taxonID	type	description	language	source
470A7D7EFF7CFFE8FE99F96817D96DBC.taxon	description	(FIGS 1 A, 8 A, B, 10 A, 11 A, 12 A, 13 B, 14 A)	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF7CFFF4FF10F93B17AF681F.taxon	description	Nomenclatural notes: This is the species referred to as ‘ Bembidion “ Ebbets Pass ” ’ in Sproul & Maddison (2017) and B. breve (specimen 1930) in Maddison (2012). Diagnosis: A shiny, medium-sized, parallel-sided, relatively flat species with strongly foveate dorsal punctures (Fig. 1 A). Forebody and elytra black or dark brown often with aeneous hue. Pronotum widest anterior to middle with obtuse hind angles; not sinuate laterally; laterobasal carina strong (Figs 11 A, 12 A). Elytra parallel-sided; dorsal punctures strongly foveate. Elytral disc commonly with uneven surface (caused by a depression in elytral surface) in basal third anterior to dorsal punctures (see pattern of shadows in the basal third of elytral disc in Fig. 1 A). Microsculpture weakened in males (Fig. 13 B) causing shiny appearance and making foveate dorsal punctures easily visible without magnification. Male genitalia with flagellum sinuate and moderately long; sclerite ‘ St’ slender, commonly hourglass-shaped (Figs 8 A, B, 10 A). Comparison with similar species: Most easily confused with B. ampliatum and B. breve. Can be distinguished from the former by having a slightly more convex pronotum and stronger laterobasal carina, more strongly foveate dorsal punctures (particularly where they cooccur with B. ampliatum in California) and unevenness in the basal third of the elytral disc anterior to the dorsal punctures, and a longer, more sinuate flagellum and more slender sclerite ‘ St’. Distinguished from the latter by having more parallel-sided elytra, the pronotum widest anterior to middle and a slen- der sclerite ‘ St’. May also be confused with B. laxatum from which it is distinguished by having a smaller body size, weaker microsculpture, a narrower pronotum relative to elytral width and sclerite ‘ St’ lacking U-shaped recurvature. Geographic distribution: From southern British Columbia along the Cascade Range to the southern Sierra Nevada. East through Idaho to the Rocky Mountains in Montana and southeastern British Columbia (Fig. 14 A). Habitat: Known from a wider range of environments than most other species in the group. It is often abundant in the damp soil below receding snow patches on alpine slopes. Also present along the shoreline of streams and lakes at high elevation. Common along the shorelines of moderate-sized rivers at somewhat lower elevation with increasing latitude (e. g. the Pacific Northwest and Alaska). Geographic variation: This species is fairly variable across its range and within populations. Notable size variation is common in multiple locations of the Oregon Cascades and in Montana, with very small females in some populations (those sequenced do not show obvious differences in the genes examined).	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF60FFF3FE94FC60166B693F.taxon	description	(FIGS 1 B, 8 C, D, 10 B, 11 B, 12 D, 15 A)	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF60FFF3FE94FC60166B693F.taxon	diagnosis	Diagnosis: This medium-sized species is parallel-sided with a distinctly flat pronotum. Forebody and hindbody black or very dark brown, often with aeneous or metallic hue. Pronotum relatively flat; hind angles obtuse; widest anterior to middle with broad lateral explanation and weak laterobasal carina (as the basal fovea beside it is not as deep, and thus the carina does not stand out so prominently); not sinuate laterally (Figs 11 B, 12 C). Elytra are more or less parallel-sided and broadly rounded at apex; elytral disc with smooth appearance (particularly in California and Oregon), in part due to weak striae and weakly foveate dorsal punctures (Fig. 1 B) which are not easily observed without magnification. Microsculpture meshes moderately etched in females, but weakly or at least unevenly etched in most males, often partially disappeared (13 D). Male genitalia with the ventral portion of sclerite ‘ St’ angled anteriorly such that the ventral extremity is even with, or anterior to, the dorsal extremity; flagellum short and somewhat sinuate (Figs 8 C, 13 D, 10 B). Comparison with similar species: Most easily confused with B. lividulum from which it can be distinguished by having a flatter pronotum with weak laterobasal carina (especially reliable in California), less pronounced elytral striae with weakly foveate dorsal punctures and a smooth elytral disc in the basal third anterior to the dorsal punctures (though often less so in specimens east of California and Oregon), and the male genitalia having a more laterally expanded sclerite ‘ St’ with the ventral portion angled anteriorly. May also be confused with B. laxatum from which it can be separated through the weakened elytral microsculpture in both sexes, by having a flatter pronotum with obtuse hind angles and weakened laterobasal carina, and lacking the U-shaped recurvature in sclerite ‘ St’ of the male genitalia. Geographic distribution: Throughout the Sierra Nevada and White Mountains in California and north throughout the eastern mountains of Oregon (Steens and Wallowas) to Alberta. Easterly through the Great Basin and Rocky Mountains to Montana, Wyoming, and Colorado, south to New Mexico and Arizona. The only breve group species known from Utah, Wyoming, Colorado, New Mexico and Arizona (Fig. 15 A). A single questionable specimen is reported from Washington (see note below about ‘ Geographic variation’). Habitat: Open alpine slopes, commonly below patches of melting snow. Geographic variation: Specimens east of California and Oregon (e. g. Great Basin and Rocky Mountains) are generally smaller-bodied, especially males, and show more intraspecific variation in external structures. Notable within-population size variation has been observed in the Manti La Sal Mountains of eastern Utah, and the Sangre de Cristo Mountains of New Mexico. The single specimen from Washington reported herein (DNA 3321) is a morphological and molecular outlier that we doubtfully include within this species.	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF67FFF3FE8DFD6A10156DE6.taxon	description	(FIGS 1 C, 8 E, F, 10 C, 11 C, 13 A, 14 B)	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF67FFF3FE8DFD6A10156DE6.taxon	diagnosis	Diagnosis: A small-bodied, convex species with strongly foveate dorsal punctures. Forebody dark brown, hindbody dark brown or reddish brown, often lighter than forebody; forebody and hindbody commonly with a metallic hue. Pronotum with fairly rounded lateral margin, widest at middle or just anterior to middle; laterobasal carina strong but often short (not proceeding far anteriorly) due to convexity of pronotum; basal fovea deep; hind angles slightly obtuse (Fig. 11 C). Elytral striae generally pronounced and dorsal punctures strongly foveate; elytra fairly short relative to length of pronotum (Fig. 1 C). Microsculpture with meshes moderately etched in both sexes (Fig. 13 A) (but note geographic variation below). Male genitalia with medium-length flagellum; sclerite ‘ St’ more or less diamond-shaped (Figs 8 E, F, 10 C). Comparison with similar species: Most easily confused with B. lividulum, B. saturatum and B. vulcanix. Distinguished from B. lividulum by having more convex pronotum and elytra with somewhat rounded lateral margin of elytra, pronotum widest closer to middle, and male genitalia with a longer, more sinuate flagellum and broader sclerite ‘ St’. Distinguished from B. saturatum and B. vulcanix by having a slightly smaller body size, more rounded lateral margin of elytra that narrows at the shoulder and more strongly foveate dorsal punctures, with pronotum larger relative to elytra and with a short laterobasal carina, and male genitalia with a shorter flagellum. Geographic distribution: The northernmost species, ranging from the Aleutian Islands south along the coastal mountains of British Columbia, and in the Cascades of Oregon and Washington, south throughout California in the Sierra Nevada (Fig. 14 B). Also known from one locality in Yoho National Park in eastern British Columbia. Habitat: Most common on small, subalpine creeks in the southern part of its range. In the north, it occurs along creeks, rivers or open slopes at high elevation. Geographic variation: Some specimens are larger-bodied in the Sierra Nevada with broader pronota relative to northern localities. Notable intraspecific variation (in body size, pronotum shape, forebody and elytral coloration) is present in northern populations (e. g. southeast Alaska). Microsculpture shape and intensity is variable; in particular, some specimens may have notably transverse meshes (e. g. Queen Charlotte Islands, British Columbia), or less deeply etched meshes such that portions of cells are partially disappeared in males (e. g. Snoqualmie Pass, Washington). Some individuals from Yoho National Park are notably small in size.	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF67FFF2FC4DF92211166AC2.taxon	description	(FIGS 1 D, 8 G, H, 10 H, 11 H, 12 B, 13 E, F, 15 B)	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF67FFF2FC4DF92211166AC2.taxon	diagnosis	Diagnosis: Large and heavy-bodied, this parallel-sided species is distinct in its dull appearance, particularly females. Forebody and hindbody dark brown, rarely black, typically with aeneous, or bluish, metallic hue. Pronotum broad and somewhat convex; widest near middle with hind angles near 90 ° or slightly obtuse (Fig. 11 H); laterobasal carina strong (Fig. 12 B). Elytra widest near middle, often narrowing towards apex; elytral striae somewhat pronounced, and dorsal punctures moderately foveate (Fig. 1 D). Microsculpture strongly etched in males; very strongly etched, often granulate, in females (Fig. 13 E, F). Male genitalia with short, weakly sinuate flagellum; sclerite ‘ St’ with U-shaped recurvature ventrally (Figs 8 G, H, 10 H). Comparison with similar species: Most easily confused with B. oromaia from which it is distinguished by having a wider pronotum relative to width of the elytra, the elytra being widest near middle, having larger protarsomeres in males, less slender, elongate legs and antennae, and by various characters in the male genitalia. May also be confused with B. lividulum and B. ampliatum. It can be distinguished from both by its larger, more convex body, duller appearance due to strong microsculpture in both sexes, broadened pronotum relative to the elytra with hind angles near 90 °, and the U-shaped recurvature of sclerite ‘ St’ in the male genitalia. Geographic distribution: Throughout the Sierra Nevada in California, north along the Cascade Range to Washington and southern British Columbia (Fig. 15 B). Habitat: Open alpine slopes commonly below patches of melting snow or along small alpine creeks. Geographic variation: Sclerite ‘ St’ in males from California localities is fairly elongate (Fig. 9 G, H). Sclerite ‘ St’ in the few Washington and British Columbia specimens available for examination appears slightly less elongate and more broadened than in California specimens.	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF66FFF1FC2AFDEB17996869.taxon	description	(FIGS 2 A, 9 A, B, 10 D, 11 D, 16 A)	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF66FFF1FC2AFDEB17996869.taxon	diagnosis	Diagnosis: This small-bodied convex species has a broad pronotum and stout appearance (Fig. 2 A). Forebody and hindbody dark brown, some specimens with an aeneous reflection. Pronotum very broad basally with hind angles near 90 °; lateral explanation broad (Fig. 11 D), laterobasal carina somewhat weak as the basal fovea beside it is quite shallow, and thus the carina does not stand out so prominently, although carina may still extend far anteriorly. Elytra long relative to length of pronotum; tapered apically in some specimens; dorsal punctures weakly foveate (Fig. 2 A). Microsculpture meshes strongly etched in females and moderately etched in males (except for populations in eastern Oregon and Nevada where microsculpture is notably weakened). Male genitalia with long sinuate flagellum (Figs 9 A, B, 10 D); sclerite ‘ St’ either lacking (Fig. 9 A), or with a hint of sclerotization in the position of sclerite ‘ St’ (Fig. 9 B). Comparison with similar species: Most easily confused with B. vulcanix and B. geopearlis, from which it is extremely difficult to separate using only external structures. It is most easily distinguished using characters in the male genitalia and geography. It is separated from both by lacking sclerite ‘ St’ (but see note on ‘ Geographic variation’ below), and by having a southern and southeastern distribution (Fig. 16). May also be confused with B. breve, from which it is distinguished by a slightly larger body with more parallel-sided elytra, weakly foveate dorsal punctures, and a longer, more sinuate flagellum of the male genitalia. Geographic distribution: From Washington south to the southern Sierra Nevada, east to the Ruby Mountains in Nevada and Steens Mountains in Oregon (Fig. 16 A). Habitat: A variety of damp environments at or below the tree line including damp meadows, along lakeshores or small creeks, or on open alpine slopes below patches of melting snow. Geographic variation: Populations from the Steens Mountains in eastern Oregon and the Ruby Mountains in Nevada (the ‘ eastern form’) are smaller-bodied, shinier and more black than dark brown, with slight sclerotization in the position of sclerite ‘ St’ evident (e. g. Fig. 9 B), but not nearly so expanded as in B. geopearlis (Figs 9 C, D, 10 E) or B. vulcanix (Figs 9 E, F, 10 F). A slight patch of sclerotization has also been observed occasionally in individuals from California (Fig. 9 B).	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF65FFF0FE91FCBB17D06A4A.taxon	description	(FIGS 2 B, 9 C, D, 10 E, 11 E, 16 B) Holotype male (in OSAC) here designated, labelled: ‘ USA: Montana: Glacier N. P., east slope Clements Mtn., 2129 m, 48.692 ° N 113.7292 ° W, 12 August 2015. JSS 2015.107 - 1 [‘ - 1 ’ handwritten]. J. S. Sproul & family’ [white paper], ‘ David R. Maddison DNA 4727 DNA Voucher’ [pale green paper], ‘ HOLOTYPE Bembidion geopearlis Sproul + Maddison 2017 ’ [partly handwritten, red paper], ‘ Oregon State Arthropod Collection OSAC _ 0002000000 [matrix code] ’ [printed on both sides of white paper]. Genitalia mounted in Euparal on small card labelled ‘ DNA 4727 ’ beneath the specimen; extracted DNA stored separately. GenBank accession numbers for DNA sequences of the holotype are: KY 950786 (28 S); KY 950914 (CAD); KY 951044 (COI), KY 951174 (MSP), KY 951301 (Topo). Type locality: USA: Montana: Glacier National Park, east slope Clements Mountain near Logan Pass, 2129 m, 48.69204 ° N 113.72920 ° W. Paratypes: Thirty-eight paratypes from the following localities, with specimens deposited in OSAC and USNM: USA: Montana: Glacier National Park, east slope Clements Mountain near Logan Pass, 2129 m, 48.69204 ° N 113.72920 ° W (25); USA: Montana: Glacier Co., Glacier National Park, Iceberg Lake (2); USA: Montana: Glacier Co., Glacier National Park, Logan Pass (1); USA: Montana: Glacier National Park (1); USA: Montana: Mineral Co., Hoodoo Creek, 1780 m (1); USA: Montana: Missoula Co., inlet to Heart Lake, 1891 m, 47.3801 ° N 113.8501 ° W (1); USA: Montana: Ravalli Co., Lost Horse Creek, 1760 m, 46.1417 ° N 114.4863 ° W (1); USA: Montana: Ravalli Co., Lost Horse Creek, 1660 m, 46.1402 ° N 114.4371 ° W (1); USA: Montana: Flathead Co., Glacier National Park, Sperry Chalets (1); Canada: British Columbia: Akamina Pass, 1740 m, 49.0261 ° N 114.0611 ° W (3); Canada: Alberta: Waterton Lakes National Park, Cameron Lake, 5440 m (1). Derivation of specific epithet: The name ‘ geopearlis’ is derived from an informal combination of letters taken from the names of JSS’s children ‘ George’ (geo) and ‘ Pearl’ (pearl) with an ending (is) to make the name euphonic. The name recognizes the contribution of George (age 10) and Pearl (age 7) to the present work. They have accompanied JSS on over 6500 miles of high-intensity collecting road trips, hiked over 40 miles to high-elevation habitats in California, Oregon and Montana, and helped collect hundreds of specimens including members of the type series of this species. JSS is indebted to them for their companionship and support. The components of the name also reference the organism: ‘ geo’ evokes something of the earth; thereby, ‘ earth pearl’ or ‘ a precious thing from the earth’ is descriptive of these seldom-collected ground beetles. Diagnosis: A small, convex species with a notably broad prothorax (Fig. 2 B). Forebody dark brown or black; hind body dark brown or reddish brown. Pronotum variable but generally very broad basally with a broad lateral explanation; hind angles near 90 °; sinuate laterally in some specimens (Fig. 11 E). Elytra widest behind middle; often but not always broadly rounded at apex; dorsal punctures weakly foveate. Microsculpture meshes strongly etched in females and moderately etched in males. Genitalia somewhat parallel-sided in outer shape having a short taper towards the apex; flagellum sinuate and moderately long; sclerite ‘ St’ expanded (Figs 9 C, D, 10 E). Comparison with similar species: Most similar in appearance to B. saturatum, B. vulcanix and B. breve. Most reliably distinguished from all three with male genitalic characters and by its eastern geographic distribution. In particular, distinguished from B. saturatum and B. vulcanix by having elytra widest behind middle resulting in a more rounded apex of the elytra, and by the apical half of the aedeagus being less curved and elongate. Distinguished in male genitalia from B. saturatum by the presence of an expanded sclerite ‘ St’. Distinguished from B. vulcanix by having a less expanded, non-arcuate sclerite ‘ St’. Distinguished from B. breve by having a broader pronotum, more weakly foveate elytral dorsal punctures, the shape and position of sclerite ‘ St’. Geographic distribution: Known only from the Rocky Mountains of Montana, the Waterton National Parks area of southern Alberta and British Columbia, and the Wallowa and Blue Mountains in northeastern Oregon (Fig. 16 B). Habitat: Collected in abundance on open slopes above the tree line at the type locality. Small series or singletons have also been collected on small creeks or depressions with damp soil below the timberline.	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF64FFF0FE8DFE7210FD6E50.taxon	description	(FIGS 2 C, 9 E, F, 10 F, 11 F, 16 A) Holotype male (in OSAC) here designated, labelled: ‘ USA: Oregon: Deschutes Co., Stream east of Todd Lake, 1952 m, 44.0282 ° N 121.6709 ° W, 24. v. 2015. JSS. 2015.030 - 1 [‘ - 1 ’ handwritten]. J. S., E. C., G. S., & P. E. Sproul’ [white paper], ‘ David R. Maddison DNA 4615 DNA Voucher’ [pale green paper], ‘ HOLOTYPE Bembidion vulcanix Sproul + Maddison 2017 ’ [partly handwritten, red paper], ‘ Oregon State Arthropod Collection OSAC _ 0002000001 [matrix code] ’ [printed on both sides of white paper]. Genitalia mounted in Euparal on small card labelled ‘ DNA 4615 ’ beneath the specimen; extracted DNA stored separately. GenBank accession numbers for DNA sequences of the holotype are: KY 950767 (28 S), KY 950895 (CAD), KY 951025 (COI), KY 951155 (MSP), KY 951282 (Topo). Type locality: USA: Oregon: Deschutes Co., Deschutes National Forest, stream east of Todd Lake, 1952 m, 44.0282 ° N 121.6709 ° W. Paratypes: Seventy-two paratypes from the following localities, with specimens deposited in BMNH, CAS, CSCA, EMEC, MNHN and USNM: USA: Oregon: Deschutes Co., stream east of Todd Lake, 1952 m, 44.0282 ° N 121.6709 ° W (57); USA: Oregon: Deschutes Co., Creek below Little Three Creek Lake, 2018 m, 44.1057 ° N 121.6347 ° W (6); USA: Oregon: Deschutes Co., NE Todd Lake, Deschutes NF road 370, 2067 m, 44.038 ° N 121.6718 ° W (1); USA: Oregon: Deschutes Co., E Todd Lake, Deschutes NF road 370, 1976 m, 44.0306 ° N 121.6683 ° W (1); USA: Oregon: Hood River Co., Mt. Hood, Hood River Meadow Ski Area, 5300 ft. (2); USA: Washington: Whatcom Co., Bagley Lakes, Mt Baker, Snoqualmie NF, 1326 m, 48.8528 ° N 121.6886 ° W (1); USA: Washington: Whatcom Co., Bagley Lakes, Mt Baker, Snoqualmie NF, 1290 m, 48.8534 ° N 121.6948 ° W (1); USA: Washington: Pierce Co., Mt. Rainier, Tipsoo Lake (1); USA: Washington: Pierce Co., Mt. Rainier, Yakima Park (1); Canada: British Columbia: Garibaldi Provincial Park, S. slope Black Tusk (1). Derivation of specific epithet: Informally derived by combining the two Latin words Vulcanis, the blacksmith god of fire and volcanoes from Roman mythology, and nix, meaning snow. The name references snow-covered volcanoes of the Cascade Range where this species can be commonly found at high elevation. Diagnosis: This convex species is recognized by a broad pronotum basally, elongate elytra and distinctive male genitalia. Forebody and hindbody dark brown in some specimens aeneous. Pronotum very broad basally; hind angles near 90 °; lateral explanation broad; laterobasal carina fairly weak due to shallow adjacent basal fovea, but may extend far anteriorly nearly parallel to lateral margin of pronotum (Fig. 11 F). Elytra long relative to length of pronotum and somewhat bullet-shaped in that they are parallel-sided, and tapering toward apex (although not readily obvious in Fig. 2 C); dorsal punctures weakly foveate; striae often pronounced (Fig. 2 C). Microsculpture strongly etched in females and moderately etched in males. Aedeagus is strongly curved, with an elongate taper towards the narrow apex; sclerite ‘ St’ in apical half, large and arcuate; flagellum long and sinuate (Figs 9 E, F, 10 F). Comparison with similar species: Most easily confused with B. saturatum and B. geopearlis. It is separated from both by having an expanded, arcuate sclerite ‘ St’ in the apical half (which is absent in the former, and not as expanded, arcuate, or apically positioned in the latter), and by its northwestern distribution (Fig. 16). May also be confused with B. breve from which it is distinguished by its generally larger body size, more parallel-sided elytra, weakly foveate dorsal punctures, and longer, more sinuate shape of the flagellum and large of sclerite ‘ St’ in the male genitalia. Habitat: Along the shorelines of subalpine small creeks or lakes. Geographic distribution: From southwestern British Columbia and the Olympic Peninsula in Washington, south along the Cascades of Washington and Oregon to the Trinity Alps in northwestern California (Fig. 16 A).	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF64FFFFFC32FA7317276D79.taxon	description	(FIGS 2 D, 9 G, H, 10 G, 11 G, 17 A)	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF64FFFFFC32FA7317276D79.taxon	diagnosis	Diagnosis: This fairly large-bodied species is most easily recognized by the inflated appearance of the elytra and narrow pronotum relative to width of the elytra (Fig. 2 D). Forebody dark brown or black; hind body from dark brown to reddish brown, often paler than forebody. Pronotum narrow relative to the width of the elytra; strongly sinuate laterally (Figs 2 D, 11 G); hind angles near 90 ° with basal fovea deeply excavated (Fig. 11 G). Elytra notably convex with strongly rounded lateral margin resulting in an inflated appearance; dorsal punctures weakly foveate, elytral striae weak with striae three and four partially disappeared in some specimens. Microsculpture strongly etched in females and moderately etched in males. Male genitalia with darkened patch of membranes apically; sclerite ‘ St’ lacking; flagellum sinuate and moderately long; ostial flag with an abbreviated sinuation not nearing the ventral surface of the aedeagus, and not extending far anteriorly (Figs 9 G, H, 10 G). Comparison with similar species: Can be confused with B. saturatum and B. breve in the northern Sierra Nevada. Distinguished from both by a larger body size, more convex elytra, and a darkened patch of scale-like structures apically in the male genitalia (see inset in Fig. 9 H). Further distinguished from the former by having a narrower pronotum at the base. Distinguished from the latter by having very weakly foveate dorsal punctures and weak striae of the elytra. Geographic distribution: Known from the Sierra Nevada and Trinity Alps in California, as well as a single locality in southern Oregon (Fig. 17 A). Habitat: Appears to be restricted to small, subalpine creeks. Geographic variation: Some specimens from the Trinity Alps in northwestern California have a slightly longer flagellum in male genitalia. The single individual we sampled from Mount Ashland, Oregon has four distinctive bases in 28 S (three of which are ambiguities in DNA 4173, but non-ambiguous and different in all other specimens), but is not notably distinct in the other genes. The Oregon specimen also has less sinuate later margins of the pronotum than typical California specimens.	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
470A7D7EFF6BFFFEFEA2F9D6140E6FEB.taxon	description	(FIGS 3, 9 I, 10 I, 11 I, 13 C, 17 B) Holotype male (in OSAC) herein designated, labelled ‘ USA: California: Tulare Co., snow field above Emerald Lake, 2851 m, 36.5959 ° N 118.6756 ° W, 21. vi. 2014. JSS 2014.064 - 12 [‘ - 12 ’ handwritten]. J. S. Sproul & Family’ [white paper], ‘ David R. Maddison DNA 4250 DNA Voucher’ [pale green paper], ‘ HOLOTYPE Bembidion oromaia Sproul + Maddison 2017 ’ [partly handwritten, red paper], ‘ Sequoia and Kings Canyon National Parks SEKI 23092 ’ [green paper], ‘ Oregon State Arthropod Collection OSAC _ 0002000002 [matrix code] ’ [printed on both sides of white paper]. Genitalia mounted in Euparal on small card labelled ‘ DNA 4250 ’ beneath the specimen; extracted DNA stored separately. GenBank accession numbers for DNA sequences of the holotype are KY 950760 (28 S), KY 950889 (CAD), KY 951019 (COI), KY 951149 (MSP), KY 951276 (Topo). Type locality: USA: California: Tulare Co., snowfield above Emerald Lake, 2851 m, 36.5959 ° N 118.6756 ° W. Paratypes: Thirty-two specimens from the following localities, with specimens deposited in CAS, OSAC, and USNM: USA: California: Tulare Co., Upper East Fk. Kaweah River, 2812 m, 36.4189 ° N 118.5927 ° W (12); USA: California: Tulare Co., snow field above Emerald Lake, 2851 m, 36.5959 ° N 118.6756 ° W (6); USA: California: Tulare Co., snowfield below White Chief Lake, 2912 m, 36.417 ° N 118.5941 ° W (3); USA: California: Tulare Co., Lower Franklin Lake, 36.4203 ° N 118.5614 ° W (1); USA: California: Tuolumne Co., Blue Canyon Creek, 2750 m, 38.3151 ° N 119.6613 ° W (2); USA: California: Tuolumne Co., stream draining N. face Leavitt Peak, 2930 m, 38.3098 ° N 119.6619 ° W (2); USA: California: Tuolumne Co., Deadman Creek, 2700 m, 38.3188 ° N 119.6634 ° W (1); USA: California: Tuolumne Co., Hwy. 108, stream SE of Chipmunk Flat, 2440 m (1); USA: California: Tuolumne Co., Deadman Creek at junction with Blue Canyon Creek, 2665 m, 38.3174 ° N 119.6652 ° W (1); USA: California: Mono Co., snow field above Ellery Lake, 37.9345 ° N 119.2318 ° W (2); USA: California: Mono Co., H. M. Hall Natural Area, Lee Vining Creek, 3020 m, 37.9591 ° N 119.2838 ° W (1). Derivation of specific epithet: Derived from Greek roots, with ‘ oro ’ meaning ‘ mountain’, and ‘ maia ’ meaning ‘ good mother’, or ‘ caregiver’. Thus, the name connotes ‘ good mother of the mountain’ or ‘ caregiver of the mountains’. The name recognizes the contribution of Elizabeth C. Sproul to the present work. Mother of George and Pearl mentioned above, Elizabeth has spent hundreds of hours traveling to breve group localities, guiding young legs up steep trails, waiting at trailheads, recording locality data, and collecting and processing specimens in support of JSS’s dissertation research, and to enable positive associations for George and Pearl. Her steady support has added tremendous physical and emotional energy to the sampling efforts and overall scope of this work. The specific epithet also references the beetles. Distributed at higher elevation than any other members of the group, their elegant appearance inspires imagery of a maternal caregiver high in mountains. Diagnosis: This large-bodied black species with long, slender appendages is the most distinctive member of the group (Fig. 3). Forebody and hindbody black, commonly with greenish or bluish metallic lustre. Pronotum narrow relative to elytral width and sinuate laterally; hind angles near 90 ° (Fig. 11 I). Elytra widest behind middle with microsculpture meshes strongly etched in females and moderately etched in males (Fig. 13 C). Legs and antennae black, long and slender; first protarsomeres in males quite small (not illustrated as Fig. 3 is a female). Male genitalia distinctive with outer shape having the appearance of being bent basally; flagellum long and sinuate; sclerite ‘ St’ elongate, bilobed and heavily sclerotized (Figs 9 I, 10 I). Comparison with similar species: Most easily confused with B. laxatum from which it is distinguished by having a pronotum which is narrower relative to the elytra, and more sinuate laterally, longer and more slender legs and antennae, smaller first protarsomeres in males, opaque black coloration, and by various male genitalic characters. Geographic distribution: Throughout the Sierra Nevada, north to Mount Lassen and the Trinity Alps in California (Fig. 17 A). Habitat: Open alpine slopes under fairly large rocks on soil, commonly below patches of melting snow. Also present along small alpine creeks. Geographic variation: The single specimen we sampled from the Trinity Alps in California has two distinctive bases in 28 S (a gene for which all other specimens have identical sequences), but is not notably distinctive in other genes.	en	Sproul, John S., Maddison, David R. (2018): Cryptic species in the mountaintops: species delimitation and taxonomy of the Bembidion breve species group (Coleoptera: Carabidae) aided by genomic architecture of a century-old type specimen. Zoological Journal of the Linnean Society 183: 556-583
