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Phylogenetic systematics of the enigmatic genus Horologion Valentine, with description of a new species from Bath County, Virginia (Coleoptera, Carabidae, Trechinae, Horologionini)
<p><em>Horologion Valentine</em>, one of the rarest and most enigmatic carabid beetle genera in the world, was until now known only from the holotype of <em>Horologion speokoites Valentine</em>, discovered in 1931 in a small cave in West Virginia. A single specimen of a new species from Virginia was collected in 1991, but overlooked until 2018. DNA sequence data from specimens of this new species, <em>Horologion hubbardi sp. nov</em>., collected in 2022 and 2023, as well as a critical examination of the external morphology of both species, allow us to confidently place Horologion in the supertribe Trechitae, within a clade containing <em>Bembidarenini </em>and <em>Trechini</em>. A more specific placement as sister to the <em>Gondwanan Bembidarenini</em> is supported by DNA sequence data. Previous hypotheses placing Horologion in or near the tribes <em>Anillini,</em> <em>Tachyini,</em> <em>Trechini</em>, <em>Patrobini</em>, and <em>Psydrini </em>are rejected. The existence of two species of Horologion on opposite sides of the high mountains of the middle Appalachians suggests that these mountains are where the ancestral Horologionpopulations dispersed from, and predicts the discovery of additional populations and species. All specimens of <em>H. hubbardi</em> were collected in or near drip pools, and most were found dead, suggesting that the terrestrial epikarst, rather than caves, is the true habitat of Horologion, which explains their extreme rarity since epikarst has not been directly sampled. We recognize the tribe Horologionini, a relict lineage without any close relatives known in the Northern Hemisphere, and an important part of Appalachian biodiversity.</p>
Phylogenetic systematics of the enigmatic genus Horologion Valentine, with description of a new species from Bath County, Virginia (Coleoptera, Carabidae, Trechinae, Horologionini)
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Figure 9 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 9 Genitalia of Horologion species AH. speokoites median lobe, left lateral aspect BH. speokoites left paramere, left lateral aspect CH. speokoites median lobe, dorsal aspect DH. hubbardi median lobe, left lateral aspect EH. hubbardi left paramere, left lateral aspect FH. hubbardi median lobe, dorsal aspect GH. hubbardi right paramere, right lateral aspect HH. hubbardi female genitalia, ventral aspect A–C after Valentine (1932). Scale bar: 0.1 mm.
Figure 8 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 8 Protarsi of HorologionAH. hubbardi male holotype, dorsal aspect BH. hubbardi female paratype, dorsal aspect CH. speokoites male holotype, dorsal aspect DH. hubbardi male holotype, ventral aspect EH. hubbardi male holotype, dorsal aspect. Scale bars: 0.10 mm.
Figure 3 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 3 Habitat of species of HorologionA surface habitat at Williams Cave, Bath Co., VA B C. Harden (yellow helmet) and T. Malabad (red helmet) in microhabitat of H. hubbardi voucher CWH-452 in Williams Cave C C. Harden examining live H. hubbardi holotype in Williams Cave D live holotype male of H. hubbardi on surface of drip pool in Williams Cave E surface habitat at Arbuckle Cave, Greenbrier Co., WV F T. Malabad examining flowstone in Arbuckle Cave. Photo credit: C. Harden (A, E), K. Kosič Ficco (B–D), M. Miller (F).
Figure 5 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 5 Support for and against our preferred hypothetical placement of Horologion. Black: clade present in maximum likelihood tree, SBS 90% or greater, UFBoot 95% or greater and SH-aLRT 80 or greater. Grey: clade present in maximum likelihood tree, SBS less than 90%, UFBoot less than 95% and/or SH-aLRT less than 80. Red: clade absent in maximum likelihood tree, most-supported contradictory clade with SBS 50% or greater, UFBoot 95% or greater and SH-aLRT 80 or greater. Pink: clade absent in maximum likelihood tree, most-supported contradictory clade with SBS less than 50%, UFBoot less than 95% and/or SH-aLRT less than 80.
Figure 12 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 12 Scanning electron micrographs of the forebody of Horologion hubbardiA head of voucher CWH-452, dorsal aspect, clypeal setae indicated by white arrows B prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed) C forebody of voucher CWH-452, ventral aspect.
Figure 13 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 13 Distribution map of species of Horologion. Black triangle: Horologion hubbardi, black circle: Horologion speokoites. Grey shaded areas represent exposed karst (Weary and Doctor 2014).
Figure 2 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 2 Horologion hubbardi female paratype, voucher CWH-452 (VMNH), dorsal habitus. Abdomen removed for DNA extraction. Scale bar: 1 mm.
Figure 11 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 11 Elytral characters of Horologion species AH. speokoites holotype, elytra, oblique left lateral aspect BH. hubbardi paratype (CMNH), ditto CH. hubbardi paratype (VMNH), SEM detail of apex of left elytron, ventral aspect DH. hubbardi paratype (VMNH), SEM photo of right elytron, dorsal aspect EH. hubbardi paratype (VMNH), SEM detail of right elytron (area indicated by black box in D). Black arrows: discal setae, white arrows: plica.
Figure 6 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 6 Support for and against previously proposed placements of Horologion. Grey: clade present in maximum likelihood tree, SBS less than 90%, UFBoot less than 95% and/or SH-aLRT less than 80. Red: clade absent in maximum likelihood tree, most-supported contradictory clade with SBS 90% or greater, UFBoot 95% or greater and SH-aLRT 80 or greater. Pink: clade absent in maximum likelihood tree, most-supported contradictory clade with SBS less than 50%, UFBoot less than 95% and/or SH-aLRT less than 80. There were not sufficient sequences available for COI and Topo from our sampled Psydrini.
Figure 14 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 14 Phylogenetic tree of a portion of Trechinae showing proposed synapomorphies of clades. Topology is that of Fig. 4.
Figure 10 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 10 Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH-452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect.
Figure 4 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 4 Summary of the maximum likelihood tree of Trechinae from IQ-TREE analysis of 8-gene concatenated matrix. Standard bootstrap support values are shown below nodes. For complete phylogenetic tree showing details within each clade, see Suppl. material 4.
Supplementary material 4 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Trees from Maximum Likelihood and Bootstrap analyses of single gene and 8-gene matrices
Supplementary material 2 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Taxa sampled for phylogenetic analyses
Supplementary material 3 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Additional photos of Horologion and Bembidarenini
Supplementary material 1 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
PCR primers and protocols
Figure 1 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 1 Horologion speokoites holotype (USNM), dorsal habitus. Scale bar: 1 mm.
Figure 7 from: Harden CW, Davidson RL, Malabad TE, Caterino MS, Maddison DR (2024) Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia. Subterranean Biology 48: 1-49. https://doi.org/10.3897/subtbiol.48.114404
Figure 7 Labels of Horologion speokoites holotype, USNM.
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