Published August 8, 2024 | Version v1

Systematics and biogeography of Appalachian Anillini, and a taxonomic review of the species of South Carolina (Coleoptera, Carabidae, Trechinae, Anillini)

  • 1. Clemson University, Clemson, United States of America

Description

In the eastern United States, 74 species of Anillini in two genera have been described, with most belonging to Anillinus Casey. Until now, no systematic framework has existed for this large genus, hampering integrative studies. Using DNA sequences from 101 Nearctic species, we present a well-resolved molecular phylogeny supporting a sound systematic framework. Sixteen species groups of Appalachian Anillinus are diagnosed, in part using newly recognized variation in the number of modified male protarsi and the state of the spermathecal duct. We present the first descriptions of Nearctic anilline larvae, which possess none of the synapomorphies of previously described anilline larvae. Within Anillinus, two major clades are mostly consistent with setation of the right paramere: a "hairy clade" with more than four setae, and a "quadrisetose clade." Throughout the phylogeny, microhabitat use varies within each clade, and several endogean lineages are phylogenetically isolated. Our work increases the South Carolina fauna by nearly five-fold. Nine new species are described, Serranillus monadnock sp. nov., Anillinus castaneus sp. nov., Anillinus choestoea sp. nov., Anillinus dentatus sp. nov., Anillinus jancae sp. nov., Anillinus mica sp. nov., Anillinus micamicus sp. nov., Anillinus seneca sp. nov., and Anillinus simplex sp. nov. Several species are newly reported from South Carolina, bringing the total to 20 described species representing seven species groups. Two endemic groups inhabit deep clay soils in the Piedmont and possess unique male sexual characters. The Anillini are a unique component of Nearctic biodiversity, with great potential as a model system for studies of biogeography, secondary male sexual modification, and endogean adaptations.

Files

ZK_article_125897.pdf

Files (17.2 MB)

Name Size Download all
md5:24db3efdc8f1e09f3563d1ffa6a2ffc1
17.2 MB Preview Download

System files (1.1 MB)

Name Size Download all
md5:a29ca3ddc6cc96ef3af5f7908b1cb385
1.1 MB Download

Linked records

Additional details

References

  • Andújar C, Grebennikov VV (2021) Endogean beetles (Coleoptera) of Madagascar: Deep soil sampling and illustrated overview. Zootaxa 4963(2): 317–334. https://doi.org/10.11646/zootaxa.4963.2.4
  • Andújar C, Grebennikov VV, Lencina JL, Serrano J (2010) A new Typhlocharis (Coleoptera: Carabidae: Anillina) from Spain: combining adult and larval morphological data with DNA information. Zootaxa 2485(1): 47. https://doi.org/10.11646/zootaxa.2485.1.4
  • Andújar C, Faille A, Pérez-González S, Zaballos JP, Vogler AP, Ribera I (2016) Gondwanian relicts and oceanic dispersal in a cosmopolitan radiation of euedaphic ground beetles. Molecular Phylogenetics and Evolution 99: 235–246. https://doi.org/10.1016/j.ympev.2016.03.013
  • Andújar C, Pérez-González S, Arribas P, Zaballos JP, Vogler AP, Ribera I (2017) Speciation below ground: Tempo and mode of diversification in a radiation of endogean ground beetles. Molecular Ecology 26(21): 6053–6070. https://doi.org/10.1111/mec.14358
  • Arndt E, Grebennikov VV, Zaballos JMP (1999) Description of the larvae of a representative of Anillina with a key to the Palearctic genera of Bembidiini. Koleopterologische Rundschau 69: 11–19.
  • Barr Jr TC (1967) Antroforceps, an eyeless cave scaritine from Mexico. Coleopterists Bulletin 21(3): 65–70. https://doi.org/10.5962/p.372482
  • Barr Jr TC (1969) Evolution of the (Coleoptera) Carabidae in the Southern Appalachians. In: Holt PC, Roane MK, Parker BC (Eds) The distributional history of the biota of the Southern Appalachians. Part I: Invertebrates. Research Monograph. Virginia Polytechnic Institute and State University, 67–92.
  • Barr Jr TC (1979) Revision of Appalachian Trechus (Coleoptera: Carabidae). Brimleyana 2: 29–75.
  • Barr Jr TC (1995) Notes on some anillines (Coleoptera, Carabidae, Bembidiinae) from Southeastern United States, with descriptions of a new genus and two new species. Special Bulletin of the Japanese Society of Coleopterology 4: 239–248.
  • Bousquet Y (2012) Catalogue of Geadephaga (Coleoptera: Adephaga) of America, north of Mexico. ZooKeys 245: 1–1722. https://doi.org/10.3897/zookeys.245.3416
  • Bousquet Y, Skelley P (2012) Description of two new species of Clivina Latreille (Coleoptera, Carabidae, Clivinini) from southeastern United States. ZooKeys 178: 43–50. https://doi.org/10.3897/zookeys.178.2817
  • Brooks DW, Carlson PH, Cornell JF, Douglass HB, King EW, McCaskill VH, Morse JC, Skelton TE, Spooner J (1979) Status report: Arthropoda other than Crustacea. In: Proceedings of the first South Carolina endangered species symposium. Charleston, S.C., 46–51.
  • Carlton CE (2008) Eight New Species of Arianops Brendel from the Southeastern United States with an Updated Key and Notes on Additional Species (Coleoptera: Staphylinidae: Pselaphinae). Coleopterists Bulletin 62(2): 297–323. https://doi.org/10.1649/1082.1
  • Carlton CE, Chandler DS, Leschen RAB, Riley EG, Skelley PE (2005) Obituary and dedication: Karl Heinz Stephan 1931–2005. Coleopterists Bulletin 59(3): 277–283. https://doi.org/10.1649/0010-065X(2005)059[0277:KHS]2.0.CO;2
  • Caterino MS, Harden CW (2024) A review of Appalachian Dasycerus Brongniart and the recognition of cryptic diversity within Dasycerus carolinensis (Coleoptera: Staphylinidae: Dasycerinae). Insect Systematics and Diversity 8(2)[ixae009]: 1–22. https://doi.org/10.1093/isd/ixae009
  • Caterino MS, Recuero E (2023) Shedding light on dark taxa in sky-island Appalachian leaf litter: Assessing patterns of endemicity using large-scale, voucher-based barcoding. Insect Conservation and Diversity 17(1): 1–15. https://doi.org/10.1111/icad.12697
  • Chakrabarty P, Warren M, Page L, Baldwin C (2013) GenSeq: An updated nomenclature and ranking for genetic sequences from type and non-type sources. ZooKeys 346: 29–41. https://doi.org/10.3897/zookeys.346.5753
  • Cornell JF (1972) The eyeless beetles of North Carolina with notes on their ecology and evolution (Insecta: Coleoptera). Journal of the Elisha Mitchell Scientific Society 88: 260–264.
  • Cornell JF (1977a) Freshwater and terrestrial arthropods: Species of special concern: Insecta; Coleoptera; Carabidae (Ground Beetles). In: Cooper JE, Robinson SS, Funderburg JB (Eds) Endangered and threatened plants and animals of North Carolina: proceedings of the Symposium on Endangered and Threatened Biota of North Carolina, 1, biological concerns, Meredith College, Raleigh, November 7–8, 1975. North Carolina State Museum of Natural History, 221–222.
  • Cornell JF (1977b) Freshwater and terrestrial arthropods: Threatened species: Insecta; Coleoptera; Carabidae (Ground Beetles). In: Cooper JE, Robinson SS, Funderburg JB (Eds) Endangered and threatened plants and animals of North Carolina: proceedings of the Symposium on Endangered and Threatened Biota of North Carolina, 1, biological concerns, Meredith College, Raleigh, November 7–8, 1975. North Carolina State Museum of Natural History, 207–208.
  • Cornell JF (1979) Endangered Coleoptera. In: Proceedings of the first South Carolina endangered species symposium. Charleston, S.C., 107–110.
  • Coyne JA, Orr HA (2004) Speciation. Sinauer Associates, Inc. , Sunderland, Massachusetts, U.S.A., 545 pp.
  • Dajoz R (2005) Les Coléoptères terricoles de trois stations du sud des Appalaches (États-Unis): Structure des peuplements et description de trois espèces nouvelles. Bulletin de la Société Entomologique de France 110(2): 201–212. https://doi.org/10.3406/bsef.2005.16223
  • Donabauer M (2009) New species of the Trechus (Microtrechus) vandykei group from the southern Appalachians. Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 61: 129–141.
  • Draney ML, Milne MA, Ulyshen M, Madriz G (2019) Revision of the spider genus Epiceraticelus (Araneae, Linyphiidae) with a description of a new species. Zootaxa 4646(1): 189–200. https://doi.org/10.11646/zootaxa.4646.1.11
  • DuRant JA, Fox RC (1966) Some arthropods of the forest floor in pine and hardwood forests in the South Carolina Piedmont region. Annals of the Entomological Society of America 59(1): 202–207. https://doi.org/10.1093/aesa/59.1.202
  • Dury C (1902) A revised list of the Coleoptera observed near Cincinnati, Ohio, with notes on localities, bibliographical references and description of six new species. The Journal of the Cincinnati Society of Natural History 20: 107–196.
  • Erwin TL, Kavanaugh D (1981) Systematics and zoogeography of Bembidion Latreille: I. The carlhi and erasum groups of western North America (Coleoptera: Carabidae, Bembidiini). Entomologica Scandinavica (Supplement 15): 33–72.
  • Erwin TL, Sims LL (1984) Carabid beetles of the West Indies (Insects: Coleoptera): A synopsis of the genera and checklists of tribes of Caraboidea, and of the West Indian species. Quaestiones Entomologicae 20: 351–466.
  • Ferro ML, Gimmel ML, Harms KE, Carlton CE (2012) Comparison of the Coleoptera communities in leaf litter and rotten wood in Great Smoky Mountains National Park, USA. Insecta Mundi 0259: 1–58.
  • Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3(5): 294–299.
  • Garrick RC, Newton KE, Worthington RJ (2018) Cryptic diversity in the southern Appalachian Mountains: Genetic data reveal that the red centipede, Scolopocryptops sexspinosus, is a species complex. Journal of Insect Conservation 22(5–6): 799–805. https://doi.org/10.1007/s10841-018-0107-3
  • Giachino PM (2011) A description of two new species of Anillina from North Carolina with notes about the geographical distribution of the genus Anillinus Casey, 1918 (Coleoptera: Carabidae: Bembidiini). Studies and Reports. Taxonomical Series 7: 109–116.
  • Giachino PM (2015) New Anillina from South Africa, Tanzania, Madagascar and Seychelles Islands (Coleoptera: Carabidae, Bembidiini). Fragmenta Entomologica 47(1): 15–31. https://doi.org/10.4081/fe.2015.130
  • Giachino PM, Vailati D (2011) Review of the Anillina of Greece (Coleoptera, Carabidae, Bembidiini). Biodiversity Journal Monograph 1: 1–112.
  • Giachino PM, Eberhard S, Perina G (2021) A rich fauna of subterranean short-range endemic Anillini (Coleoptera, Carabidae, Trechinae) from semi-arid regions of Western Australia. ZooKeys 1044: 269–337. https://doi.org/10.3897/zookeys.1044.58844
  • Grebennikov VV (2002) Description of the first-instar larva of Geocharidius (Coleoptera: Carabidae: Trechitae) with a discussion of the phylogeny of the subtribe Anillina. European Journal of Entomology 99(4): 523–527. https://doi.org/10.14411/eje.2002.068
  • Grebennikov VV, Maddison DR (2005) Phylogenetic analysis of Trechitae (Coleoptera: Carabidae) based on larval morphology, with a description of first-instar Phrypeus and a key to genera. Systematic Entomology 30(1): 38–59. https://doi.org/10.1111/j.1365-3113.2004.00259.x
  • Harden CW (2024) Distribution of Nearctic Anillini (Coleoptera, Carabidae, Trechinae) (Version 01) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.10983000
  • Harden CW, Caterino MS (2024) Revision of the elongatus and pecki species groups of Anillinus Casey, 1918 (Coleoptera, Carabidae, Trechinae, Anillini), with four new species and notes on Anillinus turneri Jeannel, 1963. Insecta Mundi 1057: 1–31.
  • Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, with description of a new species from Bath County, Virginia (Coleoptera, Carabidae, Trechinae, Horologionini). Subterranean Biology 48: 1–49. https://doi.org/10.3897/subtbiol.48.114404
  • Hedin M, Thomas SM (2010) Molecular systematics of eastern North American Phalangodidae (Arachnida: Opiliones: Laniatores), demonstrating convergent morphological evolution in caves. Molecular Phylogenetics and Evolution 54(1): 107–121. https://doi.org/10.1016/j.ympev.2009.08.020
  • Hennen DA, Means JC, Marek PE (2022) A revision of the wilsoni species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae). ZooKeys 1096: 17118. https://doi.org/10.3897/zookeys.1096.73485
  • Hildebrandt D, Maddison D (2011) A new species of Bembidion Latrielle 1802 from the Ozarks, with a review of the North American species of subgenus Trichoplataphus Netolitzky 1914 (Coleoptera, Carabidae, Bembidiini). ZooKeys 147: 261–275. https://doi.org/10.3897/zookeys.147.1872
  • Hlaváč P, Baňař P, Davranoglou L-R, Kempton J (2021) Endogean and Cavernicolous Coleoptera of the Balkans. XXIII. A new species of Paramaurops J. Müller (Coleoptera: Staphylinidae: Pselaphinae) from Greece. Zootaxa 4970(1). https://doi.org/10.11646/zootaxa.4970.1.12
  • Hoffman RL (1963) A new diplopod genus and species from Georgia (Polydesmida: Xystodesmidae). Proceedings of the Biological Society of Washington 76: 113–120.
  • Huang S, Zhou J, Tian M, Faille A (2021) Three new species of the subterranean genus Guiodytes from Guangxi, China (Coleoptera: Carabidae: Clivinini). European Journal of Taxonomy 774: 135–154. https://doi.org/10.5852/ejt.2021.774.1537
  • Jeannel R (1963a) Monographie des '"Anillini"', Bembidiides endogés [Coleoptera Trechidae]. Mémoires du Muséum National d'Histoire Naturelle Série A. Zoologie 28: 33–204.
  • Jeannel R (1963b) Supplément à la monographie des Anillini. Sur quelques espèces nouvelles de l'Amérique du Nord. Revue Française d'Entomologie 30: 145–152.
  • Katoh K, Standley DM (2013) MAFFT Multiple Sequence Alignment Software Version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780. https://doi.org/10.1093/molbev/mst010
  • Keith R, Hedin M (2012) Extreme mitochondrial population subdivision in southern Appalachian paleoendemic spiders (Araneae: Hypochilidae: Hypochilus), with implications for species delimitation. The Journal of Arachnology 40(2): 167–181. https://doi.org/10.1636/A11-49.1
  • Kippenhan MG (2023) The taxonomic status of two species of North American Trogossitidae (Coleoptera: Cleroidea) described by Roger Dajoz. Coleopterists Bulletin 77(2): 278–282. https://doi.org/10.1649/0010-065X-77.2.278
  • LaBonte JR, Maddison DR (2023) Medusapyga LaBonte and Maddison, a new genus of Anillini (Coleoptera: Carabidae: Trechinae) from the Pacific Northwest of the United States. Proceedings of the California Academy of Sciences 67: 401–432.
  • Liebherr JK (2008) Taxonomic revision of Hawaiian Bembidion Latreille (Coleoptera: Carabidae: Bembidiini) with a discussion of their reductive and derivative evolutionary specializations. Annals of the Carnegie Museum 77(1): 31–78. https://doi.org/10.2992/0097-4463-77.1.31
  • Lindroth CH (1966) The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 4. Opuscula Entomologica (Supplementum XXXIII): 409–648.
  • Maddison DR (1993) Systematics of the Holarctic beetle subgenus Bracteon and related Bembidion (Coleoptera: Carabidae). Bulletin of the Museum of Comparative Zoology 153: 143–299.
  • Maddison DR (2012) Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina). Molecular Phylogenetics and Evolution 63(3): 533–576. https://doi.org/10.1016/j.ympev.2012.01.015
  • Maddison DR, Cooper KW (2014) Species delimitation in the ground beetle subgenus Liocosmius (Coleoptera: Carabidae: Bembidion), including standard and next-generation sequencing of museum specimens. Zoological Journal of the Linnean Society 172(4): 741–770. https://doi.org/10.1111/zoj.12188
  • Maddison DR, Maddison WP (2023a) Zephyr: a Mesquite package for interacting with external phylogeny inference programs. http://zephyr.mesquiteproject.org
  • Maddison WP, Maddison DR (2023b) Mesquite: a modular system for evolutionary analysis. Available from: http://www.mesquiteproject.org
  • Maddison DR, Ober KA (2011) Phylogeny of minute carabid beetles and their relatives based upon DNA sequence data (Coleoptera, Carabidae, Trechitae). ZooKeys 147: 229–260. https://doi.org/10.3897/zookeys.147.1871
  • Maddison DR, Kanda K, Boyd OF, Faille A, Porch N, Erwin TL, Roig-Juñent S (2019) Phylogeny of the beetle supertribe Trechitae (Coleoptera: Carabidae): Unexpected clades, isolated lineages, and morphological convergence. Molecular Phylogenetics and Evolution 132: 151–176. https://doi.org/10.1016/j.ympev.2018.11.006
  • Magrini P (2008) Note sul genere Binaghites con descrizione di una nuova specie e considerazioni sugli Scotodipnus delle Alpi Occidentali (Coleoptera Carabidae). Bollettino della Società Entomologica Italiana 140: 139–150.
  • Mammola S, Giachino PM, Piano E, Jones A, Barberis M, Badino G, Isaia M (2016) Ecology and sampling techniques of an understudied subterranean habitat: The Milieu Souterrain Superficiel (MSS). Naturwissenschaften 103(11–12): 88. https://doi.org/10.1007/s00114-016-1413-9
  • Maquet D, Dahan L, Raemdonck H, Dekoninck W, Drumont A (2018) Redécouverte en région de Bruxelles-Capitale du rare carabidé endogé Anillus caecus Jacquelin du Val 1851 au Jardin botanique Jean Massart grâce à la technique du lavage de terre (Coleoptera, Carabidae, Trechinae, Bembidiini, Anillina). Lambillionea 118: 201–210.
  • Moulton JK, Wiegmann BM (2004) Evolution and phylogenetic utility of CAD (rudimentary) among Mesozoic-aged Eremoneuran Diptera (Insecta). Molecular Phylogenetics and Evolution 31(1): 363–378. https://doi.org/10.1016/S1055-7903(03)00284-7
  • Ober KA (2002) Phylogenetic relationships of the carabid subfamily Harpalinae (Coleoptera) based on molecular sequence data. Molecular Phylogenetics and Evolution 24(2): 228–248. https://doi.org/10.1016/S1055-7903(02)00251-8
  • Ortuño VM, Gilgado JD (2011) Historical perspective, new contributions and an enlightening dispersal mechanism for the endogean genus Typhlocharis Dieck 1869 (Coleoptera: Carabidae: Trechinae). Journal of Natural History 45(19–20): 1233–1256. https://doi.org/10.1080/00222933.2011.566944
  • Pérez-González S, Zaballos JP (2019) Structure of male genitalia in a lineage of minute endogean ground beetles: how four new species of Microcharidius Coiffait, 1969 (Coleoptera: Carabidae: Anillini) reveal gradual change and convergent evolution. Arthropod Systematics & Phylogeny 77: 57–86. https://doi.org/10.26049/ASP77-1-2019-04
  • Pérez-González S, Andújar C, Zaballos JP (2018) Hidden biodiversity: Total evidence phylogenetics and evolution of morphological traits in a highly diverse lineage of endogean ground beetles, Typhlocharis Dieck, 1869 (Carabidae, Trechinae, Anillini). Cladistics 34(4): 359–383. https://doi.org/10.1111/cla.12208
  • Schuler L (1971) Les trechinae de France. L'inversion de la valeur systematique des organes genitaux males et femelles. Nouvelle Revue d'Entomologie I: 265–273.
  • Shelley RM (1983) Parvulodesmus prolixogonus, a new genus and species of xystodesmid milliped from South Carolina (Polydesmida). Proceedings of the Biological Society of Washington 96: 121–126.
  • Sherwood WC, Garst FM (2016) Soils of Virginia. In: Bailey CM, Sherwood WC, Eaton LS, Powars DS (Eds) The Geology of Virginia. Special Publication. Virginia Museum of Natural History, 291–313.
  • Shull VL, Vogler AP, Baker MD, Maddison DR, Hammond PM (2001) Sequence alignment of 18S ribosomal RNA and the basal relationships of Adephagan Beetles: Evidence for monophyly of aquatic families and the placement of Trachypachidae. Systematic Biology 50(6): 945–969. https://doi.org/10.1080/106351501753462894
  • Simon C, Frati F, Beckenback A, Crespi B, Liu H, Flook P (1994) Evolution, weighting and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87: 651–701.
  • Slipinski A, Lawrence JF (2013) 1. Introduction. In: Australian Beetles Vol. 1: Morphology, Classification and Keys. CSIRO publishing, Collingwood, Victoria, Australia, 94 pp. https://doi.org/10.1071/9780643097292
  • Sokolov IM (2011) Five new species of Anillinus Casey from the Southern Appalachian Mountains and the Piedmont Plateau of eastern U.S.A. (Coleoptera: Carabidae: Trechinae: Bembidiini). Insecta Mundi 0164: 1–14.
  • Sokolov IM (2012) Five new species of Anillinus Casey from Alabama with a key to the Alabama species (Carabidae: Trechinae: Bembidiini). Annals of the Carnegie Museum 81(1): 61–71. https://doi.org/10.2992/007.081.0104
  • Sokolov IM (2013) A new genus and eight new species of the subtribe Anillina (Carabidae, Trechinae, Bembidiini) from Mexico, with a cladistic analysis and some notes on the evolution of the genus. ZooKeys 352: 51–92. https://doi.org/10.3897/zookeys.352.6052
  • Sokolov IM (2014) A new species of Anillinus Casey from the Oak Ridge area west of the Appalachian mountains in Tennessee (Coleoptera: Carabidae: Trechinae: Bembidiini). Annals of the Carnegie Museum 82(3): 225–229. https://doi.org/10.2992/007.082.0302
  • Sokolov IM (2020) Four new species of the genus Anillinus Casey (Coleoptera, Carabidae, Anillini) from Alabama, U.S.A., with a revised key to the Alabama species. Zootaxa 4808(3): 547–559. https://doi.org/10.11646/zootaxa.4808.3.9
  • Sokolov IM (2021) Two new species of the genus Anillinus Casey (Coleoptera, Carabidae, Anillini) from the southern United States. ZooKeys 1016: 63–76. https://doi.org/10.3897/zookeys.1016.61397
  • Sokolov IM (2022) Two new cavernicolous species of Anillinus Casey (Carabidae, Trechinae, Anillini) from Texas with a revised key to Texas species. Subterranean Biology 44: 153–166. https://doi.org/10.3897/subtbiol.44.91002
  • Sokolov IM (2023) Fourteen new species of the genus Nesamblyops Jeannel (Coleoptera: Carabidae: Anillini) from the South Island of New Zealand with redescription of the genus and description of a new subtribe. Zootaxa 5375(2): 151–192. https://doi.org/10.11646/zootaxa.5375.2.1
  • Sokolov IM, Carlton CE (2008) Two new species of blind, forest litter-inhabiting ground beetles from the subtribe Anillina (Carabidae: Trechinae: Bembidiini) from eastern U.S.A. Zootaxa 1740(1): 37. https://doi.org/10.11646/zootaxa.1740.1.4
  • Sokolov IM, Carlton CE (2010) New species of Anillinus Casey (Carabidae: Trechinae: Bembidiini) from the Southern Appalachians and phylogeography of the A. loweae species group. Zootaxa 2502(1): 1. https://doi.org/10.11646/zootaxa.2502.1.1
  • Sokolov IM, Carlton CE (2012) Species of Anillinus Casey (Coleoptera: Carabidae: Trechinae) described from Brazil and their relation to North American representatives of the genus. Coleopterists Bulletin 66(3): 245–249. https://doi.org/10.1649/072.066.0310
  • Sokolov IM, Schnepp KE (2021) A new subterranean species of Anillinus Casey (Carabidae, Trechinae, Anillini) from Florida. Subterranean Biology 39: 33–44. https://doi.org/10.3897/subtbiol.39.65769
  • Sokolov IM, Watrous LE (2008) A new species and the first record of the genus Anillinus (Carabidae: Trechinae: Bembidiini) from the Ozark region. Coleopterists Bulletin 62(4): 537–543. https://doi.org/10.1649/1114.1
  • Sokolov IM, Carlton C, Cornell JF (2004) Review of Anillinus, with descriptions of 17 new species and a key to soil and litter species (Coleoptera: Carabidae: Trechinae: Bembidiini). Coleopterists Bulletin 58(2): 185–233. https://doi.org/10.1649/611
  • Sokolov IM, Sokolova YY, Carlton CE (2007) New species of Anillinus Casey (Carabidae: Trechinae: Bembidiini) from Great Smoky Mountains National Park, U.S.A. and phylogeography of the A. langdoni species group. Zootaxa 1542(1): 1–20. https://doi.org/10.11646/zootaxa.1542.1.1
  • Sokolov IM, Reddell J, Kavanaugh D (2014) 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. ZooKeys 417: 71–101. https://doi.org/10.3897/zookeys.417.7733
  • Sokolov IM, Carlton CE, Watrous LE, Robison HW (2017) Anillinus alleni Sokolov and Carlton (Coleoptera: Carabidae: Trechinae: Bembidiini), a new species from the Ozark Interior Highlands of Arkansas, USA. Coleopterists Bulletin 71(2): 289. https://doi.org/10.1649/0010-065X-71.2.289
  • Talavera G, Castresana J (2007) Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments. Systematic Biology 56(4): 564–577. https://doi.org/10.1080/10635150701472164
  • Timpe EK, Graham SP, Bonett RM (2009) Phylogeography of the brownback salamander reveals patterns of local endemism in southern Appalachian springs. Molecular Phylogenetics and Evolution 52(2): 368–376. https://doi.org/10.1016/j.ympev.2009.03.023
  • Voss SR, Smith DG, Beachy CK, Heckel DG (1993) Allozyme variation in neighboring isolated populations of the plethodontid salamander Leurognathus marmoratus. Journal of Herpetology 29(3): 493–497. https://doi.org/10.2307/1565011
  • Ward PS, Downie DA (2005) The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed arboreal ants: Phylogeny and evolution of big-eyed arboreal ants. Systematic Entomology 30(2): 310–335. https://doi.org/10.1111/j.1365-3113.2004.00281.x
  • Wild AL, Maddison DR (2008) Evaluating nuclear protein-coding genes for phylogenetic utility in beetles. Molecular Phylogenetics and Evolution 48(3): 877–891. https://doi.org/10.1016/j.ympev.2008.05.023