More miscellaneous improvements, such as better/more specific phrasing, and more inclusion for locations and specimens. More info in V1.0: https://www.inaturalist.org/journal/eleodesthermopolis/87075-v1-0-a-key-to-the-eleodes-species-of-the-united-states-and-canada-outdated
1 Mesofemora and metafemora dentate or subdentate...2
1’ Mesofemora and metafemora never dentate...4
2 Elytral punctures strongly muricate...E. moesta
2’ Elytral punctures simple or slightly muricate...3
3 Pronotum widest anterior of middle, femoral spines large...E. armata
3’ Pronotum widest at middle, femoral spines small to medium sized...E. femorata
4 Body small, pronotum generally cordate and constricted at base, elytra apex not attenuate or caudate, roughly sculptured, never sulcate, sometimes with short setae, usually distributed across beaches, forests, and mountains, rarely in arid regions...subgenus Blapylis
4’ Body small or large, pronotum usually not cordate although often slightly constricted at base, elytral apex attenuate, caudate or neither, roughly sculptured or not, sulcate or not, with setae or not, widespread across all environments...5
5 Legs long with unproportionately small body, robust or not, body often flattened if not robust, never sulcate, if legs not abnormally long, then body elongate and flattened...6
5’ Legs long or short but with body fairly proportionate, only rarely flattened, sulcate or not, if legs fairly long, then not with conspicuous setae, or a small, elongate, and flattened body...22
6 Body generally slender and flattened, pronotal and elytral sculpturing simple (except in E. nigrina), never with conspicuous setae...7
6’ Body, generally robust, never flattened, pronotal and elytral sculpturing complex, usually with conspicuous setae...18
7 Elytral sculpturing muricate or scabrous, southern New Mexico to Canada, Sierra Nevada Mountains to Rocky Mountains...E. nigrina (in part)
7’ Elytral sculpturing simple, always below the northernmost edge of Colorado’s border...8
8 Range only in Inyo White Mountains...9
8’ Range widespread...10
9 Profemora with blunt tooth, anterolateral angles of pronotum blunt but projected, legs shorter...E. aalbui
9’ Profemora only sinuate, anterolateral angles of pronotum slightly projected, legs longer...E. microps
10 Range near Big Bend National Park and the Guadalupe Mountains...11
10’ Range in the Southwest United States excluding Texas and the Guadalupe Mountains...13
11 Range in the Guadalupe Mountains...E. guadalupensis
11’ Range near Big Bend National Park...12
12 Mentum with finger-like process, pronotum lacking projected anterolateral angles, widest at middle, near Rio Grande river...E. labialis
12’ Mentum without finger-like process, pronotum with slightly projected anterolateral angles, Big Bend National Park...E. easterlai
13 Range in the Mohave Desert of California, pronotum usually evenly arcuate...E. californica
13’ Range in Arizona, Utah, and Sonora, pronotum rarely evenly arcuate...14
14 Range in the Sonoran desert or the most western portion of the Arizona-Utah border...15
14’ Range in the Sky Islands or the Colorado Plateau...16
15 Anterolateral angles strongly projected, caves by the most western point of the Arizona-Utah border…E. wynnei
15’ Anterolateral angles not strongly projected, the Sonoran desert…E. delicata (E. nevadensis)
16 Body elongate, northern Arizona and southern Utah...17
16’ Body not elongate, Sky Islands in southeast Arizona...E. dissimilis
17 Strial punctures larger than punctures of intervals, restricted to the Grand Canyon...E. leptoscelis
17’ Strial punctures subequal to punctures of intervals, northern Arizona and southern Utah...E. wheeleri
18 Large tubercles present on Elytra...19
18’ Elytra without large tubercules...20
19 Range in Sierra Nevada region...E. granosa
19’ Range near the Four Corners...E. leechi
20 Pronotum evenly tuberculate, range in Oregon...E. spoliata
20’ Pronotum punctate, range widespread...21
21 Antennas robust, body generally more robust...E. hirsuta
21’ Antennas thin, body generally more slender...E. pilosa
22 Size small or sometimes medium, covered in long setae, restricted to coastal and sometimes inland California, and the Baja peninsula, or rarely in the direction of and in Reno, or the Sonoran desert...23
22’ Size small or large, usually glabrous, if with conspicuous setae then not found in coastal California, nor in the Baja peninsula, or the Sonoran desert, and if found near Reno, Nevada, then third antennomere not shortened...28
23 Males with dentate profemora and elytra caudate, Sonoran desert...E. tribulus
23’ Males nor females ever caudate or attenuate, coastal California and the Baja peninsula...24
24 Third antennomere shortened, body smaller and more slender, pill shaped, coastal California and towards and into Reno, Nevada...E. littoralis
24’ Third antennomere not shorted, body larger, California...25
25 Body larger, pronotum with constriction at base, if missing constriction then setae more dense...26
25’ Body smaller, pronotum lacking constriction at base, setae less dense...27
26 Setae orange, California and northern Baja peninsula...E. osculans
26’ Setae black or grey, California and northern Baja peninsula...E. ursus
27 Setae black, California and northern Baja peninsula...E. nigropilosa
27’ Setae yellow, Baja peninsula...E. subdeplanata
28 Elytra caudate at least in males and strongly muricate, near Tamaulipas if not caudate, size usually large...29
28’ Elytra caudate or not, but if caudate and muricately punctured then elytra only slightly muricate, elytra can be strongly muricate if not caudate and not near Tamaulipas, size large or small...31
29 Only males caudate with profemora dentate, Tamaulipas...E. fiski
29’ Both males and females caudate with mutic profemora, Four Corners and Sierra Nevada region...30
30 Elytra usually glabrous, rarely with setae, Four Corners...E. caudifera
30’ Elytra always with long setae, Sierra Nevada region...E. longipilosa
31 Elytral sculpture very rough, elytra always attenuate but never caudate, pronotum wide, restricted to California, or slightly into Nevada and Oregon...32
31’ Elytral sculpture variable but if rough then elytra never attenuate, widespread...33
32 Luster shiny and elytral apex acutely attenuate...E. marginata
32’ Luster dull and elytral apex blunt yet still attenuate...E. scabricula (E. acutangula)
33 Body oval shaped, coated in minute setae, coastal Texas and Louisiana...E. veterator
33’ Body shape variable, if small and shaped slightly oval then not found in coastal Texas or Louisiana...34
34 Body small and fusiform...35
34’ Body small or large, shape variable...37
35 Body nearly oval, clothed with minute setae, giving body a grey appearance, Great Plains of northern Texas to Canada...E. opaca
35’ Elytra highly punctured and glabrous...36
36 Body more fusiform, central and southern Great Plains...E. fusiformis
36’ Body less fusiform, the Sierra Nevadas to the Rocky Mountains, and the Arizona Sky Islands to British Colombia, northern Great Plains...E. extricata (in part)
37 Size medium to large, elytra usually inflated, wide, and large or sometimes flattened, especially in E. striolata, punctures either large, or appearing absent, although if elytra are not inflated, and attenuate, then punctures may be numerous and conspicuous, never sulcate...38
37’ Size variable but abdomen rarely inflated, if so, then punctures are small but conspicuous, sulcate or not...41
38 Elytral striae composed of large conspicuous punctures...E. goryi
38’ Elytral striae without large punctures...39
39 Elytra usually flattened, always attenuate, and heavily punctate, luster shiny, south and central Texas...E. striolata
39’ Elytra sometimes flattened, inconspicuous punctures present, luster usually dull, Arizona, New Mexico, and Sonora...40
40 Pronotum more or less parallel sided, anterolateral angles almost always projected outwards...E. subnitens
40’ Pronotum evenly arcuate, anterolateral angles only rarely projected outwards...E. madrensis
41 Size large sometimes medium or small in rare occasions, but almost always with legs long, sometimes caudate...42
41’ Size usually small, sometimes medium, if so then legs almost always short, never caudate...71
42 Body elongate, sometimes robust, pronotum not all that distinct from elytra, profemora never dentate and never with caudate elytra, punctures simple or muricate/scabrous, never sulcate...43
42’ Body fairly variable, sometimes elongate, often robust, pronotum usually quite distinct from elytra, males almost always with dentate profemora, elytra sometimes caudate, sulcate or not...48
43 Elytral punctures simple, anterolateral angles never very projected, body rarely flattened, but if so, then usually near the Sky Islands of southeast Arizona…45
43’ Elytral punctures not simple, anterolateral angles of pronotum projected, or body somewhat flattened and small, never in the Sky Islands…44
44 Anterolateral angles strongly projected, elytral punctures with tiny inconspicuous setae arising from each one, size usually not small...E. inornata
44’ Anterolateral angles not projected or only slightly projected, elytral punctures glabrous, size smaller, sometimes quite small…E. nigrina (continuation)
45 Elytra with red line running down the length of the elytra, this line never diffuses into black...E. ornatipennis
45’ Elytra entirely black, or if a red line is present along the elytra, it diffuses into black...46
46 Profemora strongly sinuate, pronotum widest slightly anterior of middle...E. hepburni
46’ Profemora simple, pronotum widest at middle...47
47 Prosternum horizontal, not projecting downwards, restricted to California...E. gigantea
47’ Prosternum projecting downwards, widespread but not found in California...E. longicollis
48 Elytra never completely smooth (muricate, irregularly punctured, sulcate even if sulci not deep), pronotum sometimes concave, body large, robust, and elytra with carina, if carina absent and body not very robust, then anterolateral angles of the pronotum rounded off, and the pronotum widest anterior of middle...49
48’ Elytra variable, often smooth, pronotum never concave, body variable, and elytra lacking carina, if elytra sulcate, or irregularly punctured, then pronotum widest at middle or with acute anterolateral angles...54
49 Pronotum concave...E. suturalis
49’ Pronotum convex...50
50 Elytra with carina...E. acuta
50’ Elytra without carina...51 (E. obscura)
51 Elytra sulcate...52
51’ Elytra not sulcate...53
52 Elytra deeply sulcate, west of the Great Plains...E. obscura sulcipennis
52’ Elytra slightly sulcate, Great Plains...E. obscura obscura
53 Elytra with simple punctures, western Texas...E. obscura glabrisulca
53’ Elytra with muricate punctures, Four Corners...E. obscura dispersa
54 Elytra with large glabrous costae, intervals minutely hirsute, often coated with white debris, males are caudate, females are sometimes slightly caudate, legs long...E. mirabilis
54’ Elytra variable but without costae, caudate or not...55
55 Pronotum very wide, humeral angles of elytra obsolete, never attenuate...E. grandicollis
55 Pronotum variable, if wide, then elytral apex attenuate at least, humeral angles of elytra not obsolete...56
56 Restricted to California and the Baja peninsula, never having deep sulci...57
56’ Widespread not found in California or the northern Baja peninsula, if found in California then elytra sulcate...63
57 Pronotum two times wider than the head...E. acuticauda
57’ Pronotum less then two times wider than the head...58
58 Pronotum more or less parallel sided, Mohave desert...59
58’ Pronotum not parallel sided, or if so then elytral apex with groove in between both elytra, coastal and inland California…60
59 Tibia strongly curved, profemoral spines blunt, never caudate...E. subcylindrica
59’ Tibia not strongly curved, profemoral spines acute, males caudate...E. mexicana
60 Elytral punctures minute, pronotum nearly parallel sided, although sometimes slightly arcuate, elytral apex with groove in between both elytra…E. discincta
60’ Elytral punctures not minute, or if so then pronotum wide…61
61 Pronotum larger, elytral striae less prominent…E. dentipes
61’ Pronotum smaller, elytral striae more prominent…62
62 Pronotum widest anterior of middle, luster more shiny, elytra with simple punctures, glabrous, striae very clear…E. gracilis distans
62’ Pronotum widest at middle, luster more dull, elytra with muricate punctures and conspicuous setae, striae not clearly defined…E. adumbrata
63 Elytra strongly sulcate, at least on elytral apex…64
63’ Elytra not strongly sulcate…65
64 Pronotum widest anterior to middle…E. hispilabris
64’ Pronotum widest at middle…E. composita
65 Body flattened, elytra and pronotum not too distinctly separated, or if so then elytra remain wide up until their humeral angles…E. knullorum
65’ Body usually not flat, elytra and pronotum clearly separate, if elytra remain wide up until their humeral angles, then elytra inflated…66
66 Elytra with muricate punctures…67
66’ Elytra with simple punctures…68
67 Punctures of elytra arranged in rows and fairly muricate, elytral apex always caudate, even if short, El Paso region…E. tenuipes
67’ Punctures of elytra irregular and strongly muricate, elytral apex only sometimes attenuate, generally around the Four Corners and western Texas…E. sponsa
68 Hind angles of pronotum appearing rounded, size large, elytra often inflated and strongly convex, or fairly parallel sided…69 (E. spinipes)
68’ Hind angles of pronotum not rounded or only slightly rounded, size medium to large, elytra never inflated, but often parallel sided…70
69 Elytra more or less parallel sided, west of the Pecos River…E. spinipes macrura
69’ Elytra ventricose, east of the Pecos River…E. spinipes ventricosa
70 Anterolateral angles of pronotum always facing straight forward, never projecting outwards, male metatibia sinuate, males caudate, females strongly attenuate, strial punctures inconspicuous…E. eschscholtzii eschscholtzii
70’ Anterolateral angles of pronotum always projecting outwards at least a little, metatibia not abnormally sinuate, elytral apex only slightly attenuate, striae composed of rows of conspicuous punctures…E. gracilis gracilis
71 Elytra with conspicuous setae…72
71’ Elytra glabrous, or only with short inconspicuous setae…76
72 Size less than 12mm, elytra shiny with complex sculpturing, and covered in golden hairs, areas with sandy soils generally around the Four Corners…E. barbata
72’ Size usually small but rarely less than 12mm, elytra usually dull, sometimes shiny, never with golden hairs, usually in deserts above the Sonoran desert…73
73 Size larger, body more robust, northern Arizona and southern Utah…E. halli
73’ Size smaller, body more slender, generally around the Great Basin and the Great Sand Dunes of Colorado…74
74 Distributed only in the Great Sand Dunes National Park…E. hirtipennis
74’ Distributed generally around the Great Basin…75
75 Setae black, generally around the great basin…E. letcheri
75’ Setae brownish, in and around Utah…E. aspera
76 Elytra flat and widened, about the same width as pronotum, sides sometimes slightly rounded or with carina…77
76’ Sides of elytra rounded evenly, generally elongate or circular…81
77 Body covered with minute setae, usually appearing grey in color…E. tricostata
77’ Body glabrous…78
78 Elytral sculpturing rough, striae vague to absent, luster generally dull…E. neomexicana
78’ Elytra with clear striae, luster shiny…79
79 Elytra with deeply impressed striae…E. pedinoides
79’ Elytra with slightly impressed striae…80 (E. wenzeli)
80 Punctures of elytra fine but easily noticeable…E. wenzeli speculicollis
80’ Punctures of elytra extremely fine and minute…E. wenzeli wenzeli
81 Both elytra with at least three large smooth costae, legs often dark red, southern Utah…E. parowana
81’ Both elytra without at least three large costae…82
82 Size small, elytral luster generally shiny, elytra only with conspicuous irregular punctures, never with tubercules…E. extricata (continuation)
82’ Size small to medium, luster variable although generally dull if elytra with irregular punctures, said irregular punctures also smaller, elytra with punctures in rows or irregular, or with tubercules or papillae…83
83 Elytra generally inflated, if not, then elytra more or less parallel sided, usually tuberculate, Pacific Northwest…E. humeralis
83’ Elytra inflated or not, but if with tubercules then elytra not inflated and not parallel sided, if without tubercules then not found in the Pacific Northwest, or if in the Pacific Northwest then pronotum not clearly widest in front of middle…84
84 Elytra with papillae, pronotum wide…E. papillosa
84’ Elytra without papillae…85
85 Elytra with large tubercules…86
85’ Elytra without large tubercules…87
86 Pronotum more or less arcuate and wider than long…E. granulata
86’ Pronotum more or less parallel sided and longer than wide…E. subtuberculata
87 Size small, pronotum evenly arcuate, lateral edges angular instead of rounded, punctures minute…E. arcuata
87’ Size variable, pronotum variable, but if arcuate, then lateral edges rounded…88
88 Luster shiny, elytral punctures large and in rows, outer protibial spur clearly larger than inner protibial spur…E. debilis
88’ Luster variable but usually not too shiny, punctures variable but rarely large, if protibial spurs disproportionate in size, then elytra with small punctures or tubercules…89
89 Longer protibial spur extending to almost to the third tarsomere…90 (E. rileyi)
89’ Longer protibial spur extending only to apex of the first tarsomere…91
90 Elytra shiny with prominent striae…E. rileyi reducta
90’ Elytra dull without very prominent striae…E. rileyi rileyi
91 “…longer protibial spur evenly tapering to a point from base to apex”…E. carbonaria (see Thomas 2012 for subspecies key)
91’ “…longer protibial spur parallel sided in basal half then tapering to a point at apex”…92
92 Elytral punctures irregular, San Francisco region…E. quadricollis
92’ Elytral punctures organized in rows, southeast Arizona…93 (E. anthracina)
93 Size larger, elytral punctures less fine…E. anthracina anthracina
93’ Size smaller, elytral punctures more fine…E. anthracina lustrans
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