identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
095A87E6FFEF7234798EF908FAFBC357.text	095A87E6FFEF7234798EF908FAFBC357.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha kavanaughi	<div><p>L. kavanaughi evidently being sister groups, a result consistent with their similar male and female genitalia.</p><p>The monophyly of the L. osculans group, the L. erasa group, L. erasa + L. australerasa + L. probata, and L. casta + L. kavanaughi + L. lindrothi are also supported by multigene analyses (STACEY Bayesian analysis, Fig. 8; maximum likelihood tree from a concatenated matrix, Supporting Information, Fig. S1). The multigene trees are also consistent with the phylogeny inferred by Erwin &amp; Kavanaugh (1981).</p><p>28S COI CAD Topo wg MSP ArgK 18S #g Lionepha 100 90 100 100 100 100 100 100 8</p><p>L. erasa group 1, -50 36, -55 56, -38 87, -5 100 3, -79 4 L. eras + aus + prob + disj 13, -50 34, -55 16, -44 72, -14 99 90 4 L. eras + aus + prob 51, -12 54, -22 1, -44 8, -50 24, -45 74, -16 3 L. erasa + australerasa 74, -5 75, -6 73, -5 85, -6 89, -3 96 8 L. casta + kav + lind 31, -32 98 100 87, -4 48, -45 23, -68 3 L. casta + kavanaughi 1, -32 25, -38 16, -45 65, -15 83, -11 93 5 L. osculans group 99 83, -16 65, -16 66, -29 24, -63 100 7</p><p>L. erasa 0, -24 0, -48 0, -61 29, -28 -97 2 L. australerasa 36, -18 0, -48 15, -51 - - 0 L. probata 79, -8 61, -16 96 100 - 6 L. casta 65, -8 55, -30 83, -10 94 81, -18 7 L. kavanaughi 92 59, -28 97 84, -11 99 7 L. lindrothi 99 92 97 23, -27 - 5 L. disjuncta 92 66, -15 100 100 - 7 L. osculans 55, -16 88, -5 - 89, -5 - 6 L. sequoiae 91 85, -7 76, -14 95 - 7 L. pseudoerasa 97 84, -7 99 18, -35 - 6 L. tuulukwa 97 99 97 52, -25 - 6</p></div>	https://treatment.plazi.org/id/095A87E6FFEF7234798EF908FAFBC357	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFF0723779C1FB7EFB49C64A.text	095A87E6FFF0723779C1FB7EFB49C64A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha Casey 1918	<div><p>Lionepha specimens sequenced 143 143 140</p><p>Legend</p><p>143 70 47 52 21</p><p>Black cells: bootstrap support for the clade 90 or above</p><p>Gray cells: bootstrap support for the clade 50-89</p><p>White cells: ML tree has clade, with bootstrap support 0-49</p><p>Pink cells: ML tree has contradictory clade, with bootstrap support 0-49</p><p>Red cells: bootstrap support against clade 50-100</p><p>INDELS IN 28S</p><p>There are data in the DNA sequences not considered in any of the phylogenetic analyses, in the form of indels.The 28S gene, in particular, shows an amount of insertions and deletions in the evolutionary history of Lionepha that is unusual within bembidiines; every species of Lionepha has a unique set of indels. Many of these indels are in a part of the D2 expansion region (Supporting Information, Fig. S2). There is variation within species as well. Lionepha disjuncta, in particular, has striking variation: among the ten specimens sequenced there are eight unique patterns of insertions and deletions over three regions of the gene. There was less variation observed in other species: four bases missing from the northernmost specimen of L. erasa and an extra two bases in the Mt. Hood specimen; an extra two bases in two Oregon and one Washington specimens of L. casta; an extra base in the Utah specimens of L. probata; four indel differences between the California and Oregon specimens of L. tuulukwa .</p><p>MORPHOLOGICAL RESULTS</p><p>The morphological diversity evident among specimens of Lionepha (depicted in Figs 9–19, and described in ‘Taxonomic treatment’, below) corresponds to the patterns evident in the gene trees, with the male genitalic structures in particular (Figs 11–14) being consistent within a proposed species and showing disparities among species.</p><p>CYTOGENETIC RESULTS</p><p>Eight of the species examined have 12 pairs of autosomes, and an X0/XX sex-chromosome system, for a complement of 25 chromosomes in males (Table 5). Preparations of the ninth species examined, Lionepha pseudoerasa (Lindroth, 1963), were insufficient to determine the sex chromosomes,but the25 chromosomes seen in the single male studied were consistent with results from other species. Re-examination of notes taken about the single male of L. casta examined by Maddison (1985) suggest that the tentative male count of 22+XY reported was in error.</p></div>	https://treatment.plazi.org/id/095A87E6FFF0723779C1FB7EFB49C64A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFF7723D798EFA91FD97C1BC.text	095A87E6FFF7723D798EFA91FD97C1BC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha Casey 1918	<div><p>Lionepha Casey, 1918: 18 .</p><p>Type species: Bembidium erasum LeConte, 1859, by original designation.</p><p>Diagnosis: Although DNA sequence data shows Lionepha to be clearly separate from the large genus Bembidion (Maddison, 2012; Maddison et al., 2019), their morphological distinctiveness is not as evident. The arrangement of internal sac sclerites of male genitalia is perhaps the best defining character of the group, but we do not understand homologies of these structures to other bembidiines in sufficient detail to propose synapomorphies of either Lionepha or Bembidion . Female genitalic evolution is even more poorly understood in general, but Lionepha has a bursal characteristic that is likely derived. The dorsal surface of the bursa of female Lionepha has a region that is darker than the surrounding membrane, and is covered with a mat of microtrichia so dense that it is brown (Fig.9). The microtrichia (Fig. 10) are arranged in short rows, point posteriorly and appear to be on the inside of the bursa. This brown region was called the ‘dorsal sclerite’ by Erwin &amp; Kavanaugh (1981), but we call it the ‘dorsal microtrichial patch’. We do not know of any other bembidiines with a similar structure, although the carabid genus Dyscolus ( Platynini) has a dense microtrichial patch encircling the bursa (Moret, 1989). Within trechites, the dorsal microtrichial patch appears to be an autapomorphy of Lionepha . As such, it does not help to place the lineage. In external characters, Lionepha are no more distinctive than many subgenera of Bembidion .</p><p>Nonetheless, adult Lionepha can in general be recognized by the following suite of characters: they are moderate-sized bembidiines (3.3 to 6.0 mm) with unspotted, brown to black bodies. In body form, the smaller members are reminiscent of Phyla (currently considered a subgenus of Bembidion). Head with frontal furrows parallel, not deep and not prolonged onto clypeus; antennae short and thick. Mentum with full, more or less triangular epilobes and triangular or subtriangular mentum tooth. Pronotum with deep basolateral foveae, bounded externally by a strong, forward-converging carina. Lateral margin of elytra not prolonged inside shoulder; recurrent groove short, preapical seta (Ed7B) free; discal setae of elytra confluent with third stria; only the first elytral stria reaches elytral apex, striae 6–8 absent or nearly so. Metasternal process completely bordered. Hind wings in most species full, but dimorphic in L. erasa and L. casta . Female bursa with a dorsal microtrichial patch consisting of a dense, brown mat of microtrichia. Twelve pairs of autosomes, in contrast to the 11 pairs of the vast majority of Bembidion; males lack a Y chromosome (and thus Lionepha have an XO/ XX sex chromosome system).</p><p>Larvae of Lionepha are not distinguishable from larvae of Bembidion . We have studied two first-instar larvae of Lionepha casta (obtained ex ovo from a culture of adults), as well as a single fieldcaught first-instar larva of Lionepha erasa (this is specimen DNA2586; see Figs 5, 6). Lionepha larvae have setae FR4 and FR5 closely approximate on the head, a synapomorphy of Amerizus, Asaphidion and Bembidion (Grebennikov &amp; Maddison, 2005; Maddison, 2012), as well as all other features of Bembidion larvae documented by Grebennikov &amp; Maddison (2005).</p><p>Lionepha is composed of two clades (Fig. 8): the L. erasa group, containing the smaller Lionepha, and the L. osculans group, containing the four species with larger adults. These two groups have distinct male genitalia, with the L. osculans group having a broad apex, and the L. erasa group having a uniform pattern of internal sac sclerites (Figs 11–13).</p><p>The following species of Lionepha are now known (the novel taxa are described below):</p></div>	https://treatment.plazi.org/id/095A87E6FFF7723D798EFA91FD97C1BC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFF9723D7A70FA58FB71C2D9.text	095A87E6FFF9723D7A70FA58FB71C2D9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha lindrothi Maddison & Sproul 2020	<div><p>L. lindrothi Maddison &amp; Sproul</p><p>L. disjuncta (Lindroth, 1963)</p></div>	https://treatment.plazi.org/id/095A87E6FFF9723D7A70FA58FB71C2D9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFF9723D79E8F8EBFE35C064.text	095A87E6FFF9723D79E8F8EBFE35C064.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha probata (Casey 1918)	<div><p>L. probata (Casey, 1918)</p><p>L. casta (Casey, 1918)</p><p>L. kavanaughi Maddison</p></div>	https://treatment.plazi.org/id/095A87E6FFF9723D79E8F8EBFE35C064	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFF9723F7A16F9EEFD91C188.text	095A87E6FFF9723F7A16F9EEFD91C188.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha osculans	<div><p>L. osculans group</p><p>L. osculans (Casey, 1918)</p><p>L. sequoiae (Lindroth, 1963)</p><p>L. pseudoerasa (Lindroth, 1963)</p><p>L. tuulukwa Maddison</p><p>IDENTIFICATION OF SPECIES</p><p>Most Lionepha specimens are difficult to identify to species using external morphological structures. Although the majority of specimens can be identified with good equipment and by the trained eye, many are only identifiable using male genitalia (Figs 11–14), elytral microsculpture (Figs 15–17), or DNA sequences. In most species, there are exceptional specimens that have prothoracic shape, elytral striae or other external characters that are outside the normal bounds of the species, and thus most species contain specimens that cannot be successfully identified using the external traits we have examined. In addition, the most important morphological characters, which are in male genitalia and elytral microsculpture, require special procedures or equipment to observe. The former requires clearing with KOH or enzymes and mounting in a medium with an appropriate refractive index (e.g. Euparal, clove oil or cedarwood oil). Within the L. osculans group, the male genitalia of each species are distinctive (Fig. 13). Within the L. erasa group, genitalia are much more similar, and natural variation in the exact position of internal sac sclerites makes comparisons more difficult, but the shape of sclerite CH 1 (Fig. 14) is often sufficient for identification. A clear view of microsculpture requires a high-quality, powerful microscope and appropriate lighting (either a ring light or diffuse light if a dissecting scope is used, or coaxial illumination if a compound scope is used).</p><p>The large number of tenerals of Lionepha that are captured, more so than is typical for bembidiines, complicates identification. The reason for the high frequency of tenerals is unclear. Perhaps they emerge after eclosion earlier than other bembidiines, and are thus more accessible to collectors, or perhaps they are soft and pale for a longer period. Whatever the cause, tenerals of otherwise dark species will look like paler species (e.g. the tibiae will be paler than a fully developed individual) and their genitalia (male or female) will be more difficult to study. The appendage colour described below pertains to fully developed individuals, not tenerals.</p>KEY TO SPECIES OF LIONEPHA 1. Specimens smaller, body length 3.29–4.42 mm (few specimens longer than 4.2 mm). Microsculpture on elytra absent or consisting of isodiametric or somewhat transverse meshes (Figs 15, 16), but never transverse meshes or close-set transverse lines causing iridescence. Male genitalia with narrower apex, and with the characteristic pattern of internal sclerites shown in Figs 11, 12 ......................................... L. erasa group (2) – Specimens larger, body length 4.26–5.90 mm (few specimens less than 4.4 mm). Microsculpture on elytra consisting of close-set transverse lines causing iridescence (e.g. Fig. 17A) or transverse meshes (e.g. Fig. 17C, D), or somewhat transverse (Fig. 17G, H); never isodiametric or absent. Male genitalia with broader apex (Fig. 13) ................................................................................................................................ L. osculans group (9) 2. Body relatively flat. Prothorax small relative to elytra (Fig. 2D). Each elytron with at least five wellimpressed striae, and with outer striae nearly effaced, but evident. Microsculpture of elytra deeply engraved, even in males (Fig. 16G), with slightly transverse or nearly isodiametric meshes....................... L. disjuncta– Body relatively convex. Prothorax larger relative to elytra (Figs 1, 2A–C). Striae less well-developed in general. Microsculpture less deeply engraved on elytral disc of males (Figs 15A, C, E, 16A, C, E), isodiametric or transverse ........................................................................................................................................................3 3. Males shiny, with microsculpture on elytral disc weak or absent, isodiametric or slightly transverse (in L. australerasa, Fig. 15C; treated under both couplets). Microsculpture on elytra of females isodiametric, or only slightly transverse (at most as in Fig. 15B, D). Aedeagus with dark, distinct nub (arrows in Fig. 14). Female bursa with triangular or lobate dorsal microtrichial patch, much wider posteriorly than anteriorly (Fig. 9A, B) ...........................................................................................................................................................4 – Microsculpture on elytral disc of males easily visible under high magnification with appropriate lighting, with clearly transverse sculpticells (Fig. 16A, C, E, G); microsculpture of females slightly transverse (Fig. 16B, D, F, H). Aedeagus without a nub. Female bursa with rectangular dorsal sclerotized region, not wider posteriorly than anteriorly (Fig. 9C, D)...............................................................................................................6 4. Prothorax broader, with more rounded sides (Fig. 1C). Legs dark, piceous, nearly as dark as body. Male genitalia with ventral surface more curved, near apex more abruptly curved downward (and thus the apex expands towards the tip; Fig. 11E, F). Internal sac of aedeagus with dark nub large (Fig. 14C); sclerite CH 1 as in Fig. 14C. Females with large dorsal microtrichial patch on bursa (Fig. 9A)........................... L. probata– Prothorax narrower relative to elytra, with less rounded sides (Fig. 1A, B). Legs lighter than body, or as dark. Male genitalia with ventral surface straighter, with apex not expanded toward tip (Fig. 11A–D). Internal sac of aedeagus with smaller nub (Fig. 14A, B); sclerite CH 1 as in Figures 14A and B. Females with smaller dorsal microtrichial patch on bursa (Fig. 9B) .....................................................................................................5 5. Prothorax wider (PW/EL 0.467 –0.478,&gt; 0.47 in most specimens), sides more rounded (Fig. 1A); hind angles in most specimens, therefore, more obtuse. Tibiae distinctly paler than body. Microsculpture of elytral disk isodiametric, or nearly so (rarely slightly transverse; Fig. 15E, F). Sclerite CH 1 as in Figure 14A. From Oregon (Coast Range, central Cascades) north to Alaska .................................................................... L. erasa – Prothorax small, narrow (PW/EL 0.438 –0.466, &lt;0.46 in most specimens), more parallel-sided, sides straighter, especially posteriorly (Fig. 1B); in most specimens hind angles thus less obtuse. Tibiae dark, same colour as body. Microsculpture slightly stronger, evident on elytral disk, although faint in males; always slightly transverse (Fig. 15C, D). Sclerite CH 1 as in Figure 14B. Sierra Nevada of California north to Crater Lake, Oregon ............................................................................................................ L. australerasa6. Legs paler, with tibiae clearly paler than body. Dorsal microtrichial patch of female bursa rectangular, not narrowing anteriorly, with multiple, deep, parallel, longitudinal folds (Fig. 9C, E) ....................................7– Legs darker, as dark as, or nearly as dark as, the body; males shinier, with less-impressed sculpticells. Dorsal microtrichial patch of female bursa rectangular and not narrowing anteriorly, without deep, parallel folds, or triangular and lobate, much wider posteriorly than anteriorly (Fig. 9B, D, F) ................8 7. Fourth stria nearly effaced, much less impressed than the first. Sclerite CH 1 as in Figure 14D. Triangular scales on left-most membrane of internal sac (Fig. 18). In the Cascades and westward, as well as the northern and coastal areas of California ............................................................................................ L. casta – Fourth stria only slightly less impressed than the first. Sclerite CH 1 as in Figure 14E. Left-most membrane of internal sac without scales. In the Blue Mountains and Wallowas of Oregon and Washington east to Montana and Wyoming ............................................................................................................. L. kavanaughi 8. Microsculpture of the elytra more deeply engraved, complete sculpticells evident in the male, although slightly effaced; females duller. Sclerite CH 1 as in Figure 14F. Dorsal microtrichial patch of female bursa rectangular and not narrowing anteriorly (Fig. 9D)..................................................................... L. lindrothi – Microsculpture slightly weaker, often without complete sculpticells in males (Fig. 15C, D). Sclerite CH 1 as in Figure 14B. Dorsal microtrichial patch of female bursa triangular and lobate, much wider posteriorly than anteriorly (Fig. 9B) .......................................................................................................... L. australerasa 9. Elytral microsculpture consisting of less transversely stretched meshes, in males with most sculpticells being about three times as wide as tall (Fig. 17G), in females brick-like and deeply engraved (Fig. 17H); thus, the elytra in females are quite dull. Prothorax narrow, only slightly wider than head. Aedeagus with reduced internal sac sclerites (Fig. 13G, H). Oregon Coast Range and Trinity Alps of California ........................................................................................................................................................ L. tuulukwa– Elytral microsculpture more transverse, with less of a tendency to form distinct sculpticells (Fig. 17A–F) .........................................................................................................................................................................10 10. Elytra not iridescent. Males with small basal protarsomeres, only slightly wider than second protarsomere (Fig. 19B) ......................................................................................................................................... L. sequoiae– Elytra slightly to notably iridescent because of transverse microsculpture. Males with large basal protarsomeres, much wider than second protarsomere (Fig. 19A, C)..........................................................11 11. Elytra with reduced elytral striation, with third stria not visible beyond posterior dorsal puncture. Elytra only slightly iridescent. Prothoracic sides less rounded (Fig. 3C), in most specimens with the posterior portion being almost parallel sided. Aedeagus with ventral surface gently curved (Fig. 13E, F) .................................................................................................................................................... L. pseudoerasa – Elytra with at least third stria visible beyond posterior dorsal puncture, and second stria almost reaching apex. Elytra distinctly iridescent. Prothoracic sides more rounded (Fig. 3A). Aedeagus deep, with ventral surface sinuate (Fig. 13A, B).......................................................................................................... L. osculans<p>SPECIES ACCOUNTS</p></div>	https://treatment.plazi.org/id/095A87E6FFF9723F7A16F9EEFD91C188	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFFB723F783CF930FD64C00E.text	095A87E6FFFB723F783CF930FD64C00E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha erasa (LeConte 1859)	<div><p>LIONEPHA ERASA (LECONTE, 1859)</p><p>(FIGS 1A, 15A, B, 11A, B, 14A, 9B, 20)</p></div>	https://treatment.plazi.org/id/095A87E6FFFB723F783CF930FD64C00E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFFB723B798EF8D3FABBC7C7.text	095A87E6FFFB723B798EF8D3FABBC7C7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Bembidium erasum LeConte 1859	<div><p>Bembidium erasum LeConte, 1859: 83 . Lectotype</p><p>♀, designated by Erwin &amp; Kavanaugh (1981: 59), in MCZ (type # 5490), examined, including genitalia and DNA. Type locality: ‘Oregon’ (original citation), restricted to Fort Klamath, Klamath County, Oregon, USA, by Erwin &amp; Kavanaugh (1981: 59). Although Fort Klamath is a reasonable type locality for Erwin &amp; Kavanaugh’s concept of Lionepha erasa (i.e. the species here called Lionepha probata), the species to which the types of Bembidium erasum belong is not known from the southern half of Oregon. In order to move the type locality within the range of the current species, we here change the type locality to Marys Peak, Oregon (44.5104°N 123.5593°W); on this mountain, Lionepha erasa is the only known species of Lionepha with isodiametric microsculpture. GenBank accession numbers for DNA sequences of the lectotype are MN401767, MN401952, MN402005, MN402129 and MN402322; accession number of sequence reads in NCBI’s Sequence Read Archive is SRR5230423.</p><p>Bembidion lindrothellus Erwin &amp; Kavanaugh, 1981: 61 . Holotype ♂ in MCZ (type # 32549), examined, including genitalia and DNA. Type locality: Haines Highway Mile 31.5, Little Boulder Creek, Alaska, USA. New synonymy. GenBank accession numbers for DNA sequences of the lectotype are MN401768, MN401784, MN401905, MN401953, MN402006, MN402130, MN402255 and MN402323; accession numbers of sequence reads in NCBI’s Sequence Read Archive are SRR5230400 and SRR5230401.</p><p>Bembidion lummi Erwin &amp; Kavanaugh, 1981: 62 . Holotype ♀, in CAS (type # 13652), examined. Type locality: Friday Harbour, San Juan Island, San Juan County, Washington, USA. Synonymy with L. chintimini established by Kanda et al. (2015).</p><p>Bembidion chintimini Erwin &amp; Kavanaugh, 1981: 63 . Holotype ♀, in CNC (type # 16452), examined. Type locality: Marys Peak, 1220 m, Benton County, Oregon, USA. New synonymy.</p><p>Nomenclatural notes: Among the non-type specimens we examined, all small Lionepha with isodiametric or effaced microsculpture from Oregon, Washington, Idaho, British Columbia (BC) and Alaska (the area containing the type localities of the four names listed above) belong to two species: a common, widespread species known from southern BC south to California, east to Montana and Colorado, but not known from Alaska or the Oregon Coast Range; and a rarer species known from the Oregon Coast Range, the Cascades of central and northern Oregon, north through Washington, BC, to near Anchorage, Alaska .</p><p>The realization that the four primary types of the abovelisted names belong to only one of these species (the latter, rarer species) took several years, and was delayed by the fact that three of the primary types are females, one of which is teneral (the lectotype of Bembidium erasum), and the fourth type (holotype of Bembidion lindrothellus) is a teneral male with unpigmented, flattened genitalia.</p><p>Investigation of the rarer species, the one here called Lionepha erasa, began in 2010. Dissection of the first recognized males from Marys Peak, Oregon (type locality of Bembidion chintimini) revealed an aedeagus indistinguishable from those from San Juan Island, Washington (type locality of Bembidion lummi). The female holotype of B. chintimini is wingless and has slightly rounded shoulders. However, the Marys Peak population is wing-dimorphic, and winged individuals are in body form no different from the type series of Bembidion lummi . The elytral microsculpture of the holotype of B. chintimini is perfectly isodiametric (against Erwin &amp; Kavanaugh, 1981), thus matching that of B. lummi . Other characters mentioned by Erwin &amp; Kavanaugh as distinguishing the two populations are not consistent with available specimens. The lack of evident morphological differences, combined with effectively identical DNA sequences in specimens from Oregon, British Columbia and Alaska suggested that the Marys Peak populations are the same species as populations further north, and for this reason, Bembidion chintimini and B. lummi were synonymized by Maddison in Kanda et al. (2015).</p><p>This left in question the specimens considered to be Bembidion lindrothellus by Erwin &amp; Kavanaugh, which are at first glance similar to the Marys Peak and other populations of ‘ Bembidion chintimini ’. Specimens classified as Bembidion lindrothellus are reported to be paler, but all specimens mentioned in Erwin &amp; Kavanaugh (1981) are teneral. The unsclerotized aedeagus of the holotype of Bembidion lindrothellus made comparison of internal sac sclerites difficult. However, the internal sac membrane that rests in the left-most position has a species-specific microsculpture in Lionepha, and the microsculpture scales of the holotype of Bembidion lindrothellus from Alaska match those of Marys Peak specimens. A non-teneral male was also collected by Lindroth at the type locality of B. lindrothellus, but was not included in the type series, perhaps as the specimen was housed in Lindroth’s collection in Lund, Sweden.This specimen is presumably the one whose genitalia Lindroth figured as Bembidion brumale (1963: fig. 127f). We have examined that specimen, and it is indistinguishable from specimens of ‘ Bembidion chintimini ’ from Alaska, British Columbia, Washington and Oregon, including details of the internal sac. Most critically, DNA sequences of the holotype of Bembidion lindrothellus are identical in eight studied genes to those of other specimens from throughout the range (Figs 5–7). It is thus evident that the holotypes of Bembidion chintimini, B. lindrothellus and B. lummi belong to a single species.</p><p>However, there is an older name. The type series of Bembidium erasum consists of four females. These specimens have traditionally been considered to belong to the common, widespread species here called Lionepha probata . Females of these two isodiametrically microsculptured species are difficult to tell apart, especially those with less-extreme prothoracic proportions (neither wide nor narrow). Although there are distinctions in the lobe of the female bursa of fully sclerotized individuals, interpretation of tenerals is more tenuous. Specimens in the type series of Bembidium erasum are all teneral, with prothoraces of moderate width, and thus there is no clear morphological evidence to place them to species. The type series was provided by George Suckley (LeConte, 1859), presumably captured during his travels as naturalist for the governor of Washington Territory during 1853–57 (Cooper &amp; Suckley, 1859). The type series is from ‘Oregon’, which at the time encompassed the current area of Oregon, the southern half of what is now Idaho and some parts of Wyoming and Montana (Barry, 1932). Suckley’s travels in Oregon included areas within the range of both species (Cooper &amp; Suckley, 1859), and thus geography provides no clues about species membership. However, DNA data from the lectotype (and two of the paralectotypes; Sproul &amp; Maddison, 2017) makes it clear that these specimens belong to the current species (Figs 5–7; Supporting Information, Fig. S1). Thus, the valid name of this species is Lionepha erasa, with Bembidion chintimini, B. lindrothellus and B. lummi as junior synonyms.</p><p>Diagnosis: A small, convex species (Fig. 1A) with isodiametric elytral microsculpture (Fig. 15A, B; effaced in most males) and pale tibiae contrasting against the darker body. Prothorax moderately wide [PW/EL 0.467 –0.478, average 0.473 (N = 6),&gt; 0.47 in most specimens]. Aedeagus with straight ventral surface, with apex not expanded (Fig. 11A, B). Internal sac of aedeagus with distinct but small nub; sclerite CH 1 as in Figure 14A. Female bursa with relatively small dorsal microtrichial patch (Fig. 9B).</p><p>Most similar to L. australerasa, but also difficult to distinguish externally from L. probata . Compared to both L. australerasa and L. probata, L. erasa is paler, with the tibiae distinctly paler than the body; L. erasa also tends to have a more convex body than in those species. In contrast to L. australerasa, sculpticells of the elytra are more perfectly isodiametric, although in a few specimens they are slightly transverse; the lateral margins of the pronotum of L. erasa are more rounded, with the sides distinctly converging posteriorly, and the hind angle always quite obtuse; the striae of elytra are slightly weaker, with smaller, less distinct punctures, often with only the three inner striae being clearly evident. Compared to L. probata, L. erasa has a narrower prothorax.</p><p>Additional characteristics: Body length 3.47– 4.41 mm, although rarely above 4.2 mm. Antenna piceous. Legs with femora rufopiceous, and tibiae pale rufous or dark testaceous, except in specimens from the Cascades, whose tibiae can be rufopiceous. Hind wings dimorphic, with some individuals fully winged (e.g. DNA2615) and others brachypterous with narrow and sloped shoulders (e.g. DNA2616).</p><p>Geographic variation: Some individuals in populations from inland British Columbia have slightly transverse microsculpture (e.g. 15 km E New Denver, Zincton Summit, BC; MZLU) and are larger than typical for L. erasa . Specimens from the Cascades are darker, with darker tibiae, than specimens from the Oregon Coast Range .</p><p>The only notable variation in DNA sequences observed is in 28S: the two Alaskan specimens (DNA 4059 and the holotype of Bembidion lindrothellus), as well as the lectotype of Bembidion erasum, are missing four bases (‘ATTA’) present in other specimens; this missing section occurs just after base 613 in the sequences of the holotype of Bembidion lindrothellus in GenBank (accession MN402323).</p><p>Note: This is the species referred to as Bembidion brumale in Lindroth (1963), but the type of Bembidion brumale is a member of Lionepha casta .</p><p>Distribution: Known from the Oregon Coast Range from Prairie Peak northward, as well as in the Cascades from Lost Prairie Campground, Oregon, northwards to Mt. St. Helens, at low elevation on the San Juan Islands and Vancouver Island, eastward to the western slopes of the Rocky Mountains and north to east of Anchorage, Alaska (Fig. 20); not known from south of 44.2°N. Found from 15 to 1500 m elevation. Collected in all months of the year except January, with lower frequency in midsummer; most commonly collected in the fall and early spring. In the Oregon Coast Range, where these beetles are found in high-elevation grasslands (e.g. on Marys Peak and Mt. Hebo), they are only active above ground once the rains start in August or September, until the rains cease in late spring. In June and July they are not in evidence, even at night.</p><p>water, a specimen has also been found along the shores of a stream (DNA4144 from Mount Hood, Oregon).</p></div>	https://treatment.plazi.org/id/095A87E6FFFB723B798EF8D3FABBC7C7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFFF72047A77FEA5FC03C49E.text	095A87E6FFFF72047A77FEA5FC03C49E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha australerasa	<div><p>LIONEPHA AUSTRALERASA MADDISON, SP. NOV.</p><p>(FIGS 1B, 15C, D, 11C, D, 14B, 20)</p><p>h t t p:/ / z o o b a n k. o r g / u r n:l s i d: z o o b a n k. o r g: a c t: 57CFECAE-CE5B-4D90-9839-59A71E0B3C07</p><p>Holotype ♂ (OSAC), herein designated, labelled: ‘ USA: California: Amador Co., Oyster Lake, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-120.1186&amp;materialsCitation.latitude=38.6711" title="Search Plazi for locations around (long -120.1186/lat 38.6711)">Silver Lake Cpgd</a>, 2205 m, 38.6711°N 120.1186°W, 31 May 2012. DRM 12.060. D.R. Maddison’, ‘ David R. Maddison DNA3844 DNA Voucher’ [pale green paper] , ‘ HOLOTYPE Lionepha australerasa David R. Maddison’ [partly handwritten, on red paper], ‘ Oregon State Arthropod Collection OSAC _0002000003 [matrix code]’ [printed on both sides of white paper]. Genitalia mounted in Euparal in between coverslips pinned with specimen; extracted DNA stored separately. GenBank accession numbers for DNA sequences of the holotype are KY 246650, KY246686, KY246720, KY246801, KY246842, MN401912, MN401959 and MN402263 .</p><p>Habitat: In the Oregon Coast Range, restricted to high-elevation grasslands (e.g. Fig. 4F), where they can be abundant under small rocks in the spring before the rains end, or early fall after the rains start. At Lost Prairie Campground in the Cascades of Oregon, found at the edge of snow melt in an open, grassy field; on Mt. St. Helens in Washington, found on the open pumice plain. Although typically found in open habitats away from</p><p>Paratypes (25): One paratype from the type locality (OSAC), as well as 24 specimens from the following localities: USA: California: Tulare Co., Sequoia Nat. Park, 6000’, Huckleberry Meadow (2, MZLU) ; USA: California: Amador Co., Carson Spur, 2430 m, 38.7047°N 120.1055°W (3, OSAC); USA, California, Amador County, Highway 88 at Carson Spur, 2430 m, 38.70459°N 120.10554°W (1, CAS); USA: California: El Dorado Co., Martin Meadow, 2305 m, 38.6958°N 120.1223°W (1, OSAC) ; California: El Dorado Co., Strawberry Valley (1, CAS) ; California: El Dorado Co., <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.14628&amp;materialsCitation.latitude=38.78329" title="Search Plazi for locations around (long 120.14628/lat 38.78329)">0.8 miles S of Sciots Camp</a> at Strawberry Creek, 38.78329 120.14628 1745 m (1, CAS: CASENT1043929) ; USA: California: El Dorado Co., trail south of Lily Lake, 2012 m, 38.874°N 120.0804°W (1, OSAC) ; USA: California: Placer Co., creek in Homewood Canyon, 1930 m, 39.0783°N 120.1610°W (3, OSAC) ; USA: California: Tehama Co., <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-121.5607&amp;materialsCitation.latitude=40.3696" title="Search Plazi for locations around (long -121.5607/lat 40.3696)">Nanny Creek</a>, Lassen NF, 1585m, 40.3696°N 121.5607°W (2, OSAC) ; USA: California: Tehama Co., tributary of Mill Creek at Hwy 89, 1996 m, 40.4210°N 121.5333°W (7, OSAC, USNM) ; USA: Oregon: Klamath Co., <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-122.1343&amp;materialsCitation.latitude=42.8987" title="Search Plazi for locations around (long -122.1343/lat 42.8987)">Munson Creek</a>, Crater Lake NP, 1981 m, 42.8987°N 122.1343°W (2, OSAC) .</p><p>Type locality: USA: California: Amador Co., Oyster Lake, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-120.1186&amp;materialsCitation.latitude=38.6711" title="Search Plazi for locations around (long -120.1186/lat 38.6711)">Silver Lake Campground</a>, 2205 m, 38.6711°N 120.1186°W</p><p>Etymology: The epithet is derived from the Latin australis, southern, and erasum, expunged, which is also the specific epithet of a similar species, Lionepha erasa . This species is a southern, close relative of Lionepha erasa .</p><p>Diagnosis: A small, dark species with slightly transverse elytral microsculpture (Fig. 15C, D). The prothorax is small [PW/EL 0.438 –0.466, average 0.45 (N = 9), &lt;0.46 in most specimens] and relatively parallel-sided (Fig. 1B), with less obtuse hind angles. Aedeagus with straighter ventral margin, with apex not expanded (Fig. 11C, D). Internal sac with distinct nub; sclerite CH 1 as in Figure 14B. Dorsal microtrichial patch of bursa relatively small, triangular, narrower anteriorly (as in Fig. 9B). The combination of transverse elytral microsculpture and having a nub in the aedeagus is unique within Lionepha .</p><p>Most similar to L. erasa, but also difficult to distinguish externally from L. probata . Compared to L. erasa, L. australerasa is darker, with tibiae piceous or black in most specimens. Sculpticells of elytra more transverse. Prothorax narrower with less rounded sides, with posterior regions more parallel-sided, and hind angle in most specimens less obtuse. Striae of elytra slightly stronger, with larger, more distinct punctures; striae 4 and 5 usually evident and occasionally outer ones as well. The two species share the same overall shape of the aedeagus (Fig. 11A–D), especially the straight, ventral margin and unexpanded apex. L. erasa and L. australerasa are not known to be sympatric, but their ranges may overlap in the Cascades of Oregon. Most easily distinguished externally from L. probata by the transverse microsculpture and the notably narrower prothorax, which is also more parallel-sided, but some L. probata approach L. australerasa in this regard. L. australerasa has a straighter ventral margin of the aedeagus, without an expanded apex. Females of L. australerasa have a smaller dorsal microtrichial patch on the bursa than those of L. probata . The internal sac sclerites, including the size of the nub and shape of sclerite CH 1, are distinctive in each of the three species (Fig. 14A–C).</p><p>Although not closely related, L. australerasa is most similar in external form to the sympatric L. lindrothi, in that they both have dark legs, slightly or moderately transverse microsculpture on the elytra (although males have somewhat or notably effaced microsculpture, making the shape of sculpticells difficult to observe), and normal-sized or narrow prothoraces. They are difficult to distinguish externally, although L. australerasa has less rounded prothoracic margins and more effaced microsculpture. The lack of a nub in the internal sac of L. lindrothi and the shape of sclerite CH 1 (Fig. 14) allows males to be distinguished; females can be distinguished by the triangular, anteriorly narrowed dorsal microtrichial patch in L. australerasa (as in Fig. 9B) compared to the rectangular dorsal microtrichial patch of L. lindrothi (Fig. 9D, F).</p><p>Additional characteristics: Body length 3.63–4.14 mm. Antenna piceous. Legs with femora piceous, with tibiae piceous or rufopiceous, rarely rufous. Hind wings full-sized. Chromosomes of male 24 + X (Table 5).</p><p>Note: Referred to as Lionepha ‘Carson Spur’ in Sproul &amp; Maddison (2017). Specimens of this species were identified as ‘ Bembidion erasum ’ by Erwin &amp; Kavanaugh (1981).</p><p>Distribution: Known only from the Sierra Nevada of California and Crater Lake, Oregon (Fig. 20); not known from north of 42.9°N. A relatively high-elevation species, found from 1570 to 2925 m elevation. Specimens have been found in May, June and July.</p><p>Habitat: As with the previous species, this species is not closely tied to flowing water, being found on damp loam in forests or their clearings. At the type locality and at Carson Spur found near small patches of damp soil in an open forest, likely where snow had recently melted. At Crater Lake, Oregon, found at night in damp soil under bushes in the small flood plain of Munson Creek.</p></div>	https://treatment.plazi.org/id/095A87E6FFFF72047A77FEA5FC03C49E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFC072057AB5FC27FA90C59E.text	095A87E6FFC072057AB5FC27FA90C59E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha probata (Casey 1918)	<div><p>LIONEPHA PROBATA (CASEY, 1918)</p><p>(FIGS 1C, 15E, F, 11E, F, 14C, 9A, 21)</p><p>Bembidion probatum Casey, 1918: 22 . Lectotype ♀, designated by Lindroth (1975), in USNM (type # 36817), examined. Type locality: Boulder County, Colorado.</p><p>Bembidion lubricum Casey, 1918: 21 . Lectotype ♂, designated by Lindroth (1975), in USNM (type # 36820), examined (including genitalia). Type locality: Truckee, Nevada County, California. Synonymy established, under the name Bembidion erasum, by Lindroth (1963).</p><p>Bembidion lascivum Casey, 1918: 21 . Lectotype ♂, designated by Lindroth (1975), in USNM (type # 36821), examined (including genitalia). Type locality: Lake Tahoe, Placer Country, California. Synonymy established, under the name Bembidion erasum, by Lindroth (1963).</p><p>Nomenclatural notes: This common and widespread species has long been known as Lionepha (or Bembidion) erasa (e.g. Lindroth, 1963; Erwin &amp; Kavanaugh, 1981; Maddison, 2012). However, the lectotype of Bembidium erasum does not belong to this taxon (see nomenclatural notes under Lionepha erasa). We consider the three Casey names listed above to be synonyms. As first revisers of this group, we choose Lionepha probata to be the name for this species. We have sequenced one of the paralectotypes of Bembidion probatum (specimen USNMENT01114822) and have confirmed that it belongs to this species (Figs 5–7; Supporting Information, Fig. S1).</p><p>Diagnosis: Specimens of this species are dark, with a broad prothorax with rounded sides (Fig. 1C) and with relatively effaced, isodiametric microsculpture on the elytra (Fig. 15E, F). As the elytral microsculpture can be nearly absent in many males, the shape of sculpticells can be difficult to observe. Ventral surface of the aedeagus curved, with apex slightly expanded (Fig. 11E, F). Internal sac of aedeagus with large, distinct nub; sclerite CH 1 as in Figure 14C. The dorsal microtrichial patch of the female bursa is lobate or triangular, narrowing anteriorly, and is much larger than in L. erasa and L. australerasa (Fig. 9A). The broad prothorax is indicated by PW/EL values, ranging from 0.463 to 0.498, with an average of 0.476 (N = 8).</p><p>From the similar L. erasa, distinguished by the darker colour, especially of the tibiae, and by the broader prothorax (in most specimens), as well as characteristics of the male genitalia (Figs 11, 14) and bursal lobe of the female genitalia (Fig. 9). For differences with the similarly dark L. australerasa, see the diagnosis under that species. Lionepha probata specimens might also be confused with those of the sympatric L. lindrothi, but the latter has clearly transverse microsculpture and, in general, narrower prothoraces.</p><p>Lionepha probata is also similar in appearance to Bembidion commotum Casey, 1918, with which it is frequently microsympatric. In contrast to L. probata, Bembidion commotum has no elytral microsculpture on the disk, and nearly complete elytral striae. Bembidion commotum also has a metasternal process bordered only laterally.</p><p>Additional characteristics: Body length 3.29–4.16 mm. Antenna piceous. Legs piceous, in a few specimens with slightly paler, rufopiceous tibiae. Hind wings full-sized. Chromosomes of male 24 + X (Table 5).</p><p>Geographic variation: The 28S gene shows two forms that differ by two bases (positions 388 and 404 in the alignment deposited on Dryad). One form (having G at both sites) is eastern, from Colorado and Montana west to the White Mountains of California and the south-eastern corner of the Sierra Nevada, and in the north-west to the Warner, Steens, Wallowas and Blue Mountains. The other form occurs in the Cascades and much of the Sierra Nevada westward. The most distinctive populations are those in Utah, with the four Utah specimens having a one-base insertion in 28S; three of the specimens also differ at two sites from all other L. probata, with the fourth Utah specimen having double-peaks at both of those sites. The four Utah specimens form a distinct, well-supported clade in the COI tree (Fig. 5), as well as the STACEY analysis (Fig. 8).</p><p>Distribution: This widespread species is common from southern British Columbia south to the San Bernadino and San Jacinto Mountains near Los Angeles, east to Colorado (Fig. 21). It is found in the broadest elevation range of any Lionepha species, from 65 to 3400 m. Specimens have been collected from April through November, with the majority being found in July and August.</p><p>Habitat: Found in many habitats close to some form of water. This species is common around the edges of melting snowfields in the Sierra Nevada and Cascades, as well as on the shores of creeks in forested areas (especially among mosses on sand or silt on the upper bank), but it can also occur in other habitats, such as in leaf litter around pools in an open forest floor.</p></div>	https://treatment.plazi.org/id/095A87E6FFC072057AB5FC27FA90C59E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFC172007A97FD34FDB3C37A.text	095A87E6FFC172007A97FD34FDB3C37A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha casta (Casey 1918)	<div><p>LIONEPHA CASTA (CASEY, 1918)</p><p>(FIGS 2A, 16A, B, 12A, B, 14D, 18, 22)</p><p>Bembidion castum Casey, 1918: 20 . Lectotype ♂, designated by Lindroth (1975), in USNM (type # 36818), examined. Type locality: Santa Cruz Mountains, Santa Clara County, California.</p><p>Bembidion serenum Casey, 1918: 21 . Lectotype ♀, designated by Lindroth (1975), in USNM (type # 36819), examined. Type locality: Arcata, Humboldt Country, California. Synonymy established by Lindroth (1963).</p><p>Bembidion brumale Casey, 1918: 22 . Lectotype ♀, designated by Lindroth (1975), in USNM (type # 36823, specimen USNMENT01114819), examined (including genitalia and DNA). Synonymy established by Erwin &amp; Kavanaugh (1981: 52), confirmed through DNA sequences. Type locality: Metlakatla, British Columbia. GenBank accession numbers for DNA sequences of the lectotype are MN402439, MN402246 and MN401772; accession number of sequence reads in NCBI’s Sequence Read Archive is SRR5230408.</p><p>Bembidion vacivum Casey, 1918: 22 . Lectotype ♀, designated by Lindroth (1975), in USNM (type # 36822), examined. Type locality: Skeena River at Terrace, British Columbia. Synonymy established, under the name B. brumale Casey, by Lindroth (1963: 262).</p><p>Bembidion nescium Casey, 1918: 30 . Lectotype ♂, designated by Lindroth (1975), in USNM (type # 36845), examined. Type locality: Metlakatla, British Columbia. Synonymy established by Lindroth (1963).</p><p>Diagnosis: Specimens of this species have relatively pale tibiae, with evident, transverse microsculpture on the elytra; the elytral striae are less impressed, with the fourth stria much less impressed than the first. Sclerite CH 1 as in Figure 14D. Females share with L. kavanaughi a dorsal microtrichial patch of the bursa that is rectangular (not narrowed anteriorly) and with deep, longitudinal, parallel folds (Fig. 9C).</p><p>This species is similar in appearance and genitalia to Lionepha kavanaughi, which with it shares pale tibiae, although the femora of L.casta are on average paler than in L. kavanaughi . However, L. casta has less impressed elytral striae. The most definitive characteristic of the male genitalia is the presence of distinct, triangular scales on the left-most membrane of the internal sac (Fig. 18); in contrast, there are no scales on membranes between sclerite CH 1 and the outer, left wall of the aedeagus in L. kavanaughi or L. lindrothi . They are also geographically disjunct, with L. casta occurring only in the Cascades and westward (Fig. 22), and L. kavanaughi being found only in the north-east corner of Oregon and adjacent Washington east to Montana and Wyoming. From the sympatric L. probata it is most easily distinguished by the narrower prothorax, paler legs and sculpticells that are clearly transverse.</p><p>Additional characteristics: Body length 3.45–4.22 mm. Antenna piceous. Femora rufous or rufopiceous; tibiae rufous or dark testaceous. Hind wings full-sized in most specimens, although a few brachypterous individuals are known (Erwin &amp; Kavanaugh, 1981). Chromosomes of male 24 + X (Table 5).</p><p>Geographic variation: At the easternmost edge of the known range of this species (e.g. from Taneum Creek Campground in Wenatchee National Forest, Washington) specimens are unusually large and dark.</p><p>Distribution: One of the more widespread Lionepha, from coastal California north to southern Alaska (Fig. 22). It has not been found east of the Cascades. We agree with Erwin &amp; Kavanaugh (1981) that Lindroth’s (1963) record from Barkerville, BC, is doubtful. A lower elevation species, found between 0 and 1515 m, with the majority of specimens from below 200 m. It has been collected from March through November, with most specimens having been found in the summer.</p><p>Habitat: In numerous microhabitats often near water: on the sand or gravel shores of creeks in forests; in marshy areas along creeks in mountain forests. They can also be found during springtime in damp areas in high-elevation grasslands far from water.</p></div>	https://treatment.plazi.org/id/095A87E6FFC172007A97FD34FDB3C37A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFC4720179EDFB65FD08C291.text	095A87E6FFC4720179EDFB65FD08C291.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha kavanaughi Maddison 2020	<div><p>LIONEPHA KAVANAUGHI MADDISON, SP. NOV.</p><p>(FIGS 2B, 16C, D, 12C, D, 14E, 9C, E, 22)</p><p>h t t p: / / z o o b a n k. o r g / u r n: l s i d: z o o b a n k. org:act: B7402931-D60B-484A-8231-6DEB41460D52</p><p>Holotype ♂ (OSAC), herein designated, labelled: ‘ USA: Oregon: Wallowa Co., Lostine River Valley, 1483 m 45.3485°N 117.4152°W, 28 July 2016. DRM 16.069. D.R. &amp; W.P. Maddison’, ‘ David R. Maddison DNA5000 DNA Voucher’ [pale green paper] , ‘ HOLOTYPE Lionepha kavanaughi David R. Maddison’ [partly handwritten, on red paper], ‘ Oregon State Arthropod Collection OSAC _0002000004 [matrix code]’ [printed on both sides of white paper]. Genitalia mounted in Euparal in between coverslips pinned with specimen; extracted DNA stored separately. GenBank accession numbers for DNA sequences of the holotype are MN401771, MN401889, MN401946, MN401989, MN402108, MN402235, MN402308 and MN402428 .</p><p>Paratypes (91): A total of 75 additional specimens from the type locality (deposited in OSAC, CAS, CNC, CMNH, USNM, MCZ, MNHN, MZLU, NHMUK, UASM, EMEC, JRLC), as well as 16 specimens from the following localities: USA: Oregon: Union Co., Little Phillips Creek at NF-3734, 1140 m, 45.6285°N 118.0163°W 16.072 (2, OSAC); USA: Oregon: Wallowa Co., Lostine River, Two Pan Trailhead, 1709m 45.249°N 117.3762°W (3, OSAC); USA: Oregon: Wallowa Co., Lostine River Valley, 1526 m, 45.3181°N 117.4015°W (1, OSAC); USA: Oregon: Wallowa Co., Lostine River, Two Pan Trailhead, 45.2490°N 117.3763°W, 1728 m (3, OSAC); USA: Oregon: Wallowa Co, Lostine River (1, OSAC); USA: Washington: Blue Mountains, Lewis Peak (1, OSAC); USA: Montana: Ravalli Co., Nez Perce Fork of Bitterroot River, 3.1 miles E of Nez Perce Pass on Nez Perce Pass Road, 45.73086°N 114.48828°W, 1785 m (1, CAS); USA: Montana: Ravalli Co., Lost Horse Creek, 17.1 miles W of Highway 93 on Lost Horse Road 46.14142°N 114.48584°W, 1752 m (1, CAS); USA: Montana: Ravalli Co., Lost Horse Creek, 12.3 miles W of Highway 93 on Lost Horse Creek, 46.13404°N 114.39418°W, 1513 m (1, CAS); USA: Montana: Sanders Co., Prospect Creek, 5.1 miles E Thompson Pass, 47.57539°N 115.64034°W (1, CAS: CASENT 1048646); USA: Wyoming: Grand Teton National Park (1, UASM).</p><p>Type locality: USA: Oregon: Wallowa Co., Lostine River Valley, 1483 m, 45.3485°N 117.4152°W. The type locality is along a small tributary of the Lostine River, both above and below where it is crossed by Upper Lostine Road (Fig. 4D).</p><p>Etymology: It gives the first author great pleasure to name this species for David H. Kavanaugh, a superb carabidologist, good friend and collector of the first recognized specimens of this species.</p><p>Diagnosis: Specimens of this species have pale tibiae and evident, transverse microsculpture on the elytra (Fig. 16C, D); the elytral striae are relatively impressed (Fig. 2B), with the fourth stria similar in depth to the first. Aedeagus (Fig. 12C, D) without nub on internal sac (Fig. 12E); sclerite CH 1 as in Figure 14E. Females share with L. casta a dorsal microtrichial patch of the bursa that is rectangular (not narrowed anteriorly) and with deep, longitudinal, parallel folds (Fig. 9C).</p><p>This species is similar in appearance and genitalia to Lionepha casta, which with it shares pale tibiae. However, L. kavanaughi has more impressed elytral striae, especially the fourth. They are most easily distinguished by the left-most membrane of the internal sac: L. kavanaughi lacks the obvious triangular scales present in L. casta (Fig. 18). L. kavanaughi occurs further east than any known localities for L. casta (Fig. 22).</p><p>Additional characteristics: This is the largest member of the erasa group, with a body length of 3.81–4.42mm. Antenna piceous. Femora piceous or rufopiceous; tibiae paler, rufous or dark testaceous. Hind wings full-sized. Chromosomes of male 24 + X (Table 5).</p><p>Geographic variation: The two specimens sequenced from Montana are reconstructed as somewhat distinct in the STACEY tree (Fig. 8). These specimens differ by three bases in COI and three bases in wg, all representing synonymous differences in those protein-coding genes.</p><p>Note: Referred to as Lionepha ‘Bitterroots’ in Sproul &amp; Maddison (2017). The specimen referred to in Erwin &amp; Kavanaugh (1981: 54) as an ‘Anomalous record’ of Lionepha casta from Lewis Peak, Blue Mountains, Washington, is a female of this species.</p><p>Distribution: Known from the Bitterroot Mountains along the Montana – Idaho border and Grand Teton National Park in Wyoming, west to the Wallowas and Blue Mountains of north-eastern Oregon and south-eastern Washington. It likely occurs throughout the intervening regions of Idaho, but it has not yet been recorded there. There is a single female labelled ‘Up. Truckee R 8-19-52 PSBartholomew’ in CAS that externally appears most like L. kavanaughi, and it has a folded microtrichial patch matching that of L. kavanaughi or L. casta, but it is far enough outside the known distribution of either L. kavanaughi or L. casta that we consider the record doubtful, and have not included it on the map. Elevational range of the few known localities is between 1085 and 2060 m. Specimens were collected in June and July.</p><p>Habitat: In the Wallowa and Blue Mountains of Oregon, found on the gravel and sand shores of small creeks in forests (e.g. at the type locality, shown in Fig. 4D). Lionepha probata and L. disjuncta are also abundant at the type locality, along with Bembidion kuprianovii Mannerheim.</p><p>LIONEPHA LINDROTHI MADDISON &amp; SPROUL,</p></div>	https://treatment.plazi.org/id/095A87E6FFC4720179EDFB65FD08C291	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFC6720379F3F943FE65C071.text	095A87E6FFC6720379F3F943FE65C071.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha disjuncta (Lindroth 1963)	<div><p>LIONEPHA DISJUNCTA (LINDROTH, 1963)</p><p>(FIGS 2D, 16G, H, 12G, H, 23)</p><p>Bembidion disjunctum Lindroth, 1963: 264 . Holotype ♂ in MCZ (type # 32533), examined. Type locality: Sonora Pass, Tuolumne County, California.</p><p>Diagnosis: A flat, parallel-sided species with more complete striae than other Lionepha (Fig. 2D), and with relatively strong, slightly transverse elytral microsculpture (Fig. 16G, H). Aedeagus (Fig. 12G, H) similar to other members of the erasa group, with a more or less ovoid CH 1 sclerite.</p><p>In appearance L. disjuncta resembles a small member of Bembidion subgenus Plataphus, or a Bembidion nebraskense LeConte, 1863 . From sympatric Bembidion (Plataphus) specimens it can be distinguished externally by the completely bordered metasternal process (bordered at the sides only in Plataphus). The elytral striae are less distinct in L. disjuncta . In Plataphus, multiple striae will be easily evident near the elytral apex. The aedeagus is different from Plataphus, without the evident flagellum present in members of that subgenus. From Bembidion nebraskense it can be immediately distinguished by the presence of elytral microsculpture: B. nebraskense lacks elytral microsculpture and is thus glossy.</p><p>Additional characteristics: Body length 3.62–4.32 mm. Antenna piceous. Femora rufopiceous or piceous, tibiae rufopiceous or rufous. Hind wings full-sized. Chromosomes of male 24 + X (Table 5).</p><p>Geographic variation: This species shows notable variation in 28S, with numerous insertion and deletion events evident (Supporting Information, Fig. S1). Of the ten specimens sequenced, seven have unique indels, and the remaining three share an eighth pattern of insertions and deletions. In addition, the specimen from the Wallowas that was sequenced has a unique amino acid, threonine, within COI, although other L. disjuncta have alanine at that position (base 356 in the sequence of specimen DNA3848, GenBank accession number MN402196).</p><p>Distribution: A widespread species, from the northern Sierra Nevada and Trinity Alps of northern California, north to southern British Columbia and east to Montana (Fig. 23). Found between 650 and 2930 m in elevation. Most records are from late summer and early fall, with specimens having been collected in June through September, as well as one record from April.</p><p>Habitat: This species is usually encountered as a specimen or two amongst more common Bembidion on gravel and cobble shores of cold, clear creeks and rivers. There are two localities where numerous specimens have been found: Summit Creek west of Creston, British Columbia, and in the Lostine River valley in the Wallowas of Oregon. At the latter locality, 36 specimens were found in the drying bed of a small creek in a forest (Fig. 4D), along with numerous Lionepha kavanaughi and L. probata, as well as Bembidion kuprianovii .</p></div>	https://treatment.plazi.org/id/095A87E6FFC6720379F3F943FE65C071	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFC7720C7A8FF931FDE7C019.text	095A87E6FFC7720C7A8FF931FDE7C019.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha osculans (Casey 1918) DNA 2638	<div><p>LIONEPHA OSCULANS (CASEY, 1918)</p><p>(FIGS 3A, 17A, B, 13A, B; 19A, 24A)</p><p>Bembidion osculans Casey, 1918: 20 . Lectotype ♀, designated by Lindroth (1975: 116), in USNM (type # 36816), examined. Type locality: Marin County, California, as restricted by Lindroth (1963).</p><p>Bembidion speculum Casey, 1918: 20 . Lectotype ♀, designated by Lindroth (1975: 116), in USNM (type # 36815), examined. Synonymy established by Lindroth (1963). Type locality: Marin County, California.</p><p>Diagnosis: This is the largest species of Lionepha, with some females reaching nearly 6 mm in length. It is also the broadest, with a wide, rounded prothorax (Fig. 3A). The elytral microsculpture is more transversely stretched than other Lionepha (Fig. 17A, B), yielding a notable iridescence, especially in the males. Elytra with at least the third stria visible beyond the posterior dorsal puncture and the second stria almost reaching the elytral apex. It tends to be slightly darker, with darker legs, than other members of the L. osculans group, but some specimens are paler. Aedeagus deep, with a bulbous appearance because of the sinuate ventral surface (Fig. 13A, B).</p><p>Additional characteristics: Body length 4.68– 5.90 mm. Antennae dark, piceous, although the first antennomere can be dark rufous on the underside. Legs in most specimens rufopiceous, occasionally rufous, darker at the joints. Hind wings full-sized. Chromosomes of male 24 + X (Table 5).</p><p>Geographic variation: The six specimens sequenced from Oregon have three unique bases within the 28S gene; this is evident in the 28S tree in Figure 5, where the Oregon specimens form a distinct clade .</p><p>Note: Hering (1998) reported on the food consumed by ‘ Bembidion osculans ’ on Knowles Creek in Oregon; we examined a selection of specimens from his study, and the majority belong to Lionepha tuulukwa, with a few specimens belonging to L. osculans</p><p>Distribution: A widespread species, common in the Sierra Nevada and coastal areas of California, north to Washington and Idaho (Fig. 24A). As noted by Erwin &amp; Kavanaugh (1981), the record from the Olympic Peninsula in Washington is somewhat doubtful. Commonly encountered between 0 and 2000 m, with a few specimens found up to 2400 m. Found in all months of the year except February; most common in the middle of summer.</p><p>Habitat: By far the commonest large Lionepha, found in a variety of habitats associated with water, including along the shores of creeks, especially in forests and on the edge of melting snowfields in open conifer forests in the Sierra Nevada.</p></div>	https://treatment.plazi.org/id/095A87E6FFC7720C7A8FF931FDE7C019	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFC8720C7A51FF29FC09C2B0.text	095A87E6FFC8720C7A51FF29FC09C2B0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha sequoiae (Lindroth 1963)	<div><p>LIONEPHA SEQUOIAE (LINDROTH, 1963)</p><p>(FIGS 3B, 17C, D, 13C, D, 19B, 24B)</p><p>Bembidion sequoiae Lindroth, 1963: 260 . Holotype ♂ in MCZ (type # 32532), examined. Type locality: Sequoia National Park, California.</p><p>Diagnosis: The most distinctive external feature of this moderately large Lionepha is the exceptionally small basal protarsomeres in males (Fig. 19B), which are only slightly wider than the second protarsomeres, much smaller than in other Lionepha (see other images in Fig. 19). Elytra not iridescent, as sculpticells are not sufficiently transverse (Fig. 17C, D). Aedeagus with broad apex, and with clearly visible dark scales on the internal sac membranes, yielding a speckled appearance (Fig. 13C, D).</p><p>Additional characteristics: Body length 4.31–4.95 mm. Antenna piceous, with first antennomere rufous, at least on the underside. Legs rufous, with darker joints. Hind wings full-sized. Chromosomes of male 24 + X (Table 5).</p><p>Distribution: This species has been found in the Sierra Nevada of California, the Cascades of Oregon and north to Vancouver, British Columbia (Fig. 24B). There are, in addition, two low-elevation records from coastal California (Sonoma County); although one of these records (from 2.5 miles N of Cazadero on King Ridge, Big Austin Creek; CAS) is relatively recent, both records are unexpected and should be confirmed. Found from 30 to 2230 m in elevation. Specimens were collected from March through September.</p><p>Habitat: Found along the gravel and cobble shores of creeks in forests or in forest clearings. Over 80 specimens were found along partly shaded areas of School Creek in the Cascades of Oregon, with most found at the gravel and cobble edge of a splash pool below a culvert .</p></div>	https://treatment.plazi.org/id/095A87E6FFC8720C7A51FF29FC09C2B0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFC8720E7A49FA32FD03C7C4.text	095A87E6FFC8720E7A49FA32FD03C7C4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha pseudoerasa (Lindroth 1963)	<div><p>LIONEPHA PSEUDOERASA (LINDROTH, 1963)</p><p>(FIGS 3C, 17E, F, 13E, F, 19C, 24C)</p><p>Bembidion pseudoerasum Lindroth, 1963: 260 . Holotype ♂ in USNM (type # 76638), examined. Type locality: Truckee, Nevada County, California.</p><p>Diagnosis: This species is most easily distinguished externally by the reduced elytral striation, with third stria not visible beyond the posterior dorsal puncture. Elytra only slightly iridescent. Prothoracic sides moderately rounded, in most specimens with the posterior portion being almost parallel sided (Fig. 3C). Basal protarsomeres of male large, significantly wider than the second tarsomere (Fig. 19C). Aedeagus with ventral surface gently curved (Fig. 11E, F), with internal sac sclerites reduced (although not as much so as in L. tuulukwa).</p><p>In comparison to L. tuulukwa, the prothorax is relatively wider compared to the head (Fig. 3C), and the elytral microsculpture is more transverse (Fig. 17E, F). In comparison to L. osculans, narrower and with a less rounded prothorax, and with less iridescent elytra. Additional characteristics: Body length 4.41– 5.25 mm, with most specimens&gt; 4.6 mm. Antenna piceous, with underside of first antennomere rufous. Legs rufous or dark rufous, with darker joints. Hind wings full-sized. Male with 25 chromosomes (Table 5).</p><p>Distribution: This species is only known from the Sierra Nevada and Trinity Alps of California (Fig. 24C). A mid-elevation species, found from 1200 to 2720 m, from May through August.</p><p>Habitat: This rarely collected species has been found on sand and silt near streams and ponds in forests.</p></div>	https://treatment.plazi.org/id/095A87E6FFC8720E7A49FA32FD03C7C4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
095A87E6FFCA720F79E7FEC8FEA8C7C7.text	095A87E6FFCA720F79E7FEC8FEA8C7C7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lionepha tuulukwa	<div><p>LIONEPHA TUULUKWA MADDISON, SP. NOV.</p><p>(FIGS 3D, 17G, H, 13G, H, 19D, 24C)</p><p>h t t p: / / z o o b a n k. o r g / u r n: l s i d: z o o b a n k. org:act: BC118CC8-0693-4497-87F8-167098DC6EF4</p><p>Holotype ♂ (OSAC), herein designated, labelled: ‘ USA: Oregon: Benton Co., Marys Peak Rd, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-123.5286&amp;materialsCitation.latitude=44.4746" title="Search Plazi for locations around (long -123.5286/lat 44.4746)">Alder Creek Falls</a>, 700 m, 44.4746°N 123.5286°W, 24 September 2010. DRM 10.133. D.R. Maddison’, ‘ David R. Maddison DNA2643 DNA Voucher’ [pale green paper] , ‘ HOLOTYPE Lionepha tuulukwa David R. Maddison’ [partly handwritten, on red paper], ‘ Oregon State Arthropod Collection OSAC _0002000006 [matrix code]’ [printed on both sides of white paper]. Genitalia mounted in Euparal in between coverslips pinned with specimen; extracted DNA and chromosome slide stored separately. GenBank accession numbers for DNA sequences of the holotype are MN401816, MN401919, MN401967, MN402036, MN402162, MN402273 and MN402355 .</p><p>Paratypes (70): In addition to 33 paratypes from the type locality (deposited in OSAC, CAS, CNC, CMNH, USNM, MCZ, MNHN, MZLU, NHMUK, UASM) , we designate 37 paratypes from the following localities: USA: Oregon: Benton Co., Marys Peak, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-123.5282&amp;materialsCitation.latitude=44.4745" title="Search Plazi for locations around (long -123.5282/lat 44.4745)">Alder Creek Falls</a>, 700 m, 44.4745°N 123.5282°W (1, OSAC) ; USA: Oregon: Benton Co., Marys Peak Rd, nr <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-123.528&amp;materialsCitation.latitude=44.4748" title="Search Plazi for locations around (long -123.528/lat 44.4748)">Alder Creek Falls</a>, 700 m, 44.4748°N 123.5280°W (21, OSAC) ; USA: Oregon: Benton Co., Waterfall at mile 2.2 <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-123.5247&amp;materialsCitation.latitude=44.4748" title="Search Plazi for locations around (long -123.5247/lat 44.4748)">Marys Peak Rd</a>, 700 m, 44.4748°N 123.5247°W (5, OSAC) ; USA: Oregon: Lane Co., <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-123.788&amp;materialsCitation.latitude=44.0136" title="Search Plazi for locations around (long -123.788/lat 44.0136)">Knowles Creek</a>, 6.8 km SE Mapleton, 48 m, 44.0136°N 123.7880°W (7, OSAC) ; USA: Oregon: Lane Co., Knowles Creek (3, OSAC) .</p><p>The single specimen from the Trinity Alps of California (USA: California: Trinity Co., Canyon Creek, 1440 m, 40.9490°N 123.0179°W; OSAC) was not designated a paratype.</p><p>Type locality: The type locality is Alder Creek Falls, on the south-east slope of Marys Peak, in the Oregon <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-123.5286&amp;materialsCitation.latitude=44.4746" title="Search Plazi for locations around (long -123.5286/lat 44.4746)">Coast Range</a>, at 44.4746°N 123.5286°W (Fig. 4A) .</p><p>Etymology: The epithet is derived from tuu-lukwa, meaning ‘waterfall’ in the Santiam dialect of Kalapuya (Swadesh, 1965), and it is thus most likely the same word in the Marys River dialect (Henry Zenk, pers. comm., 2013). In addition to describing the habitat of these beetles at the type locality, the name also honours the first people of Marys Peak, who knew these lands so well.</p><p>Diagnosis: Within the L. osculans group it is distinguished by the less transversely stretched meshes on the elytra; in males, most sculpticells are about three times as wide as tall (Fig. 17G); in females, they are brick-like and deeply engraved (Fig. 17H). Because of the exceptionally dull lustre of their elytra, females are among the easiest Lionepha to identify to species. In contrast, the elytral microsculpture is more transverse than any of the smaller Lionepha ( erasa group). Prothorax narrow, only slightly wider than the large head (Fig. 3D). Basal protarsomeres of male large, significantly wider than the second tarsomere (Fig. 19D). Aedeagus (Fig. 13G, H) most similar to that of L. pseudoerasa, but with even more reduced internal sac sclerites.</p><p>Additional characteristics: Body length 4.26– 5.07 mm, with most specimens&gt; 4.5 mm. Surface of elytra in many specimens with a slight metallic reflection, brassy or greenish in colour. Antennae dark, piceous, except for base of first antennomere, which can be dark rufous on the underside. Legs rufous or rufopiceous, darker at the joints. Hind wings full-sized. Chromosomes of male 24 + X (Pflug et al. in review).</p><p>Geographic variation: The single specimen from the Trinity Alps of California (DNA4113) is distinctive in both 28S and COI. In 28S, the California specimen has three insertions not present in Oregon L. tuulukwa, totalling six bases; it also has one five-base deletion relative to the Oregon specimens. In COI, this same specimen has a triplet that codes for valine (at position 305 within its sequence in GenBank, accession MN402204), as do all Lionepha from other species, whereas the Oregon L. tuulukwa have the derived state of isoleucine at that position.</p><p>Note: Referred to as Lionepha ‘Waterfalls’ in Sproul &amp; Maddison (2017) and Pflug et al. (in review). See also the note under Lionepha osculans .</p><p>Distribution: This species is currently known from only three localities: two in the Oregon Coast Range and one in the Trinity Alps of California (Fig. 24C). The Coast Range localities are 50 and 700 m in elevation, with the Trinity Alps locality at 1440 m. Specimens have been found in April, June, August and September. At Alder Creek Falls, they are most commonly encountered in late August and September, after the first autumn rains.</p><p>Habitat: At the type locality, found in the splash zone of Alder Creek Falls (Fig. 4A), and on nearby seeps running down large rockfaces. They crawl among wet moss and soil in these areas, and are most easily found at night. At the other two known localities (Knowles Creek west of Eugene, Oregon, and Canyon Creek in the Trinity Alps of California) they were found under rocks on the gravel and cobble shores of small creeks in forests (Fig. 4B).</p></div>	https://treatment.plazi.org/id/095A87E6FFCA720F79E7FEC8FEA8C7C7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Maddison, David R.;Sproul, John S.	Maddison, David R., Sproul, John S. (2020): Species delimitation, classical taxonomy and genome skimming: a review of the ground beetle genus Lionepha (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1313-1358
