species by the pentagonal hypopharyngeal sclerome, lack of a caudal tooth on the pygidium, presence of 8–14 marginal spines on the pygidium, subquadrate prothoracic tergum, and having abdominal sternite I moderately clothed in long erect setae to posterior pigmented band.