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

Abstract 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 Eleodes 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), Eleodes 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 Eleodes, 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 Eleodes extricatus (Say, 1824), Eleodes hispilabris (Say, 1824), Eleodes obsoletus (Say, 1824), Eleodes opacus (Say, 1824), and Eleodes suturalis (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 Eleodes species, only seven have been previously described in the larval stages.

Taxonomic history
Relatively few Eleodes larvae have been described or characterized (Table 1). Gissler (1878) provided the first larval descriptions in the genus for Eleodes dentipes Eschscholtz, 1833 and Eleodes giganteus (Mannerheim, 1843). Hyslop (1912) described the larvae of Eleodes vandykei Blaisdell, 1909 (then listed as a subspecies of Eleodes letcheri) and Eleodes pimelioides Mannerheim, 1843 from the Pacific Northwest. McColloch (1918) described Eleodes tricostatus (Say), 1824. Wade and St. George (1923) described Eleodes suturalis (Say, 1824), followed closely by illustrations, without additional descriptions, of the pygidia of Eleodes carbonarius (Say, 1824), Eleodes opacus (Say, 1824), and E. tricostatus by St. George (1924). Blaisdell (1909) redescribed the larvae of E. dentipes in greater detail and described the pupa of Eleodes clavicornis Eschscholtz, 1833. The most recent larval description was provided by Thomas (1984) for Eleodes armatus LeConte, 1851. In most cases, these descriptions are insufficient to reliably diagnose Eleodes larvae to species.  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 Eleodes, 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. Total length (TL) was measured from the anterior edge of the clypeus to the dorsomedial apex of abdominal segment IX. Prothoracic width (PW) and length (PL) were measured dorsally across the widest and longest points on the segment respectively, head capsule width (HW) 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.
Rearing. Adult Eleodes 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 Eleodes 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 Eleodes caudiferus, 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 Eleodes larvae, plus two outgroup species (Tenebrio molitor Linnaeus and Zophobas morio (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 parsimonyinformative 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 Eleodes was relatively strongly supported (BS=87, Bremer=8). Eleodes extricatus was placed at the base of the genus with the rest of the Eleodes, excluding a reversal in E.
hispilabris + E. tenuipes, having moderately punctate clypei (11:1). While the backbone of the clade had little support, several groupings were supported in the analyses.
Eleodes armatus + (E. tenuipes + E. hispilabris) was supported (BS=60, Bremer=3), and represents most of the members of the nominate subgenus Eleodes 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). Eleodes caudiferus, another species currently in the nominate subgenus, is lacking both characters and was (BS=64, Bremer=4) supported in a sister relationship with Eleodes pilosus from the subgenus Tricheleodes. Both E. caudiferus and E. pilosus 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. Eleodes caudiferus also had one unusual autapomorphy in the presence of longitudinal tomen- 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. tose bands of setae along the margins of the abdominal sternites (Fig. 13A), which may also be an adaptation for living primarily on unconsolidated dunes. Eleodes tribulus 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 E. goryi + E. subnitens from the subgenus Promus. Determining whether E. caudiferus and E. tribulus belong in the subgenus Eleodes requires further analyses of additional data.

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.
Eleodes Eschscholtz, 1829 http://species-id.net/wiki/Eleodes Material examined. Over 1,400 larval Eleodes specimens were examined for this study from 14 Eleodes species. In addition, historical descriptions and Eleodes 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.
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. Eleodes 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 Eleodes 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 Eleodes larvae cannot yet be separated from other Amphidorini larvae due to a lack of specimens.

Subgenus Blapylis Horn, 1870
Eleodes (Blapylis) nigropilosus (LeConte, 1851) http://species-id.net/wiki/Eleodes_nigropilosus Figure 4A Material examined. Larval E. nigropilosus specimens were reared from adults with the following collecting information: "USA: CA: San Diego Co. / Oceanside beach / 33.1865, -117.3778 / 14.May.2011, ADSmith". A total of 29 eggs and larvae were reared and examined for this study, of which 34 survived to the 2nd instar or beyond. The following description is based on a detailed examination of three 8-11th instar specimens.  Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color light to dark tan, same or nearly the same as body segments; punctation minute, moderately dense, separated by 2-4 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons faintly rugose. Epicranial plates weakly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of four to five long setae along anterior margin near buccal cavity, not confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, not swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2-4 puncture diameters. Labrum not swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae subapical setal row with seven to eight erect setae, anterior margin straight to weakly emarginate. Epipharynx anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular diagonal rows; four subanterior sensory papillae present, arranged as transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetric, left torma smaller. Ligula apex and subapical dorsal surface densely micro-setose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length subequal or greater than maximum width. Antenna three segmented, cylindrical, length of first segment subequal to second.
Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs medium brown, thoracic sternites posterior to prolegs light brown. Prothoracic tergum subquadrate, 1.5× length of meso-or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, forming a gradient from darker brown anteriorly to lighter brown along posterior border. Meso-and metathoracic tergites wider than long, each with a faintly indicated sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso-and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout spines ventromedially; prothoracic femur with ventromedial row of three spines, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of three to four spines, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with two ventromedial spines. Abdomen. Abdominal tergites and sternites light tan with darker transverse striated bands present along posterior margins of segments I-VIII, forming near contiguous band around segments, bands dark along anterior edge, fading to segment color posteriorly. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex not forming a distinct tooth, moderately clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 14-18 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I-VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 9-12 erect setae.
Diagnosis. Eleodes nigropilosus larvae can be separated from the other currently known Eleodes species by having the posterior pigmented band around the abdominal segments forming a color gradient from dark along anterior edge and fading to the color of the rest of the segment posteriorly. 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, ante-rolateral 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 mesoand 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.
Diagnosis. 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. 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 granu-lated 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.
Diagnosis. Eleodes armatus larvae can be separated from the other currently known Eleodes species by presence of an apical tooth on the pygidium and the absence of stout spiniform setae on the anterolateral margins of the epipharnyx. Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width narrower than prothorax; sides rounded; strongly constricted before occipital foramen; color dark tan, same or nearly the same as on body segments; punctation minute, moderately dense, separated by 2-4 puncture diameters. Epicranial suture stem length approximately one-fourth to one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons rugose. Epicranial plates rugose dorsally; lateral portions densely setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, puncta- tion 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.

Eleodes (Eleodes) caudiferus
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.
Diagnosis. Eleodes caudiferus larvae can be separated from the other currently known Eleodes species by the presence of longitudinal tomentose bands along the lateral margins of abdominal sternites I-VIII. Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width narrower than prothorax; sides rounded; strongly constricted before occipital foramen; color ferruginous, more heavily pigmented than body segments; punctation minute, dense, separated by 1-2 puncture diameters. Epicranial suture stem length approximately one-fourth head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons rugose. Epicranial plates rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of four to five long setae along anterior margin near buccal cavity, not confluent with setae on lateral portions of plates, with a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation dense, separated by 1-2 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with 10-14 erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 9C) anterior setal row with eight or more stout spiniform setae, anterolateral margins with stout spinose setae; six anterior sensory papillae present, arranged in two irregular rows, each with two posterior papillae and one near the anterior margin; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; seven to eight posterior sensory papillae present, arranged in an irregular cluster. Tormae strongly asymmetric, left torma larger. Ligula apex lacking microsetae, two long subapical setae present ventrally, eight or more subapical setae present dorsally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length less than maximum width. Antenna three segmented, cylindrical, first segment longer than second.
Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs light tan to ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum wider than long, 1.2× or more length of meso-or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso-and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso-and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with one or two stout ventromedially spines; prothoracic femur with ventromedial row of six to ten spines, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of eight to eleven spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with four to five ventromedial spines.
Abdomen. Abdominal tergites and sternites light tan, with slightly darker transverse striated bands present along posterior margins of segments I-VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex forming a distinct tooth, sparsely clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 17-23 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I-VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 9-12 erect setae.
Diagnosis. Eleodes hispilabris larvae can be separated from the other currently known Eleodes species by the presence of an apical tooth on the pygidium, stout spiniform setae on the anterolateral margins of the epipharnyx, and a row of 6-10 ventromedial spines on the prothoracic femur. 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 onefourth head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons rugose. Epicranial plates rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation dense, separated by 1-2 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with 10-14 erect setae, anterior margin straight to weakly emarginate. Epipharynx anterior setal row with eight or more stout spiniform setae, anterolateral margins with stout spinose setae; six anterior sensory papillae present, arranged in two irregular rows, each with two posterior papillae and one near the anterior margin; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae strongly asymmetric, left torma smaller. Ligula apex lacking microsetae, two long subapical setae present ventrally, eight or more subapical setae present dorsally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length less than maximum width. Antenna three segmented, cylindrical, first segment longer than second.
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.
Diagnosis. Eleodes tenuipes larvae can be separated from the other currently known Eleodes species by the presence of an apical tooth on the pygidium, stout spiniform se-tae on the anterolateral margins of the epipharnyx, and a row of 13-14 ventromedial spines on the prothoracic femur. It is further differentiated from E. hispilabris by having a row of three ventromedial spines on the mesotarsus and having the ventral portion of the epicranial plates with a row of six or more long setae along anterior margin near buccal cavity, confluent with setae on lateral portions of plates.
Remarks. 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. Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides angular; strongly constricted before occipital foramen; color light tan to medium brown, more heavily pigmented than body segments ; punctation minute, moderately dense, separated by 2-4 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons rugose. Epicranial plates weakly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2-4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae subapical setal row with six to seven erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 9D) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; five to six anterior sensory papillae present, arranged in two irregular longitudinal rows or an irregular cluster; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; seven to eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetric, left torma larger. Ligula apex and subapical dorsal surface densely micro-setose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length subequal or greater than maximum width. Antenna three segmented, cylindrical, first segment length subequal to second.
Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs medium brown, thoracic sternites posterior to prolegs light brown. Prothoracic tergum subquadrate, 1.5× length of meso-or metaterga; lateral margins with distinct granulated band, darker than protergal disc; anterior transverse striated band present, darker than tergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso-and metathoracic tergites wider than long, each with a faintly indicated sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso-and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout spines ventromedially; prothoracic femur with ventromedial row of two spines and three to five longer setae, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of three to four spines, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with three ventromedial spines.
Abdomen. Abdominal tergites and sternites light tan with darker transverse striated bands present along posterior margins of segments I-VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I moderately clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex not forming a distinct tooth, moderately clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 8-14 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I-VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 11-15 erect setae.
Diagnosis. Eleodes tribulus larvae can be separated from the other currently known Eleodes species based on the pentagonal hypopharyngeal sclerome, lack of a caudal tooth on the pygidium, presence of 8-14 marginal spines on the pygidium, and the angular, nearly straight sides of the head capsule. Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color light tan, same or nearly the same as body segments; punctation minute, dense, separated by 1-2 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons faintly rugose. Epicranial plates faintly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen or not, unicolorous, minutely punctate, punctation dense, separated by 1-2 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with six to seven erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 10A) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae symmetrical or weakly asymmetrical with left torma smaller. Ligula apex densely microsetose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length less than maximum width. Antenna three segmented, cylindrical, first segment longer than second.
Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum subquadrate, 1.5× length of meso-or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso-and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso-and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout ventromedially spines; prothoracic femur with ventromedial row of two spines and three to five longer setae, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of three to four spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with four to five ventromedial spines.
Abdomen. Abdominal tergites and sternites light tan, with slightly darker transverse striated bands present along posterior margins of segments I-VIII, forming near contiguous unicolorous band around segments. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex lacking a distinct tooth, sparsely clothed in short and mid length erect setae, sclerotized uniformly throughout, lacking maculations; marginal row of 17-23 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I-VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 11-15 erect setae.
Diagnosis. Eleodes extricatus larvae can be separated from the other currently known Eleodes species based on the pentagonal hypopharyngeal sclerome, small or absent apical tooth on the pygidium, lateral margins of prothoracic tergum with a distinct granulated band, and having antennal segment I longer than antennal segment II.
Remarks. Eleodes extricatus is a widespread species found on dunes and at high elevations. Specimens from Arizona and Texas showed no population differences in the larval stage. Adults varied in the presence or prominence of muricate tubercles on the elytra. Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital fo-ramen; color medium brown to brown-grey, nearly as on body segments; minute punctation moderately dense dorsally. Epicranial stem approximately one-third head capsule length; frontal arms U-shaped, not obscured by sculpturing. Frons and dorsal portion of epicranial plates faintly rugose; lacking non-primary setae.  Lateral portions of epicranial plates moderately setose; setae golden, erect, length equal to or longer than antennal segment 2; ventral portions of epicranial plates with a row of four long setae along anterior margin near buccal cavity with a patch of short setae medially forming a triangular pattern with its base near the anterior margin; two stemmata present on each plate, pigmented spots often faded. Clypeus trapezoidal; not swollen, moderately punctate, darker medially in basal half. Labrum not swollen, basal half more darkly pigmented; sides rounded; two transverse rows of seven to eight erect setae present medially and subapically; anterior margin straight. Epipharynx (Fig. 3) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular diagonal rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in two irregular rows.Tormae asymmetrical, left torma smaller. Ligula with four long setae near apex. Hypopharyngeal sclerome trapezoidal. Gula distinct, trapezoidal, widest in basal half. Antenna three segmented, cylindrical; first segment longer than second.
Thorax. Grey-brown to medium brown dorsally and anterior to legs on prothoracic sternite, tan on rest of sternites; lighter transverse striated band present along anterior fourth of prothoracic tergum; thin darkly sclerotized transverse line present on anterior fifth of meso-and metathoracic tergites; striated bands present along posterior 5th of all thoracic tergites, color forming a gradient from darker brown anteriorly to lighter brown along posterior border. Eight evenly arranged setae present on dorsal surface of each thoracic terga, lateral margins more densely setose. Prothoracic tergum subquadrate, 1.5× length of meso-or metaterga; lateral margins lacking pigmented band. Meso-and metaterga wider than long, lacking pigmented bands along lateral margins; mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracle; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Prothoracic leg slightly longer, much thicker than meso-and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; trochanter with row of two stout spines and two longer setae ventromedially, tibia with ventromedial row of two spines and four to five longer setae, tarsus with ventromedial row of four spines. Dorsal surface of protibia (at rest) with faintly indicated basal sclerotized band; dorsal surface of protarsus slightly more sclerotized than ventral surface.
Abdomen. Tergites grey-brown to medium brown dorsally, lightening towards lateral margins, sternites light to dark tan; transverse striated bands not visible on abdominal sternites, barely visible on posterior 5th of terga I-VIII, nearly concolorous with rest of tergites. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline, abdominal segments II-VIII each with two sparse transverse bands of long erect setae, posterior margin of segment 8 denser setal band. Abdominal laterotergites concolorous with tergites, lacking distinct pigmented margins. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, sparsely clothed in short and mid length erect setae, dorsally more sclerotized in apical two-thirds with faint maculations; marginal row of 14-18 socketed spines present apical half, apex not forming distinct sclerotized projection. Pygopods short, subconical, each with 11-15 erect spines.
Variation. Little variation was observed between specimens beyond the number of spines on the legs and pygidium, and the overall degree of sclerotization.
Diagnosis. Eleodes anthracinus larvae can be separated from most currently known Eleodes species based on their darker dorsal coloration on all segments, the absence of pigmented bands along the lateral margins of the thoracic terga, and the lack of a distinct sclerotized tooth at the apex of the pygidium. They can be distinguished from E. carbonarius larvae by their lighter ventral segments and lack of distinct posterior pigmented bands on the abdominal terga. Larvae of Eleodes tricostatus (Say), another species in the subgenus Melaneleodes, are mentioned as being "nearly black" by Mc-Colloch (1918). However, no other diagnostic characters are mentioned that would separate them from the other E. anthracinus or E. carbonarius.  Figures 3C, 6B, 9A, 11A, 12A Material examined. Larval E. carbonarius specimens were reared from adults with the following collecting information: "USA: CO: Montezuma Co. / Ute RA off Hwy 160 / 37.3535, -108.44385 / 05 Jun 2011, ADSmith". A total of 129 eggs and larvae were reared and examined for this study, with 45 surviving until the 2nd instar or later. The following description is based on a detailed examination of five 8-11th instar specimens.
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 nonprimary setae. Lateral portions of epicranial plates moderately setose; setae golden, erect, length equal to or longer than antennal segment 2; ventral portions of epicranial plates with a row of four to five long setae along anterior margin near buccal cavity and a patch of short setae medially forming a triangular pattern with its base near the anterior margin; two stemmata present on each plate, pigmented spots often faded. Clypeus trapezoidal; not swollen, moderately punctate, darker medially in basal half. Labrum not swollen, basal half more darkly pigmented; sides rounded; two transverse rows of six to seven erect setae present medially and subapically; anterior margin straight to weakly emarginate. Epipharynx (Fig. 9A) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular diagonal rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetrical, left torma larger. Hypopharyngeal sclerome trapezoidal. Ligula with four long setae near apex. Gula distinct, trapezoidal, widest in basal half. Antenna three segmented, cylindrical; first segment longer than second.
Thorax. Dark brown to ferruginous dorsally and anterior to legs on prothoracic sternite, lighter brown on rest of sternites; distinct longitudinally striated band present along anterior fourth of prothoracic tergum; thin darkly sclerotized transverse line present on anterior fifth of meso-and metathoracic tergites; striated bands present along posterior 6th of all thoracic tergites, darker than rest of surface. Eight evenly arranged setae present on dorsal surface of each thoracic terga, lateral margins more densely setose. Prothoracic tergum wider than long, 1.5× length of meso-or metaterga; lateral margins lacking pigmented band. Meso-and metaterga wider than long, lacking pigmented bands along lateral margins; mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracle; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Prothoracic leg slightly longer, much thicker than meso-and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; trochanter with two stout spines ventromedially, tibia with ventromedial row of three to four spines and four to five longer setae, tarsus with ventromedial row of five spines. Dorsal surface of protibia (at rest) with basal sclerotized band; dorsal surface of protarsus more sclerotized than ventral surface.
Abdomen. Tergites dark brown to ferruginous, concolorous or lightly lighter than tergites; longitudinally striated bands not visible on abdominal sternites, distinct on posterior 5th of terga 1-8. Abdominal sternite I sparsely clothed in long erect setae from anterior margin to near midline, abdominal segments 2-8 each with two sparse transverse bands of long erect setae, posterior margin of segment 8 denser setal band. Abdominal laterotergites concolorous with tergites, lacking distinct pigmented margins. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, sparsely clothed in short and mid length erect setae, apical two-thirds with faint maculations; marginal row of 18-20 socketed spines present apical half, apex not forming distinct sclerotized projection. Pygopods short, subconical, each with 9-12 erect spines. Variation. Little variation was observed between specimens beyond the number of spines on the legs and pygidium, and the overall degree of sclerotization.
Diagnosis. Eleodes carbonarius larvae can be separated from most currently known Eleodes species their darker dorsal coloration on all segments, the absence of pigmented bands along the lateral margins of the thoracic terga, and the lack of a distinct sclerotized tooth at the apex of the pygidium. They can be further distinguished from E. anthracinus larvae as outlined in that species diagnosis.
Remarks. Eleodes carbonarius adult morphology is notoriously variable across the species range and even within populations. Nine subspecies are currently recognized (Triplehorn and Thomas 2011). The specimens examined were all reared from a few females of E. carbonarius knausi Blaisdell collected at a single locality. Larval characters showed little variation; however, this may change as more specimens are reared from other localities and subspecies. Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color ferruginous to dark brown, more heavily pigmented than body segments; punctation minute, moderately dense, separated by 2-4 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms U-shaped, not obscured by sculpturing. Frons faintly rugose. Epicranial plates faintly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates, and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker medially in basal half, minutely punctate, punctation moderately dense, separated by 2-4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with six to seven erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig.  10B) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae strongly asymetrical with left torma larger. Ligula apex densely microsetose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length subequal or greater than maximum width. Antenna three segmented, cylindrical, first segment subequal to second.

Subgenus
Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs ferruginous to medium brown, thoracic sternites posterior to prolegs medium brown. Prothoracic tergum wider than long, 1.2× or more length of meso-or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, unicolorous brown. Meso-and metathoracic tergites wider than long, each with a heavily sclerotized transverse line present on anterior fifth. Thoracic tergites sparsely setose on dorsal surfaces, lateral margins more densely setose. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracles; reduced metathoracic spiracle visible, less than one-fourth size of mesothoracic spiracle. Legs. Prothoracic leg slightly longer, much thicker than meso-and metathoracic legs; prothoracic tarsungulus strongly sclerotized, sickle-shaped; prothoracic trochanter with two stout ventromedially spines; prothoracic femur with ventromedial row of two spines and three to five longer setae, dorsal surface with faintly indicated basal sclerotized band; prothoracic tibia with ventromedial row of three to four spines or spinose setae, dorsal surface slightly more sclerotized than ventral surface. Mesotibia with three ventromedial spines.
Abdomen. Abdominal tergites and sternites 1-7 light tan, with slightly darker transverse striated bands present along posterior margins of segments I-VIII, forming near contiguous unicolorous band around segments. Abdominal tergite 8 more darkly pigmented than preceding segments. Abdominal sternite I moderately clothed in long erect setae from anterior margin to near midline. Abdominal laterotergites with lateral margins distinctly pigmented. Abdominal segment IX (pygidium) triangular in dorsal view, gradually reflexed to apex, urogomphi absent, apex lacking a distinct tooth, moderately clothed in short and mid length erect setae, dorsally more sclerotized in apical two-thirds with faint maculations; marginal row of 18-20 socketed spines present, arranged as single row around posterior two-thirds to one half of segment. Abdominal sternites I-VIII lacking longitudinal tomentose bands along lateral margins. Pygopods short, subconical, each with 11-15 erect setae.
Diagnosis. Eleodes goryi larvae can be separated from the other currently known Eleodes species based on the darkly pigmented eighth and ninth abdominal tergites. It is further distinguished by the pentagonal hypopharyngeal sclerome, lack of a caudal tooth on the pygidium, and the presence of 3-4 ventromedial spines on the protibia.
Description Head. Prognathous or weakly declined; weakly dorsoventrally flattened; width nearly equal to prothorax; sides rounded; strongly constricted before occipital foramen; color ferruginous, more heavily pigmented than body segments; punctation minute, moderately dense, separated by 2-4 puncture diameters. Epicranial suture stem length approximately one-third head capsule length; frontal arms sinuate, not obscured by sculpturing. Frons faintly rugose. Epicranial plates faintly rugose dorsally; lateral portions moderately setose; ventral portion of each plate with row of six or more long setae along anterior margin near buccal cavity confluent with setae on lateral portions of plates and a patch of short setae medially, forming a triangular pattern with its base near the anterior margin. Two stemmata present on each epicranial plate, pigmented spots often faded. Clypeus trapezoidal, swollen, darker in apical half, minutely punctate, punctation moderately dense, separated by 2-4 puncture diameters. Labrum swollen, sides rounded, basal half more darkly pigmented, medial setal row with six to seven erect setae, subapical setal row with seven to eight erect setae, anterior margin straight to weakly emarginate. Epipharynx (Fig. 10C) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; six anterior sensory papillae present, arranged in two irregular rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; eight posterior sensory papillae present, arranged in an irregular cluster. Tormae asymetrical with left torma smaller. Ligula apex densely microsetose, two long subapical setae present ventrally. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half, length subequal or greater than maximum width. Antenna three segmented, cylindrical, first segment subequal to second.
Thorax. Thoracic tergites light tan, prothoracic sternite anterior to legs ferruginous, thoracic sternites posterior to prolegs light brown. Prothoracic tergum wider than long, 1.2× or more length of meso-or metaterga; anterior transverse striated band present, darker than protergal disc; lateral margins with distinct granulated band, darker than protergal disc. Posterior transverse striated band present on all thoracic tergites, uni- colorous 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.
Diagnosis. Eleodes subnitens larvae can be separated from the other currently known Eleodes species by the pentagonal hypopharyngeal sclerome, prothoracic tergum wider than long, 8th and 9th abdominal tergites not darker than proceeding segments, lack of a caudal tooth on the pygidium, and the presence of 5-6 ventromedial spines on the protibia. Head. Prognathous, weakly flattened, narrower than prothorax; sides rounded, strongly constricted before occipital foramen; color nearly as in body segments. Epicranial stem short, one-fourth head capsule length; frontal arms U-shaped, partially obscured by sculpturing. Frons and dorsal portion of epicranial plates rugose; sparsely setose; densely punctate, punctures minute, lacking setae. Ventrolateral portions of epicranial plates densely setose; setae golden, erect, most longer than antennal segment 2, interspersed with shorter setae; two stemmata present on each plate, pigmented spots often faded. Clypeus trapezoidal; swollen, weakly transversely raised medially; moderately punctate, rugose in basal half. Labrum slightly swollen, basal fourth darkly pigmented; sides rounded, minutely tomentose; two transverse rows of six to eight erect setae present medially and subapically; anterior margin straight. Epipharynx (Fig. 10D) anterior setal row with six stout spiniform setae, anterolateral margins with micro-setation; seven anterior sensory papillae present, arranged in two irregular longitudinal rows; four subanterior sensory papillae present, arranged as a transverse row subtended by two spinose setae; seven posterior sensory papillae present, arranged in an irregular cluster. Tormae asymmetrical, left side torma smaller with or without a small spine near emergent edge. Hypopharyngeal sclerome pentagonal, tricuspidate. Gula distinct, trapezoidal, widest in basal half. Antenna three segmented, cylindrical; first segment longer than second.
Thorax. Light to dark tan, darker longitudinally striated bands present on anterior margin of prothoracic tergum and posterior 5th of all thoracic tergites. Sparsely setose along dorsal margins of terga near striated bands, lateral margins more densely setose. Prothoracic tergum subquadrate, 1.5× length of meso-or metaterga; lateral margins with pigmented band along entire length. Mesothoracic spiracle simple, ovate, approximately 1.5× size of abdominal spiracle; reduced metathoracic spiracle visible, less than onefourth size of mesothoracic spiracle. Prothoracic leg slightly longer, much thicker than meso-and metathoracic legs; tarsungulus strongly sclerotized, sickle-shaped; trochanter with two stout spines ventromedially, tibia and tarsus each with a ventromedial row of four to seven spines, number of spines often differing between prolegs. Dorsal surface of protibia (at rest) with basal sclerotized band; dorsal surface of protarsus sclerotized.
Abdomen. Light to dark tan, darker longitudinally striated bands present on posterior 5th of segments I-VIII. Abdominal sternite I moderately clothed in long erect setae, sparser medially, extending to posterior pigmented band, abdominal tergite I and segments II-VIII sparsely clothed in short to mid length setae. Lateral margins of abdominal laterotergites I-VIII darkly pigmented, ventral margin with two pigmented bands. Abdominal segment IX (pygidium) triangular in dorsal view, moderately clothed in long erect setae; marginal row of 10-20 socketed spines present apical half, apex not forming distinct sclerotized projection. Pygopods short, subconical, each with 16-20 erect spines.

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

Discussion
As adult morphology in many Eleodes 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 Eleodes. The subgenus Melaneleodes was well supported based on several synapomorphies present in the two included species. The subgenus Eleodes 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 E. tribulus and E. caudiferus 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 Eleodes 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 tenebrio-nidbase.org portal, including an online multi-entry key (http://tenebrionidbase.org/public/ clave) to the currently known Eleodes larvae. Characters and states from the matrix and key will also be linked to the developing Coleoptera Anatomy Ontology project (ColAO).