Larvae of the genus Eleodes (Coleoptera, Tenebrionidae): matrix-based descriptions, cladistic analysis, and key to late instars.

Darkling beetle larvae (Coleoptera, Tenebrionidae) are collectively referred to as false wireworms. Larvae from several species in the genus Eleodes are considered to be agricultural pests, though relatively little work has been done to associate larvae with adults of the same species and only a handful of species have been characterized in their larval state. Morphological characters from late instar larvae were examined and coded to produce a matrix in the server-based content management system mx. The resulting morphology matrix was used to produce larval species descriptions, reconstruct a phylogeny, and build a key to the species included in the matrix. Larvae are described for the first time for the following 12 species: Eleodes anthracinus Blaisdell, Eleodes carbonarius (Say), Eleodes caudiferus LeConte, Eleodes extricatus (Say), Eleodes goryi Solier, Eleodes hispilabris (Say), Eleodes nigropilosus LeConte, Eleodes pilosus Horn, Eleodes subnitens LeConte, Eleodes tenuipes Casey, Eleodes tribulus Thomas, and Eleodes wheeleri Aalbu, Smith & Triplehorn. The larval stage of Eleodes armatus LeConte is redescribed with additional characters to differentiate it from the newly described congeneric larvae.

Introduction

Species of the genus are among the most iconic and recognizable insects of the western United States. Flightless, almost always black in color, and medium to large sized (~10-50 mm), are perhaps most closely associated with head-standing. While this behavior, linked to the exudation or squirting of a concoction of noxious defensive chemicals from paired reservoirs near the tip of the abdomen, is not unique to , it has been the source of common names for the genus such as stink or circus beetles.

Larvae of the family Tenebrionidae are known as false wireworms. Feeding on seeds, roots, and subterreanean stems, a number of them are considered agricultural pests, including (Say, 1824), (Say, 1824), (Say, 1824), (Say, 1824), and (Say, 1824) (Calkins and Kirk 1975). A summary of species considered agricultural pests and the crops they attack is given by Allsopp (1980). In spite of ecological and agricultural interest in false wireworms, and their potential contribution of a whole suite of characters for phylogenetic and taxonomic studies, knowledge of their morphology, development, and habits remain limited. Doyen (1988) estimated that approximately 240 genera and 300 species of darkling beetle larvae had been described which, as far as we know, remains a reasonable approximation. Of the 190+ currently valid species, only seven have been previously described in the larval stages.

Taxonomic history

Relatively few larvae have been described or characterized (Table 1). Gissler (1878) provided the first larval descriptions in the genus for Eschscholtz, 1833 and (Mannerheim, 1843). Hyslop (1912) described the larvae of Blaisdell, 1909 (then listed as a subspecies of ) and Mannerheim, 1843 from the Pacific Northwest. McColloch (1918) described (Say), 1824. Wade and St. George (1923) described (Say, 1824), followed closely by illustrations, without additional descriptions, of the pygidia of (Say, 1824), (Say, 1824), and by St. George (1924). Blaisdell (1909) redescribed the larvae of in greater detail and described the pupa of Eschscholtz, 1833. The most recent larval description was provided by Thomas (1984) for LeConte, 1851. In most cases, these descriptions are insufficient to reliably diagnose larvae to species.

Table 1.

Previous publications describing or illustrating larvae.

Species	Publication	Remarks
Eleodes armatus (LeConte), 1851	Thomas 1984	egg, larva, and pupa described, larva and pupa imaged
Eleodes dentipes (Eschscholtz), 1833	Gissler 1878; Blaisdell 1909	larva briefly described in Gissler (1878); larva redescribed and illustrated in Blaisdell (1909)
Eleodes giganteus (Mannerheim), 1843	Gissler 1878	egg and larva characterized; larva illustrated
Eleodes opacus (Say), 1824	St. George 1924	pygidium imaged; no description
Eleodes pimelioides (Mannerheim), 1843	Hyslop 1912	egg, larva, and pupa described; pygidium of larva imaged
Eleodes suturalis (Say), 1824	Wade and St. George 1923	egg, larva, and pupa described, larval natural history discussed, egg and pupa imaged
Eleodes tricostatus (Say), 1824	McColloch 1918; St. George 1924	egg, larva, and pupa briefly characterized, larval natural history discussed; right mandible and pygidium of larva imaged in St. George (1924)
Eleodes vandykei Blaisdell, 1909	Hyslop 1912	egg, larva, and pupa described, egg, larva, and pupa imaged; species listed as Eleodes letcheri vandykei

Matrix-based taxonomy

A number of modern taxonomic works on insects have produced descriptions based on matrices of morphological characters, including Winterton (2009), Yoder et al. (2009), PageBreakTalamas et al. (2011), and Mullins et al. (2012) to name a few. The advantages of this methodology include a structured and explicit differentia between the character states exhibited by each species or other taxonomic units, the ability to score new species into the matrix, an option to further utilize the matrix for phylogenetic analyses, and the capability to turn a matrix into a multi-entry key or link it to other data sources, such as an anatomy ontology. The present work is intended as a first step to describe the larvae of the genus , define important characters for species and subgeneric differentiation, provide a first glimpse into evolutionary relationships within the genus, and provide a platform to link character states to the developing Coleoptera Anatomy Ontology (ColAO).

Methods and Materials

Morphological parameters. Measurements were taken using either digital calipers, an optical micrometer attached to a Nikon SMZ 1500 stereomicroscope, or measurement scales set in Photoshop specific to the camera and lens used to take measurements from images. was measured from the anterior edge of the clypeus to the dorsomedial apex of abdominal segment IX. and were measured dorsally across the widest and longest points on the segment respectively, was measured dorsally across the widest portion of the head (generally near the apex of the cranial stem). Terminology primarily follows Lawrence (1991). Dissections were performed using fine forceps and a sharpened #0 insect pin.

Total length

Prothoracic width

length

head capsule width

Photographs of specimens or characters were taken using a BK Plus or Passport Imaging system (R. Larimer, www.visionarydigital.com). Montaged images were assembled using Zerene Stacker (zerenesystems.com/stacker/) and backgrounds were cleaned up in Adobe Photoshop CS5. Confocal laser images were taken on a Zeiss LSM 710.

Rearing. Adult specimens were hand collected from throughout the southwestern United States. Specimens were maintained in separate plastic containers for each species, locality, and collecting event on a substrate of sand. Every one to two weeks, containers were sifted for eggs and larvae. Larvae were reared on a sand/food substrate in plastic containers, with either plaster of Paris at the bottom watered though a vinyl tube to maintain a moisture gradient (Brown 1973) or with daily watering. A study detailing rearing regimes, instar numbers, and life histories for the reared species is forthcoming (Dornburg, Smith & Wheeler, in preparation).

Matrix-based descriptions. To allow for easier direct comparisons between larvae of different species and provide a framework for the addition of larvae from more species in the future, descriptions were produced from a morphological character matrix and edited for traditional telegraphic description format. The character matrix was built in mx (Yoder et al. 2010), based on 86 morphological characters (Appendices 1 and 2). Mx was also used to produce the initial descriptions. Single state characters included in the descriptions, were also included in the matrix. All specimens scored in the matrix were classified as late (7th–11th) instar larvae based on their size or observed number of molts. The one exception was , in which only third instar larvae were available.

Phylogeny. A modified subset of the morphology matrix consisting of 48 characters scored for 13 species of larvae, plus two outgroup species ( Linnaeus and (Fabricius)), was exported to TNT (Goloboff et al. 2008) and Winclada-NONA (Nixon 2002, Goloboff 1999) for phylogenetic analyses. Invariant characters and characters judged to be potentially highly variable between specimens (e.g. many characters involving color) were excluded from the analyses. Some character states were reordered and/or combined in the matrix used for analyses (Appendices 3 and 4) to reflect outgroup scoring and to clarify discrete parsimony-informative states. Characters and states from Appendices 3 and 4 are abbreviated in the text as (character:state).

Traditional searches were run with 10,000 random additions and TBR branch swapping. New technology searches were also performed using a variety of settings for the Sectorial Search, Rachet, Drift, and Tree fusing functions. Standard bootstrap (10,000 replicates) and Bremer support were assessed in TNT.

Results

The phylogenetic analyses returned one most parsimonious tree (Fig. 1). The genus PageBreak was relatively strongly supported (BS = 87, Bremer = 8). was placed at the base of the genus with the rest of the , excluding a reversal in + , having moderately punctate clypei (11:1). While the backbone of the clade had little support, several groupings were supported in the analyses.

Figure 1.

Most parsimonious tree (L = 141, CI = 0.5, RI = .53) based on larval morphology. Numbers not associated with rectangles are bootstrap support values (above branches) and Bremer support values (below branches). Smaller numbers above rectangles on branches represent character number; numbers below rectangles represent character state. Black rectangles correspond to non-homoplasious character state changes. White rectangles correspond to homoplasious character state changes. All character states were unambiguously optimized on the tree.

+ , representing the only members of the subgenus in the analyses, was well supported (BS = 99, Bremer = 10). The presence of four long setae on the ligula (18:4, Fig. 11A) and a trapezoidal hypopharyngeal sclerome (19:1; Fig. 12A) may represent synapomorphies for the subgenus.

Figure 11.

Ligulas of three species: A B C . Scale bars = 200 µm.

Figure 12.

Hypopharyngeal scleromes of two species: A B . Scale bars = 200 µm.

+ ( + ) was supported (BS = 60, Bremer = 3), and represents most of the members of the nominate subgenus in the analyses. The three species share two synapomorphies within the species sampled. One, the arrangement of anterior sensory papillae (16:1, Fig. 9B–C); and two, the presence of a distinct apical tooth on the pygidium (42:1, Fig. 14A). , another species currently in the nominate subgenus, is lacking both characters and was (BS = 64, Bremer = 4) supported in a sister relationship with from the subgenus . Both and adults are found on sand dunes, and the two larval synapomorphies the species share in the matrix (8:1 and 26:1) are based on the presence of dense setation, a common adaptation to living on sand. Hence, it is possible these character codings represent convergence based on larval habitat. also had one unusual autapomorphy in the presence of longitudinal tomentose bands of setae along the margins of the abdominal sternites (Fig. 13A), which may also be an adaptation for living primarily on unconsolidated dunes. was suggested as a member of the nominate subgenus (Triplehorn and Aalbu 1987), but also lacks the two synapomorphies found in other species of the subgenus. It was weakly supported in a relationship with + from the subgenus . Determining whether and belong in the subgenus requires further analyses of additional data.

Figure 9.

Epipharynges of four species: A B C D . Scale bars = 1 mm.

Figure 14.

Pygidia of two species: A B . Scale bar = 1 mm.

Figure 13.

Abdominal sternites I and II for two species: A B . Scale bars = 1 mm.

Larval descriptions

Larvae are described or redescribed to include differential characters to separate species within the genus. Verbatim locality label data are listed with “/” indicating line breaks on the label.

Eschscholtz, 1829

http://species-id.net/wiki/Eleodes

Material examined.

Over 1,400 larval specimens were examined for this study from 14 species. In addition, historical descriptions and specimens for which the species could not be confirmed due to a lack of positive association between adults and larvae also conform to the generic description provided.

Description.

Integument strongly sclerotized, light tan to nearly black in color; setose, with hair-like setae throughout and spinose setae on legs and abdominal tergite IX. Thoracic and abdominal segments subcylindrical, surface coriaceous (Figs 2A–D, 3A–D, 4A–C, 5A–C, 6A–C).

Figure 2.

Dorsal habitus of four species: A ; B C D . Scale bar = 5 mm.

Figure 3.

Dorsal habitus of four species: A B C D . Scale bar = 5 mm.

Figure 4.

Lateral habitus of three species: A B C . Scale bar = 5 mm.

Figure 5.

Lateral habitus of three species: A B C . Scale bar = 5 mm.

Figure 6.

Lateral habitus of three species: A B C . Scale bar = 5 mm.

Head. Prognathous or slightly declined (Fig. 7A–C), weakly dorsoventrally flattened, strongly constricted before occipital foramen. Epicranial stem one-fourth to one third head capsule length; frontal arms U-shaped or sinuate, occasionally obscured by sculpturing. Frons and dorsal portion of epicranial plates weakly to moderately rugose; punctate, punctures minute, lacking setae. Ventrolateral portions of epicranial plates setose; setae golden, erect; two stemmata present on each plate, pigmented spots often faded. Clypeus trapezoidal, often weakly transversely raised medially. Labrum with two transverse rows of six to fourteen erect setae present medially and subapically; anterior margin straight or weakly emarginate. Epipharynx (Figs 8, 9A–D, 10A–D) with stout spiniform setae along anterior margin, an anterior cluster of four to six variably arranged spinules, a subanterior transverse row of four small spinules subtended by two spinose setae and posterior cluster of six to eight small spinules; tormae symmetrical or asymmetrical. Mandible apex bidentate, mola concave. Ligula small, setation variable (Fig. 11A–C). Hypopharyngeal sclerome pentagonal or trapezoidal (Fig. 12A–B). Gula distinct, trapezoidal, widest in basal half. Antenna three segmented, cylindrical.

Figure 7.

Lateral habitus of the head and thoracic segments of three species: A B C . Scale bar = 5 mm.

Figure 8.

, epipharnyx. asp = anterior spines, msp = medial spines, mst = microsetae, pap = sensory papillae, tor = tormae. Scale bar = 1 mm.

Figure 10.

Epipharynges of four species: A B C D . Scale bars = 1 mm.

Thorax. Prothoracic tergum 1.2× or more length of meso- or metaterga (Figs 2A–D, 3A–D); anterior transverse striated band present, generally darker than protergal disc; lateral margins with granulated band either distinct or barely visible (Fig. 7A–C). Posterior transverse striated band present on all thoracic tergites. Meso- and metathoracic tergites wider than long. 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; dorsal surface of prothoracic femur with faintly indicated basal sclerotized band; dorsal surface of prothoracic tibia slightly more sclerotized than ventral surface.

Abdomen. Abdominal tergites and sternites I–VIII with transverse striated bands present along posterior margins. Abdominal sternite I setose (Fig. 13A–B). Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apical tooth present or absent (Fig. 14A–B); marginal row of socketed spines present around posterior two-thirds to one half of segment. Abdominal segment X located ventrally; pygopods short, subconical, each with erect setae.

Variation.

larvae can vary greatly in pigmentation, size, number of spines on the legs and pygidium, and the overall degree of sclerotization. Characters in the matrix relating to general integument coloration (6, 24, 45, 46, 47, 69) can vary greatly between specimens depending on age of specimen, length of time since last instar, and preservation method. There may also be genetic variation, though specimens from our populations were generally homogeneous.

Diagnosis.

All known larvae share the following combination of characters: head capsule weakly dorsoventrally flattened, strongly constricted before occipital foramen; prothoracic tergum 1.2× or more length of meso- or metaterga, anterior transverse striated band present, lateral margins with granulated band either distinct or barely visible; prothoracic leg slightly longer and much thicker than meso- and metathoracic legs; 8–38 socketed spines on the pygidial margin, pygopods short, subconical, each with erect setae. However, the known larvae cannot yet be separated from other Amphidorini larvae due to a lack of specimens.

Subgenus Horn, 1870

(LeConte, 1851)

http://species-id.net/wiki/Eleodes_nigropilosus

Fig. 4A

Material examined.

Larval 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.

Description.

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 PageBreakto 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.

Diagnosis.

larvae can be separated from the other currently known 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.

Subgenus Triplehorn, 1975

Eleodes Aalbu, Smith, & Triplehorn, 2012

http://species-id.net/wiki/Eleodes_wheeleri

Figs 2A , 4B

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, antePageBreakrolateral 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.

Remarks.

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

Subgenus Eschscholtz, 1829

LeConte, 1851

http://species-id.net/wiki/Eleodes_armatus

Figs 2B , 4C , 9B , 11B

Larval 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 granuPageBreaklated 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.

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

LeConte, 1858

http://species-id.net/wiki/Eleodes_caudiferus

Figs 2C , 5A , 13A

Larval 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, PageBreakmoderately 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, punctaPageBreaktion 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.

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

(Say, 1824)

http://species-id.net/wiki/Eleodes_hispilabris

Figs 9C , 14A

Larval 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 PageBreaksetose 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.

larvae can be separated from the other currently known 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.

Casey, 1890

http://species-id.net/wiki/Eleodes_tenuipes

Larval 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, PageBreakforming 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.

larvae can be separated from the other currently known 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 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.

Thomas, 2005

http://species-id.net/wiki/Eleodes_tribulus

Figs 2D , 5B , 9D

Larval 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, PageBreakwidest 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.

larvae can be separated from the other currently known 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.

Subgenus Blaisdell, 1909

(Say, 1823)

http://species-id.net/wiki/Eleodes_extricatus

Figs 3A , 5C , 7B , 10A , 12B , 13B

Larval PageBreak 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, PageBreakless 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.

larvae can be separated from the other currently known 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.

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.

Blaisdell, 1909

http://species-id.net/wiki/Eleodes_anthracinus

Figs 3B , 6A , 7A , 8

Larval 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 foPageBreakramen; 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 PageBreakPageBreakportions 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 PageBreakpresent 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.

larvae can be separated from most currently known 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 larvae by their lighter ventral segments and lack of distinct posterior pigmented bands on the abdominal terga. Larvae of (Say), another species in the subgenus , are mentioned as being “nearly black” by McColloch (1918). However, no other diagnostic characters are mentioned that would separate them from the other or .

Blaisdell

Figs 3C , 6B , 9A , 11A , 12A

Larval 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 PageBreakthan 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.

larvae can be separated from most currently known 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 larvae as outlined in that species diagnosis.

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 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.

Subgenus LeConte, 1862

Solier, 1848

http://species-id.net/wiki/Eleodes_goryi

Figs 10B , 11C

Larval 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 PageBreakseven 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 PageBreakgranulated 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.

larvae can be separated from the other currently known 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.

LeConte, 1851

http://species-id.net/wiki/Eleodes_subnitens

Figs 7C , 10C

Larval 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, PageBreakdarker 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.

larvae can be separated from the other currently known 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.

Horn, 1870

http://species-id.net/wiki/Eleodes_pilosus

Figs 3D , 6C , 10D , 14B

Larval 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, PageBreakinterspersed 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, PageBreaksparser 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.

larvae can be separated from the other currently known 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.

Notes on additional species

One specimen of (TB08942) was reared to a late instar, 9th or 10th, in the lab. However, the specimen apparently died in its rearing container while molting and suffered some damage, thus obscuring many characters. What could be seen of the epipharnyx, ligula, and abdominal segment IX, including the presence of an apical tooth on the pygidium, place it with , , and in the subgenus . The presence of spinose setae along the anterolateral margins of the epipharnyx placed it closest to , and .

Previously described species

Little comparative data to separate the species can be drawn from Gissler (1878). Blaisdell (1909) provides a more detailed description, but likewise does not include many characters currently necessary to differentiate the species.

Little comparative data to separate this species from the other currently described larvae can be drawn from Gissler (1878).

Little comparative data to separate this species from the other currently described larvae can be drawn from Hyslop (1912).

The thorough description in Wade and St. George (1923) easily places this species within the subgenus based on the epipharnyx and the apical tooth on the pygidium. It also appears to have spinose setae on the anterolateral margins of the epipharnyx, as in , , and .

McColloch (1918) describes the larvae as black in color after the first instar. The larvae of and , the only other larvae known, are similarly dark. The picture provided by McColloch (plate 5, image B), also looks similar to in gestalt.

Little comparative data to separate this species from the other currently described larvae can be drawn from Hyslop (1912).

Discussion

As adult morphology in many PageBreak species may be heavily influenced by participation in mimicry rings with co-occurring species (Doyen and Somerby 1974), the addition of characters from larval morphology may help produce a more accurate phylogeny based on morphological data than one using adult morphology alone. The presented phylogeny (Fig. 1) demonstrates the utility of larval morphology in resolving at least some relationships within the genus . The subgenus was well supported based on several synapomorphies present in the two included species. The subgenus showed two synapomorphies supporting a relationship for three of the included species, but did not recover a clade containing all of the current or presumed species from the nominate subgenus. Both and are somewhat unusual members of the subgenus based on adult morphology as well; hence further research is needed to accurately place them within the subgeneric classification. The inclusion of more taxa should increase phylogenetic accuracy and help illuminate the currently unresolved relationships between the subgenera (Heath et al. 2008). It is likely that extensive modifications to the matrix and key will be needed as more larvae become known. It is also possible that some species, particularly closely related ones, cannot be separated based on larval characters alone.

By producing matrix-based descriptions within mx, we are creating a growing repository of digital morphological and specimen data, already available through the tenebrionidbase.org portal, including an online multi-entry key (http://tenebrionidbase.org/public/clave) to the currently known larvae. Characters and states from the matrix and key will also be linked to the developing Coleoptera Anatomy Ontology project (ColAO).

1	Lateral margins of abdominal sternites I–IX pigmented, with golden tomentose setae (Fig. 13A)	Eleodes caudiferus LeConte
1’	Lateral margins of abdominal sternites I–IX unpigmented, lacking tomentose setae (Fig. 13B)	2
2 (1’)	Apex of pygidium attenuated and sclerotized, forming a distinct projection (Fig. 14A)	3
2’	Apex of pygidium somewhat attenuated, lightly or unsclerotized, not forming distinct projection (Fig. 14B)	5
3 (2)	Anterolateral margins of epipharynx with micro-setation (Figs 8, 9A–B, D, 10A–D), lacking stout setae	Eleodes armatus LeConte
3’	Anterolateral margins of epipharynx with stout setae (Fig. 9C)	4
4 (3)	Mesotarsus with row of three ventromedial spines; ventral portion epicranial plates with row of six or more long setae along anterior margin near buccal cavity, confluent with setae on lateral portions of plates; prothoracic femur with ventromedial row of 13–14 spines	Eleodes tenuipes Casey
4’	Mesotarsus with two ventromedial spines, ventral portion epicranial plates with row of four to five long setae along anterior margin, not confluent with setae on lateral portions of plates; prothoracic femur with ventromedial row of 6–10 spines	Eleodes hispilabris (Say)
5 (2’)	Granulated band along lateral margins of protergum faint, concolorous with protergal disc (Fig. 7A)	6
5’	Granulated band along lateral margins of protergum distinct, darker than protergal disc (Fig. 7B–C)	8
6 (5)	Hypopharyngeal sclerome pentagonal, tricuspidate (Fig. 12B); ligula apex lacking microsetae, two long subapical setae present ventrally, eight or more subapical setae present dorsally (Fig. 11B); pigmented band present along posterior margin of abdominal sterna, integument usually tan	Eleodes wheeleri Aalbu, Smith, & Triplehorn
6’	Hypopharyngeal sclerome trapezoidal (Fig. 12A); ligula apex glabrous, four long subapical setae present, two ventrally and two dorsally (Fig. 11A); pigmented band absent along posterior margin of abdominal sterna, integument usually dark brown	7
7 (6)	Terga dark brown, nearly black, throughout; prothoracic tergum wider than long, 1.2× or more length of meso- or metaterga; posterior pigmented band on abdominal terga I–VIII darker than rest of segment; abdominal sternites nearly concolorous with tergites	Eleodes carbonarius (Say)
7’	Terga medium brown, lighter towards margins; prothoracic tergum subquadrate, 1.5× length of meso- or metaterga; posterior pigmented band on abdominal terga I–VIII concolorous with rest of segment; abdominal sternites lighter than tergites	Eleodes anthracinus (Blaisdell)
8 (5’)	Pigmented band around posterior margin of abdominal segments dark along anterior edge, fading to segment color posteriorly (Fig. 4A)	Eleodes nigropilosus (LeConte)
8’	Pigmented band around posterior margin of abdominal segments unicolorous, darker than rest of segment throughout	9
9 (8’)	Abdominal sternite I moderately clothed in long erect setae to posterior pigmented band; seven anterior sensory papillae present on epipharnyx in all specimens examined	Eleodes pilosus (Horn)
9’	Abdominal sternite I with sparsely setose on at most anterior half; six anterior sensory papillae present on epipharnyx in all specimens examined	10
10 (9’)	Antennal segment I longer than antennal segment II (Fig. 7A–B); gula length less than maximum width	Eleodes extricatus (Say)
10’	Antennal segment I subequal to antennal segment II (Fig. 7C); gula length subequal or greater than maximum width	11
11 (10’)	Pygidium with marginal row of 8–14 socketed spines; prothoracic tergum subquadrate, 1.5× length of meso- or metaterga	Eleodes tribulus Thomas
11’	Pygidium with marginal row of 18–20 socketed spines; prothoracic tergum wider than long, 1.2× or more length of meso- or metaterga	12
12 (11’)	Eighth and ninth abdominal tergites more darkly pigmented than preceding segments, protibia with ventromedial row of 3–4 spines	Eleodes goryi Solier
12’	Eighth and ninth abdominal tergites with same pigmentation as preceding segments; protibia with ventromedial row of 5–6 spines	Eleodes subnitens LeConte

Characters with “{ }” indicate polymorphic codings

Eleodes_nigropilosus	01100501202110100000210101201100021111100110411211221010100015000100022120111021001000
Eleodes_tribulus	01110{04}112022201100012111011011{04}{03}0{12}1021100110411212221010100010000000022321111021401010
Eleodes_wheeleri	0110040130211001000121110120113102111211001044{01}212221100000016100200022511110211001010
Eleodes_armatus	01100211312110110001211102201102021012100000{24}212121111101000132003{03}0{02}1{12}311111111301010
Eleodes_caudiferus	010005012{01}22202100012{01}11044021000211011000102212120030101000121000{03}0022431111221610020
Eleodes_hispilabris	010002113122201000013111014020020{12}10221000004{24}12121111101000{12}3210200022121111111201000
Eleodes_tenuipes	0{01}1002113122201100013{12}11014020020210121000004212121211101000242103000221111111215010?0
Eleodes_extricatus	0110040130211011000{01}3011011011000211{01}1100000421212111010100010000200022121111011201010
Eleodes_anthracinus	01000{01}01200110100000210102201100020010000000{01}{01}01111110000000000000{12}1{01}1{01}120110010{014.0}01010
Eleodes_carbonarius_knausi	01000{23}01200110100000210101101100021120000000{23}{23}11221111000000111002{34}0{01}20121120010101000
Eleodes_goryi	01100{23}112001101100012111011011000210211001104{12}22{23}2111110100010000000022321001020101010
Eleodes_subnitens	01100211202110110001231101201100021021100110421232121110100010100200022221111021101020
Eleodes_pilosus	01000{45}0131122121000120110{12}{12}01111011031100000412232222110100012100100022211111021201020
Tenebrio_molitor	01100201302110020001311006202000021014120120222232111010101115000400022511001331721132
Zophobas_morio	01100{23}114122201000013211051012010211131210211122420010001011113102301215000010307211?0

Cladistic Character Codings

Eleodes_nigropilosus	012001100111102002011202200101100011201100211110
Eleodes_tribulus	002000001010002012011202200101100013211100211110
Eleodes_wheeleri	013001101010102003010202201011110010111112111110
Eleodes_armatus	003101101010102113010101101101120213111101111110
Eleodes_caudiferus	112100011010212002010200010101110014311102211210
Eleodes_hispilabris	103100000000211113010101101101121011211101111111
Eleodes_tenuipes	00310000100021111301010120110112101111110121111?
Eleodes_extricatus	013001101000002002010101100101100011211100111110
Eleodes_anthracinus	112011100111102014110111100011100101201110101110
Eleodes_carbonarius_knausi	112011100111002004110111101011110101211210101110
Eleodes_goryi	002011101010002012011201101101100013210000201110
Eleodes_subnitens	002001101010102012011201201101110012211100211111
Eleodes_pilosus	113110011010[01]02012010102211101110012111100211111
Tenebrio_molitor	013001102000111011001001100100000010110003010000
Zophobas_morio	000100000000000000000000000000031100000000000000