Evidence for admixture and rapid evolution during glacial climate change in an alpine specialist.

The pace of current climate change is expected to be problematic for alpine flora and fauna, as their adaptive capacity may be limited by small population size. Yet despite substantial genetic drift following post-glacial recolonization of alpine habitats, alpine species are notable for their success surviving in highly heterogeneous environments. Population genomic analyses demonstrating how alpine species have adapted to novel environments with limited genetic diversity remain rare, yet are important in understanding the potential for species to respond to contemporary climate change. In this study, we explored the evolutionary history of alpine ground beetles in the Nebria ingens complex, including the demographic and adaptive changes that followed the last glacier retreat. We first tested alternative models of evolutionary divergence in the species complex. Using millions of genome-wide SNP markers from hundreds of beetles, we found evidence that the Nebria ingens complex has been formed by past admixture of lineages responding to glacial cycles. Recolonization of alpine sites involved a distributional range shift to higher elevation, which was accompanied by a reduction in suitable habitat and the emergence of complex spatial genetic structure. We tested several possible genetic pathways involved in adaptation to heterogeneous local environments using genome scan and genotype-environment association approaches. From the identified genes, we found enriched functions associated with abiotic stress responses, with strong evidence for adaptation to hypoxia-related pathways. The results demonstrate that despite rapid demographic change, alpine beetles in the N. ingens complex underwent rapid physiological evolution.

Phylogeny of the supertribe Nebriitae (Coleoptera, Carabidae) based on analyses of DNA sequence data.

The phylogeny of the carabid beetle supertribe Nebriitae is inferred from analyses of DNA sequence data from eight gene fragments including one nuclear ribosomal gene (28S), four nuclear-protein coding genes (CAD, topoisomerase 1, PEPCK, and wingless), and three mitochondrial gene fragments (16S + tRNA-Leu + ND1, COI ("barcode" region) and COI ("Pat/Jer" region)). Our taxon sample included 264 exemplars representing 241 species and subspecies (25% of the known nebriite fauna), 39 of 41 currently accepted genera and subgenera (all except Notiokasis and Archileistobrius), and eight outgroup taxa. Separate maximum likelihood (ML) analyses of individual genes, combined ML analyses of nuclear, nuclear protein-coding, and mitochondrial genes, and combined ML and Bayesian analyses of the eight-gene-fragment matrix resulted in a well-resolved phylogeny of the supertribe, with most nodes in the tree strongly supported. Within Nebriitae, 167 internal nodes of the tree (out of the maximum possible 255) are supported by maximum-likelihood bootstrap values of 90% or more. The tribes Notiophilini, Opisthiini, Pelophilini, and Nebriini are well supported as monophyletic but relationships among these are not well resolved. Nippononebria is a distinct genus more closely related to Leistus than Nebria. Archastes, Oreonebria, Spelaeonebria, and Eurynebria, previously treated as distinct genera by some authors, are all nested within a monophyletic genus Nebria. Within Nebria, four major clades are recognized: (1) the Oreonebria Series, including eight subgenera arrayed in two subgeneric complexes (the Eonebria and Oreonebria Complexes); (2) the Nebriola Series, including only subgenus Nebriola; (3) the Nebria Series, including ten subgenera arrayed in two subgeneric complexes, the Boreonebria and Nebria Complexes, with the latter further subdivided into three subgeneric subcomplexes (the Nebria, Epinebriola, and Eunebria Subcomplexes)); and (4) the Catonebria Series, including seven subgenera arrayed in two subgeneric complexes (the Reductonebria and Catonebria Complexes). A strong concordance of biogeography with the inferred phylogeny is noted and some evident vicariance patterns are highlighted. A revised classification, mainly within the Nebriini, is proposed to reflect the inferred phylogeny. Three genus-group taxa (Nippononebria, Vancouveria and Archastes) are given revised status and seven are recognized as new synonymies (Nebriorites Jeannel, 1941 and Marggia Huber, 2014 = Oreonebria Daniel, 1903; Pseudonebriola Ledoux & Roux, 1989 = Boreonebria Jeannel, 1937; Patrobonebria Bänninger, 1923, Paranebria Jeannel, 1937 and Barbonebriola Huber & Schmidt, 2017 = Epinebriola Daniel & Daniel, 1904; and Asionebria Shilenkov, 1982 = Psilonebria Andrewes, 1923). Six new subgenera are proposed and described for newly recognized clades: Parepinebriola Kavanaugh subgen. nov. (type species: Nebria delicata Huber & Schmidt, 2017), Insulanebria Kavanaugh subgen. nov. (type species: Nebria carbonaria Eschscholtz, 1829), Erwinebria Kavanaugh subgen. nov. (type species Nebria sahlbergii Fischer von Waldheim, 1828), Nivalonebria Kavanaugh subgen. nov. (type species: Nebria paradisi Darlington, 1931), Neaptenonebria Kavanaugh subgen. nov. (type species: Nebria ovipennis LeConte, 1878), and Palaptenonebria Kavanaugh subgen. nov. (type species: Nebria mellyi Gebler, 1847). Future efforts to better understand relationships within the supertribe should aim to expand the taxon sampling of DNA sequence data, particularly within subgenera Leistus and Evanoleistus of genus Leistus and the Nebria Complex of genus Nebria.

A high-quality carabid genome assembly provides insights into beetle genome evolution and cold adaptation.

The hyperdiverse order Coleoptera comprises a staggering ~25% of known species on Earth. Despite recent breakthroughs in next generation sequencing, there remains a limited representation of beetle diversity in assembled genomes. Most notably, the ground beetle family Carabidae, comprising more than 40,000 described species, has not been studied in a comparative genomics framework using whole genome data. Here we generate a high-quality genome assembly for Nebria riversi, to examine sources of novelty in the genome evolution of beetles, as well as genetic changes associated with specialization to high-elevation alpine habitats. In particular, this genome resource provides a foundation for expanding comparative molecular research into mechanisms of insect cold adaptation. Comparison to other beetles shows a strong signature of genome compaction, with N. riversi possessing a relatively small genome (~147 Mb) compared to other beetles, with associated reductions in repeat element content and intron length. Small genome size is not, however, associated with fewer protein-coding genes, and an analysis of gene family diversity shows significant expansions of genes associated with cellular membranes and membrane transport, as well as protein phosphorylation and muscle filament structure. Finally, our genomic analyses show that these high-elevation beetles have endosymbiotic Spiroplasma, with several metabolic pathways (e.g., propanoate biosynthesis) that might complement N. riversi, although its role as a beneficial symbiont or as a reproductive parasite remains equivocal.

Notes on the genus Xestopus from China, with description of a new species (Carabidae,Sphodrini, Dolichina).

The genus Xestopus Andrewes, 1937 in China is reviewed, with the description of a new species: X. gutangensis Zhu & Kavanaugh, sp. nov. (type locality: Xizang: Mêdog, 29.46414°N, 95.73563°E). The male of X. cyaneus Sciaky & Facchini, 1997 is described for the first time, and the first record of this species in Yunnan, China, represents an eastward range extension for the species. A key is provided for the eight known species of the genus.

Drainage basins serve as multiple glacial refugia for alpine habitats in the Sierra Nevada Mountains, California.

The evolutionary histories of alpine species are often directly associated with responses to glaciation. Deep divergence among populations and complex patterns of genetic variation have been inferred as consequences of persistence within glacier boundaries (i.e., on nunataks), while shallow divergence and limited genetic variation are assumed to result from expansion from large refugia at the edge of ice shields (i.e., massifs de refuge). However, for some species, dependence on specific microhabitats could profoundly influence their spatial and demographic response to glaciation, and such a simple dichotomy may obscure the localization of actual refugia. In this study, we use the Nebria ingens complex (Coleoptera: Carabidae), a water-affiliated ground beetle lineage, to test how drainage basins are linked to their observed population structure. By analysing mitochondrial COI gene sequences and genome-wide single nucleotide polymorphisms, we find that the major drainage systems of the Sierra Nevada Mountains in California best explain the population structure of the N. ingens complex. In addition, we find that an intermediate morphotype within the N. ingens complex is the product of historical hybridization of N. riversi and N. ingens in the San Joaquin basin during glaciation. This study highlights the importance of considering ecological preferences in how species respond to climate fluctuations and provides an explanation for discordances that are often observed in comparative phylogeographical studies.

After 157 years, a second specimen and species of the phylogenetically enigmatic and previously monobasic genus Nototylus Gemminger & Harold, 1868 (Coleoptera, Carabidae, Nototylini).

The enigmatic beetle tribe Nototylini (Carabidae) is revised and a key to species is provided. Two species from South America are included in the genus. One species, Nototylus fryi (Schaum), is reviewed and a second, Nototylus balli Erwin & Kavanaugh, sp. nov., is described as new. Each species is known from a single specimen, neither of which is in good condition. The possible function of what appears to be a unique antennal grooming structure on the front femur is discussed.

ResumenLa tribu enigmática de escarabajos Nototylini (Carabidae) se revisa y se provee una clave para las especies. Dos especies de América del Sur están incluidas bajo este género. Se revisa una especie, Nototylus fryi (Schaum), y una segunda, Nototylus balli sp. nov., se describe como nueva. Ambas especies se conocen de un solo espécimen, ninguno de los cuales está en buenas condiciones. Se discute la posible función de lo que aparece ser una estructura para acicalamiento antenal el fémur anterior.

RésuméUne révision taxonomique de la tribu énigmatique de coléoptères Nototylini (Carabidae) est présentée et une clé pour l'identification d'espèces est fournie. Deux espèces d'Amérique du Sud sont incluses dans le genre. Une espèce est examinée (Nototylus fryi (Schaum)) et, une deuxième est décrite comme nouvelle espèce (Nototylus balli sp. nov.). Chaque espèce est connue à partir d'un seul spécimen dont aucun est en bon état. La possible fonction de ce qui semble être une structure de toilette antennaire sur le pro-fémur est discutée.

ResumoUma enigmática tribo de besouros, Nototylini (Carabidae), é revisada, e uma chave para as espécies de Nototylus é fornecida. O gênero agora inclui duas espécies: Nototylus fryi (Schaum), aqui redescrita, e Nototylus balli sp. nov., descrita como nova. Ambas são conhecidas por somente um espécimen em mal estado de conservação. Discute-se a função de uma estrutura singular do femur anterior, possivelmente usada na escovagem da antena.

Unexpected findings in the Eocene Baltic amber forests: Ground beetle fossils of the tribe Nebriini (Coleoptera: Carabidae).

Occurrence of the ground beetle tribe Nebriini in the Baltic amber forest is confirmed. The nebriine taxon Archaeonebria inexspectata gen. n., sp. n., is described and imaged based on four fossil specimens preserved in the Eocene Baltic amber using light microscopy and X-ray micro-computed tomography. The new taxon shares likely synapomorphies with the amphipacific genus Nippononebria and is therefore proposed as its sister group. The occurrence of Archaeonebria in the Baltic amber forests supports previous assumptions that these forests grew under temperate, at least warm-temperate, climatic conditions.

Beetles in bamboo forests: community structure in a heterogeneous landscape of southwestern Amazonia.

Amazonian bamboo forests dominated by large woody bamboo plants in the genus Guadua cover approximately 180,000 km2 and represent a key resource for many organisms. In southwestern Amazonia, native bamboo forests differ in structure, biodiversity, and growth dynamics from other forest types in the region. However, with the exception of a few species in which habitat specialization or a strong habitat association has been demonstrated, little is known about how bamboo forests influence animal community structure. In an effort to understand more about the animal assemblages associated with Amazonian bamboo forests, we characterized the structure of ground-dwelling beetle assemblages living in bamboo forests and adjacent terra firme forests in a lowland rainforest site in Peru. We conducted intensive pitfall trap surveys in 13 bamboo habitat patches and 13 adjacent terra firme habitat patches to determine if there were differences in the abundance and richness of beetle species in these two habitat types. Additionally, given that southwestern Amazonia experiences distinct dry and wet seasons, we conducted our study during the dry and wet season of one year to account for differences in seasonality. We found a distinct beetle assemblage associated with each forest type, and identified a set of dominant species that significantly contributed to the distinctness in beetle community structure between bamboo and terra firme forest. The terra firme forest had a greater number of rare species than the bamboo forest. Several beetle species exhibited a strong association with the bamboo forest, including a large species of Scarabaeidae that appears to be specializing on bamboo. We also found marked differences in beetle assemblages between dry and wet seasons. Our results support the prediction that beetle community structure in bamboo forest differs from that of terra firme in terms of species richness, abundance, and composition. Bamboo-associated animal communities require more exploration and study, and must be included in regional conservation plans seeking to protect entire animal communities in southwestern Amazonia.

The integripennis species group of Geocharidius Jeannel, 1963 (Carabidae, Bembidiini, Anillina) from Nuclear Central America: a taxonomic review with notes about biogeography and speciation.

Our review recognizes 15 species of the integripennis species group of Geocharidius from Nuclear Central America, include three species previously described (Geocharidiusgimlii Erwin, Geocharidiusintegripennis (Bates) and Geocharidiuszullinii Vigna Taglianti) and 12 described here as new. They are: Geocharidiusandersoni sp. n. (type locality: Chiapas, Chiapas Highlands, Cerro Huitepec) and Geocharidiusvignatagliantii sp. n. (type locality: Chiapas, Motozintla, Sierra Madre de Chiapas, Benito Juárez) from Mexico; Geocharidiusantigua sp. n. (type locality: Sacatepéquez, 5 km SE of Antigua), Geocharidiusbalini sp. n. (type locality: Suchitepéquez, 4 km S of Volcan Atitlán), Geocharidiuserwini sp. n. (type locality: Quiché Department, 7 km NE of Los Encuentros), Geocharidiusjalapensis sp. n. (type locality: Jalapa Department, 4 km E of Mataquescuintla), Geocharidiuslonginoi, sp. n. (type locality: El Progreso Department, Cerro Pinalón), and Geocharidiusminimus sp. n. (type locality: Sacatepéquez Department, 5 km SE of Antigua) from Guatemala; and Geocharidiuscelaquensis sp. n. (type locality: Lempira Department, Celaque National Park), Geocharidiuscomayaguanus sp. n. (type locality: Comayagua Department, 18 km ENE of Comayagua), Geocharidiusdisjunctus sp. n. (type locality: Francisco Morazán, La Tigra National Park), and Geocharidiuslencanus sp. n. (type locality: Lempira Department, Celaque National Park) from Honduras. For all members of the group, adult structural characters, including male and female genitalia, are described, and a taxonomic key for all members of the integripennis species group is presented based on these characters. Behavioral and biogeographical aspects of speciation in the group are discussed, based on the morphological analysis. In all cases of sympatry, pairs of closely related species show greater differences in sizes than pairs of more remotely related species. Integripennis group species occupy six different montane areas at elevations above 1300m, with no species shared among them. Major faunal barriers in the region limiting present species distributions include the Motagua Fault Zone and a gap between the Guatemalan Cordillera volcanic chain and the Honduran Interior Highlands no higher than 900m in elevation. Highest species diversity is in the Guatematan Cordillera (six species), second highest in the Honduran Interior Highlands area (four species).

Life beneath the surface of the central Texan Balcones Escarpment: genus Anillinus Casey, 1918 (Coleoptera, Carabidae, Bembidiini): new species, a key to the Texas species, and notes about their way of life and evolution.

The Texas fauna of the genus Anillinus Casey, 1918 includes three previously described species (A. affabilis (Brues), 1902, A. depressus (Jeannel), 1963 and A. sinuatus (Jeannel), 1963) and four new species here described: A. acutipennis Sokolov & Reddell, sp. n. (type locality: Fort Hood area, Bell County, Texas); A. comalensis Sokolov & Kavanaugh, sp. n. (type locality: 7 miles W of New Braunfels, Comal County, Texas); A. forthoodensis Sokolov & Reddell, sp. n. (type locality: Fort Hood area, Bell County, Texas); A. wisemanensis Sokolov & Kavanaugh, sp. n. (type locality: Wiseman Sink, Hays County, Texas). A key for identification of adults of these species is provided. The fauna includes both soil- and cave-inhabiting species restricted to the Balcones Fault Zone and Lampasas Cut Plain and adjacent areas underlain by the Edwards-Trinity Aquifer. Based on morphological and distributional data, we hypothesize that four lineages of endogean Anillinus species extended their geographical ranges from a source area in the Ouachita-Ozark Mountains to the Balconian region in central Texas. There the cavernous Edwards-Trinity aquifer system provided an excellent refugium as the regional climate in the late Tertiary and early Quaternary became increasingly drier, rendering life at the surface nearly impossible for small, litter-inhabiting arthropods. Isolated within the Edwards-Trinity aquifer system, these anilline lineages subsequently differentiated, accounting for the currently known diversity. The paucity of specimens and difficulty in collecting them suggest that additional undiscovered species remain to be found in the region.

Inventory of the carabid beetle fauna of the Gaoligong Mountains, western Yunnan Province, China: species of the tribe Zabrini (Coleoptera, Carabidae).

A ten-year multidisciplinary, multi-national and multi-institutional biodiversity inventory project in the Gaoligong Shan region of western Yunnan Province, China generated more than 35,000 specimens of the beetle (Coleoptera) family Carabidae. In this report, first of a planned series, we focus on diversity in tribe Zabrini. Our study of just over 1300 specimens of zabrine carabids from the project, all in genus Amara Bonelli, found a total of 13 species, all previously described, to occur in the study area, with none of them strictly endemic. We present a key for identification of adults of these species, as well as nomenclatural data, diagnoses, illustrations of dorsal habitus and male genitalia, and information about geographical, altitudinal and habitat distributions within the study area and overall geographical distribution for each species. Distributions of the species within the study area are compared, and broader geographical range patterns are characterized. We also discuss a possible role of the Gaoligong Shan region as one source area for the present-day fauna of the Himalaya and southern edge of the Qinghai-Xizang (Tibetan) Plateau.

Contribution to knowledge of the genus Chydaeus in Xizang Autonomous Region [Tibet] and Yunnan Province, China (Coleoptera, Carabidae, Harpalini).

Five new species of the genus Chydaeus Chaudoir, 1854 are described from China: Chydaeus fugongensissp. n. (Shibali, Fugong County, Yunnan Province), Chydaeus gutangensissp. n. (Gutang, Medog County, Xizang Autonomous Region [Tibet]), Chydaeus hanmiensissp. n. (Hanmi, Medog County, Xizang Autonomous Region [Tibet]), Chydaeus asetosussp. n. (NE of Fugong, Yunnan Province), and Chydaeus baoshanensissp. n. (N of Baoshan, Yunnan Province). Taxonomic and faunistic notes on eleven other species occurring in Xizang and Yunnan are also provided. Chydaeus shunichii Ito, 2006 is re-described, based on specimens from Lushui County, Yunnan. Chydaeus kumei Ito, 1992 is treated as a subspecies of Chydaeus andrewesi Schauberger, 1932 [NEW STATUS]. The taxonomic status of Chydaeus guangxiensis Ito, 2006 is discussed. The following taxa are recorded from China for the first time: Chydaeus obtusicollis Schauberger, 1932 (Xizang and Yunnan), Chydaeus malaisei Kataev & Schmidt, 2006 (Yunnan), Chydaeus semenowi (Tschitschérine, 1899) (Xizang and Yunnan), Chydaeus andrewesi andrewesi Schauberger, 1932 (Xizang and Yunnan), Chydaeus andrewesi kumei Ito (Yunnan), Chydaeus bedeli interjectus Kataev & Schmidt, 2002 (Xizang), and Chydaeus bedeli vietnamensis Kataev & Schmidt, 2002 (Yunnan).

A new species of Lesticus Dejean, 1828 (Coleoptera, Carabidae) from the Finisterre Range, Papua New Guinea and a key to the genera of pterostichine-like Harpalinae of New Guinea.

Lesticus finisterrae (Carabidae: Pterostichini) sp. n. (type locality: Finisterre Range, Papua New Guinea), is described and characters to differentiate it from other "Trigonotomi" species are given. A key to the genera of pterostichine-like Harpalinae of the island, including all genera of Morionini, Cratocerini, Drimostomatini, Abacetini, Loxandrini and Pterostichini, is provided. The genus Rhytisternus (Pterostichini) is for the first time reported from New Guinea, represented by the likely adventive species Rhytisternus laevis (Macleay). The previously unknown male of Stegazopteryx ivimkaensis Will (Drimostomatini) is described.

A new species of Amara (Coleoptera, Carabidae, Zabrini) from Sichuan Province, China, with additional records for other Amara species from the region.

A new species, Amara (Bradytulus) shalulishanica Hieke & Kavanaugh, sp. n. (type locality: Haizishan Yakou, 29.47366°N, 100.21921°E, 4623 m, Shalulishan, Zhuosang Township, Litang County, Sichuan Province, China) is described and diagnosed. Additional records are provided for 16 other Amara species, each of which represents one of five different geographical distribution types, which are discussed.

A key to species of subgenus Lithochlaenius (Coleoptera, Carabidae, Chlaeniini, Chlaenius), with descriptions of three new species.

Three new species of genus Chlaenius Bonelli subgenus Lithochlaenius Kryzhanovskij are described from China: Chlaenius chuanqianensis Liu & Liang, sp. n. (type locality: Xishui, Guizhou Province), Chlaenius linwensini Liu & Liang, sp. n. (type locality: Fujian Province), and Chlaenius propeagilis Liu & Kavanaugh, sp. n. (type locality: Gaoligongshan, Yunnan Province). Seven species of the subgenus are redescribed: Chlaenius agiloides Jedlicka, Chlaenius formosensis Lorenz, Chlaenius agilis Chaudoir, Chlaenius leishanensis Kirschenhofer, Chlaenius noguchii Bates, Chlaenius rambouseki Lutshnik, and Chlaenius wrasei Kirschenhofer. Additional taxonomic changes include the following: Chlaenius formosanus Jedlicka is treated as a junior synonym of Chlaenius rambouseki Lutshnik and Chlaenius anchomenoides Bates, syn. n. and Chlaenius nuristanus Jedlicka as junior synonyms of Chlaenius agilis Chaudoir, syn. n.Chlaenius latroLaFerté-Sénectère is considered a nomen nudum stat. n. and unavailable, leaving Chlaenius agilisChaudoir as the next available name. Chlaenius nuristanusaberration rubridipesJedlicka is also an unavailable name. Chlaenius formosensisLorenz (=Chlaenius formosanusHabu) is returned to species status stat. n. A key to adults of the 10 known species of subgenus Lithochlaenius is provided.

Two new species of Pterostichus Bonelli subgenus Pseudoferonina Ball (Coleoptera, Carabidae, Pterostichini) from the mountains of central Idaho, U.S.A.

Two new species of Pterostichus Bonelli subgenus Pseudoferonina Ball, are described from the mountains of central Idaho: Pterostichus bousqueti Bergdahl [type locality = small tributaries of South Fork of Payette River watershed, ca. 1170 m (3840 ft), 44.0675°; -115.6822°, near Lowman, Salmon River Mountains, Boise County, Idaho, U.S.A.] and Pterostichus lolo Bergdahl [type locality = Cottonwood/Orogrande Creek, ca. 870 m (2850 ft), 46.5528°; -115.5522°, North Fork of Clearwater River watershed, Clearwater Mountains, near Bungalow, Clearwater County, Idaho, U.S.A.]. Males of Pterostichus bousqueti and Pterostichus lolo are easily distinguished from each other and the seven previously described Pseudoferonina species by the form of the median lobe of the aedeagus, and from most individuals of the other species of Pseudoferonina in Idaho by features of pronotal shape and macrosculpture. Both species appear to be obligate ripicolous hygrophiles, restricted in distribution primarily to the margins of small montane streams in forested areas. Widespread intensive stream surveys for Pseudoferonina over many years indicate the geographic ranges of both species are highly localized, and additional undescribed species may occur in Idaho.

A re-consideration of the taxonomic status of Nebria lacustris Casey (Coleoptera, Carabidae, Nebriini) based on multiple datasets - a single species or a species complex?

This study gathered evidence from principal component analysis (PCA) of morphometric data and molecular analyses of nucleotide sequence data for four nuclear genes (28S, TpI, CAD1, and Wg) and two mitochondrial genes (COI and 16S), using parsimony, maximum likelihood, and Bayesian methods. This evidence was combined with morphological and chorological data to re-evaluate the taxonomic status of Nebria lacustris Casey sensu lato. PCA demonstrated that both body size and one conspicuous aspect of pronotal shape vary simultaneously with elevation, latitude, and longitude and served to distinguish populations from the southern Appalachian highlands, south of the French Broad, from all other populations. Molecular analyses revealed surprisingly low overall genetic diversity within Nebria lacustris sensu lato, with only 0.39% of 4605 bp varied in the concatenated dataset. Evaluation of patterns observed in morphological and genetic variation and distribution led to the following taxonomic conclusions: (1) Nebria lacustris Casey and Nebria bellorum Kavanaugh should be considered distinct species, which is a NEW STATUS for Nebria bellorum. (2) No other distinct taxonomic subunits could be distinguished with the evidence at hand, but samples from northeastern Iowa, in part of the region known as the "Driftless Zone", have unique genetic markers for two genes that hint at descent from a local population surviving at least the last glacial advance. (3) No morphometric or molecular evidence supports taxonomic distinction between lowland populations on the shores of Lake Champlain and upland populations in the adjacent Green Mountains of Vermont, despite evident size and pronotal shape differences between many of their members.