A peer-reviewed open-access journal ZooKeys | 137: 133-175 (2022) ANE Rae Te #ZooKeys https:/ / ZOO keys. pensoft.net Launched to accelerate biodiversity research First report of the Euconnus Thomson subgenus Cladoconnus Reitter in the New World, represented by thirteen new Appalachian species (Coleoptera, Staphylinidae, Scydmaeninae) Michael S. Caterino! | Dept of Plant & Environmental Sciences, Clemson University, Clemson, SC 29634, USA Corresponding author: Michael S. Caterino (mcateri@clemson.edu) Academiceditor: Jan Klimaszewski | Received 3 November2022 | Accepted27 November 2022 | Published 22 December2022 https://z00 bank. org/DFA4D0BD-CBA6-49D4-87AC-C741E6C216E9 Citation: Caterino MS (2022) First report of the Euconnus Thomson subgenus Cladoconnus Reitter in the New World, represented by thirteen new Appalachian species (Coleoptera, Staphylinidae, Scydmaeninae). ZooKeys 1137: 133-175. https://doi.org/10.3897/zookeys. 1137.97068 Abstract Thirteen new species of Euconnus Thomson (Staphylinidae: Scydmaeninae: Glandulariini) are described from the southern Appalachian Mts, USA: Euconnus megalops sp. nov., E. vexillus sp. nov., E. cumberlandus sp. nov., £. vetustus sp. nov., E. adversus sp. nov., E. astrus sp. nov., E. cultellus sp. nov., E. falcatus sp. nov., £. cataloochee sp. nov., E. kilmeri sp. nov., E. draco sp. nov., E. tusquitee sp. nov., and E. attritus sp. nov. These share a number of morphological characters with the Old World subgenus Cladoconnus Reitter, representing a diversification of species distinct from anything previously known from the western hemisphere. Most of the species occur at higher elevations, some at the tops of the region’s highest moun- tains, and a few are single-peak endemics. No females of these species are winged, and in several species neither sex is winged. A preliminary phylogeny suggests the wingless species represent a clade within a clade of wing-dimorphic species. Keywords Biodiversity, dark taxa, leaf litter, metabarcoding Copyright Michael S. Caterino. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 134 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Introduction Following a century of near neglect, the hyperdiverse scydmaenine (Staphylinidae) ge- nus Euconnus Thomson has seen something of a resurgence of interest. With over 2500 described species, the genus ranks as one of the most species-rich genera of organisms on the planet. Pawel Jaloszynski (2012 and others) has recently attempted to bring some order to the classification of these species on a global scale, revising the types of most of the 30+ subgenera, elevating, synonymizing, or validating the species of most, motivated by and also acknowledging the fact that many hundreds of undescribed species exist, many of uncertain placement in the cumbersome and inconsistent intra- generic taxonomy. On the North American front, Stephan et al. (2021) recently revised the ‘then’- valid subgenus Napochus, describing and redescribing more than 100 species in the group. Jaloszynski’s (2021) subgeneric revision subsequently demoted Napochus to synonymy with the nominate Euconnus s. str., finding no consistent characters to dis- tinguish them. Synonymizing the monotypic Pycnophus in the same paper, following several other changes (e.g. Jaloszyniski 2017) leaves Euconnus with only three valid sub- genera in the Nearctic, Psomophus Casey, Tetramelus Motschulsky, and Euconnus s. str., compared with 10 subgenera recognized in American Beetles (O’Keefe 2001). Apart from those formerly contained in Napochus, none of these has been revised since the foundational treatment of Casey (1897), with few species even having been described in the interim. Recent work in the higher parts of the southern Appalachian mountains of the southeastern USA revealed numerous new Euconnus species. This on its own was not particularly surprising, but the difficulty of assigning all of them to subgenus was. Several of these species exhibit a sexual dimorphism that turns out to be known from only a single subgenus, Cladoconnus Reitter, hitherto known only from the Palaearctic region (Hlavaé and Stevanovié 2013; Jaloszynski 2018), the males having longitudinal carinae on the inner edges of the 8" and 9" antennomeres. Careful comparison of these species with the other defining characters of the group in Jatoszynski (2018, 2019) sug- gest that this subgenus is, in fact a member of the North American fauna. The subgenus Cladoconnus comprises thirty-seven extant species distributed pre- dominantly in the Western Palaearctic region (although recent discoveries by Hoshina and colleagues (Hoshina 2004; Hoshina et al. 2018; Hoshina and Park 2020) suggest that the east Asian diversity is considerable as well) and one fossil species from ~ 40 MY Baltic amber (Jaloszynski and Perkovsky 2021). Here I report this subgenus for the first time from the western hemisphere, represented by 13 previously undescribed species. At the same time, these species require a redefinition of the subgenus, because they exhibit considerable variability in morphology, notably not all having the most conspicuous defining character of the subgenus, the carinae of the male 8" and 9° antennomeres. This group also has significance from a regional, faunistic perspective, as it seems to represent a wholly undocumented high Appalachian radiation. Such endemic radia- tions are known in numerous other taxa, both arthropods (Barr 1979; Gusarov 2002; Appalachian Euconnus (Cladoconnus) 135 Miller and Wheeler 2005; Wheeler and Miller 2005) and other organisms. Cladoconnus are somewhat unusual among even these, however, in having no other close relatives (at least as far as currently documented) elsewhere in North America. Other regional species of Euconnus sensu stricto (where these species might otherwise have keyed out, in e.g., O’Keefe 2001) do not appear to be closely related, most in the southeastern US exhibiting a distinct suite of sexual dimorphisms, particularly on the male abdominal ventrites. On a regional scale, these separate lineages show no geographical overlap, with Euconnus s. str. records coming mainly from low-lying areas in the coastal plain. Most records of the species assigned to Cladoconnus are from ~ 1000 m or above, and some subgroups primarily above 1500 m in the highly restricted spruce-fir habitats of the highest peaks. In addition to describing these species, I report COI barcode sequences for most of them, several being represented by sequences from multiple localities, allowing a preliminary assessment of their internal phylogenetic relationships, as well as an even more preliminary exploration of their possible relationships among Euconnus species on a broader scale. Materials and methods Specimens used in this paper originated in or are deposited in the following collections: AMNH The American Museum of Natural History, New York; FMNH The Field Museum, Chicago; CNCI The Canadian National Collection of Insects, Ottawa; CUAC The Clemson University Arthropod Collection, Clemson; UNHC The University of New Hampshire Collection, Durham. Most of the 679 specimens used in this study were collected as part of a larger in- ventory of leaf litter inhabiting arthropods. Litter samples were obtained from numer- ous high elevation localities (> 3300 ft or 1000 m) across five states (Virginia, North Carolina, Tennessee, South Carolina, Georgia) during the years 2015-2021, visiting most sites on two different dates, roughly in spring and fall timeframes, and collecting at least three separate litter samples on each visit. Many of the highest elevation sam- ples come from spruce-fir forest, where litter consists of deep decomposing needles, with minor components of deciduous leaves and fine woody debris. But at lower sites, deciduous leaf litter or Rhododendron litters were often sampled. Litter was sifted down to the soil surface (or to a depth where litter was so decayed as to be indistinguish- able from soil, where the interface was not a hard boundary), over an area of ca. 1 m’, through an 8-mm mesh, until a bag of ~ 6 L was filled. Samples were processed in the lab using Berlese-Tullgren funnels, running subsamples until thoroughly dry, ~ 12 hours per batch. Specimens were collected directly into 100% ethanol, and moved to -20 °C storage after each subsample was complete. 136 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Conventions Despite availability of many more specimens of most species, type series are generally restricted to single localities, acknowledging the possibility of cryptic diversity within some of these species. Type specimens are all dissected males or specimens that we've been able to associate with dissected males through DNA sequences. Label data for primary and secondary types are quoted; data for non-type specimens are summarized. Full specimen-level data for all material examined, along with voucher codes, extraction codes, and GenBank accession numbers are detailed in an Excel supplement. Following Hlavac and Stevanovic (2013), measurements conform with the follow- ing conventions: e BL - total body length is a sum of the lengths of head (anterior labral margin to summit of vertex), pronotum and elytra measured separately; e HW - width of head is measured across eyes; e PW & PL - pronotal width and length are the maximum widths and lengths; e EW - elytral width is the maximum width; e EL -— length of elytra along suture. Sequencing Many specimens reported here were processed through a voucher-based high-through- put sequencing protocol in an attempt to generate a barcode database for the high Appa- lachian litter arthropod fauna. Subsequently, I also selected additional specimens, many dry, from older collections to complement these, representing additional localities and potential species. Most barcoded specimens were imaged prior to extraction, with images archived on a Flickr page: https://www.flickr.com/search/?user_id=183480085%40N02 &text=Cladoconnus. The supplementary specimens were not imaged, but other sequenc- ing procedures were similar. Dry specimens were removed from points by soaking in 100% ethanol. Every specimen was subdivided or punctured to permit tissue digestion, and placed in a separate well in a 96-well plate. Tissues were digested with lysis buffer and proteinase K (Omega BioTek, Norcross, GA). The liquid fraction was removed to a new plate, leaving behind the voucher remains for further dissection and archiving. Follow- ing digestion, remains of extracted specimens were recombined with any non-extracted body parts, labelled, assigned unique CUAC (Clemson University Arthropod Collec- tion) identifiers, and curated into the CUAC. ‘The digested tissue mixture was purified using Omega BioTek’s MagBind HDQ Blood and Tissue kit on a Hamilton Microlab Star automated liquid handling system, eluting with 150 pL elution buffer. Analyses reported here include sequences from two separate sequencing ap- proaches, both based on a ‘mini-barcode’, a 421 bp fragment of the mitochondrial COI gene amplified using the primers BF2-BR2 (GCHCCHGAYATRGCHTTYCC & TCDGGRIGNCCRAARAAYCA, respectively; Elbrecht and Leese 2017), corre- sponding to the downstream two-thirds of the standard barcoding region. Primer pairs for each well were tagged with a unique combination of forward and reverse 9 bp indexes Appalachian Euconnus (Cladoconnus) LaF to allow multiplexing, synthesized as part of the primer by Eurofins Genomics (Louis- ville, KY). These indexes were derived from a list provided by Meier et al. (2016). All PCRs were conducted in 12.5 pL volumes (5.6 uL water, 1.25 wL Taq buffer, 1.25 wL dNTP mix [2.5 mM each], 0.4 uwL MgCl [50 mM], 1.5 uL each primer, 0.05—0.09 pL Platinum Tag polymerase, 1-2 pL DNA template, with a 95 °C initial denaturation for 5 minutes, followed by 35-45 cycles of 94 °C (30 sec), 50 °C (30 sec), 72 °C (30 sec), and a5 minute 72 °C final extension on an Eppendorf Gradient Mastercycler. Earlier barcodes (pre-2022) were generated on an Illumina MiSeq, later ones on an Oxford Nanopore MinION sequencer. For Illumina library preparation, PCR products were combined and purified using Omega Bio-Tek’s Mag-Bind Total Pure NGS Kit, in a ratio of 0.7:1 (enriching for fragments >300 bp). Illumina adapters and sequencing primers were ligated to PCR products using New England BioLab’s Blunt/TA Ligase Master Mix. The amplicon+adapter library was again purified using Mag-Bind Total Pure NGS, and quantified using a Qubit fluorometer. This library was sequenced on an Illumina MiSeq using a v.3 2x300 paired-end kit. Nanopore libraries were prepared using the ligation sequencing kit LSK-112 (Oxford Nanopore Technologies, Oxford, UK) and sequenced using a v10.4 flowcell. Illumina reads were processed with bbtools software package (https://jgi.doe.gov/ data-and-tools/bbtools/; v38.87, Bushnell et al. 2017) to merge paired read ends, remove PhiX reads, trim Illumina adapters, filter reads for the correct size, remove reads with quality score<30, cluster sequences by similarity allowing 5 mismatches (~1%), and generate a final matrix in FASTA format. Nanopore reads were basecalled using the ‘super-accurate’ algorithm of Guppy (v6.1.2), then demultiplexed using ONTbarcoder v0.1.9 (Srivathsan et al. 2021), with minimum coverage set at 5. Final barcode sequences were submitted to GenBank, under accession numbers OP779401-— OP779518 (see Suppl. material 1 for details). Phylogeny FASTA files from all sequencing methods were combined and aligned with the online version of Mafft v7 (Katoh et al. 2017) using the ‘auto’ strategy. To attempt to determine the placement of putative Appalachian Cladoconnus species among global Euconnus diversity, sequences were combined with other barcode-region sequences for Euconnus available on BOLD and GenBank, making a combined matrix of 389 sequences. ‘These include 117 American putative Cladoconnus, two European Cladoconnus, 265 other Euconnus sequences (multiple individuals for many species, as well as many unidentified) representing other subgenera (Neonapochus, Tetramelus, Napochus, and Euconnus s. stt., Old World and New), and five outgroups from the also-Glandulariini genus Brachycepsis Brendel. Phylogenetic reconstructions were performed using maximum likelihood (ML) with W-IQ-TREE v. 2.0 (Trifinopoulos et al. 2016), available at http://iqtree.cibiv. univie.ac.at. This program was used also to determine the best substitution model for the data. Analyses used a perturbation strength of 0.4. and an IQ-TREE stopping rule value at 200. Branch support is based on an ultrafast bootstrap analysis (Minh et al. 2013), run with 1,000 bootstrap replicates with a minimum correlation coefficient of 0.99. 138 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Results Taxonomy Genus Exconnus Thomson, 1859 Subgenus Cladoconnus Reitter, 1909 Cladoconnus Reitter, 1909: 226, as subgenus of Euconnus Thomson. Type species: Scy- dmaenus motschulskii Motschulsky, 1837 (subsequent designation by Newton and Franz 1998: 145). Euconnus (Cladoconnus) Reitter: Hlavac and Stevanovi¢ 2013 (diagnosis); Jaloszynski 2018 (diagnosis). Diagnosis. Previous diagnoses of Palaearctic Cladoconnus have focused primarily on the presence in males of serrulate carinae on the inner margins of antennomeres VIII and [X, a character not known elsewhere in Euconnus. Jatoszynski (2018) further noted the unique presence of a bell-shaped, setose pronotum with 3 basal foveae in an inward- pointing triangular arrangement on each side of a short median basal pronotal carina, straddling short sublateral carinae. He also noted a unique mandibular shape in which the outer margin is interrupted where the apical portion narrows abruptly. The species described here do not entirely conform to this diagnosis, showing variability in most of these characters among them. All are small, with total body lengths ranging from 1.3 to 1.7 mm, not differing much between the sexes (Table 1). The highly distinctive male antennal carinae are evident in most of these species (Figs 4E, 6, 9E, 14C, 16C), but are completely absent in a couple that otherwise seem to be related by other morphological characters and by DNA sequences (Figs 1E, 8A, 13E). Most are consistent in having a bell-shaped pronotum, narrowed basally in some, with the triangle pattern of posterior foveae and at least short sublateral carinae (Figs 4F, 9F). A median basal pronotal carina is evident in several (e.g., Fig. 4F), but in others it is so weak as to be essentially absent (e.g., Fig. 9F). Only one species (E. vetustus) exhibits the distinctive bi-arcuate mandible, the others appearing to have simply arcuate mandibles. Similar to all Palaearctic spe- cies, American Cladoconnus have male genitalia with complex, asymmetrical endophallic armature. Metathoracic wings have not been described for European species, but illus- trations and descriptions of body and metathoracic shape suggest that most described species are fully winged in both sexes. Hoshina et al. (2018) notes apparent sexual di- morphism in wingedness in three Asian species. Several of the new American species are flightless in both sexes, and some appear to be flightless in females only. Females associ- ated by sequencing show reduced eyes relative to males, and have smaller, unmodified antennomeres, in several species having the club reduced to only three antennomeres (e.g., Fig. 9D). Due to this variability, it is likely that future revision of Cladoconnus will be necessary, but for now these new species seem to have clear relationships to those in the Old World, and are relatively easy to recognize among New World Euconnus. Appalachian Euconnus (Cladoconnus) | Be) Table |. Measurements of EF. (Cladoconnus) species, in mm. Each cell has separate averages for males / females, with total number measured in the last column. HL HW PL PW EL EW BL n E. megalops 0.4/0.4 0.3/0.3 0.4/0.4 0.3/0.4 0.9/0.9 0.6/0.6 1b. fbs7, 3458 E. falcatus 0.3/0.3 0.2/0.2 0.3/0.3 0.3/0.3 0.7/0.7 0.5/0.5 1.3/1.3 3L3 E. cataloochee 0.3/0.3 0.2/0.2 0.3/0.3 0.3/0.3 0.7/0.7 0.5/0.5 1.4/1.3 3/3 E. tusquitee 0.3 0.3 0.3 0.3 0.7 0.4 1.3 1/0 E. kilmeri 0.3 0.3 0.3 0.3 0.7 0.4 1.3 1/0 E. draco 0.3/0.3 0.2/0.2 0.3/0.3 0.3/0.3 0.7/0.7 0.5/0.5 1.3/1.3 3/2 E. vetustus 0.3/0.4 0.3/0.3 0.3/0.4 0.4/0.4 0.9/0.8 0.6/0.6 ils y leo, eM ote, E. attritus 0.3/0.3 0.3/0.3 0.4/0.4 0.3/0.3 0.8/0.8 0.5/0.5 1.5/1.5 Bd 3 E. astrus 0.4/0.4 0.3/0.3 0.3/0.4 0.3/0.3 0.8/0.9 0.6/0.6 1.5/1.6 S163 E. vexillus 0.4/0.4 0.3/0.3 0.4/0.4 0.4/0.4 0.9/0.9 0.7/0.7 L7hle7 S23 E. adversus 0.4/0.4 0.3/0.3 0.4/0.4 0.3/0.4 0.9/0.9 0.6/0.6 1.6/1.6 3 43 E. cumberlandus 0.4/0.4 0.3/0.3 0.4/0.4 0.4/0.4 0.9/0.9 0.6/0.6 ea by 3/3 E. cultellus 0.3/0.3 0.3/0.2 0.3/0.3 0.3/0.3 0.7/0.7 0.5/0.5 1.3/1.3 ae There is little point writing a key to these species because most can only definitely be identified by male genitalia, with a few externally similar species even sympatric in a few places. There are three main morphotypes, dark and stout (Euconnus vexillus and EF. vetustus), dark with rufescent highlights, more gracile (Euconnus megalops), and small and pale (flightless), with a mix of modified and non-modified male antenno- meres (all remaining species). All share a generally similar form of male genitalia: the basal bulb is large and voluminous, narrowing at the shoulders to a variously tapered median lobe (sensu Stephan et al. 2021; equivalent to the dorsal apical projection of Jatoszynski 2012. The opposite side (morphologically ventral) exhibits a thin, weakly sclerotized compressor plate (sensu Stephan et al. 2021; equivalent to the ventral api- cal projection of Jaloszynski 2012). They all have a crescent-shaped diaphragm plate sclerite, and relatively thin, weakly curving parameres, bearing a small number of api- cal setae. They vary much more, however, in the sclerites of the endophallus, which are always asymmetrical, including two or more long, curving, often opposing hooks or spikes. These may bear secondary hook-like processes, or other variously acute projec- tions. In descriptions ‘upper’ and ‘lower’ are used to refer to the diaphragm and fora- men sides, respectively, and ‘left’ and ‘right’ referring to the structures as drawn (upper side up), not to true morphological position. Euconnus (Cladoconnus) megalops sp. nov. https://zoobank.org/92355F93-458D-455B-B19E-CFEOD3DFE12E Figs, 2As=3 Type material. Holotype 3, deposited in FMNH: “USA: NC: Haywood Co., 35.6721°N, 83.1760°W, Smoky Mts NP, 6150’, Big Cataloochee Mt., xi.5.2020, sift- ed litter, M.Caterino & EEtzler” / “[QR code] CLEMSON-ENT CUAC000135174” 140 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) / “Caterino DNA Voucher Extraction MSC6486, Morphosp. BCat.B.316”. Paratypes (33, CUAC, FMNH, CNCI, UNHC) — 3 9, 6 : same data as type; 1 9, 3 d: “USA: NC: Haywood Co., 35.6686°N, 83.1749°W, Smoky Mts NP, 5725’, Big Cataloochee Mt., vii.14.2020, sifted litter, M.Caterino, EEtzler”; 3 2: “USA: NC: Haywood Co., 35.6722°N, 83.1758°W, Smoky Mts NP, 6155’, Big Cataloochee Mt., vii.14.2020, sifted litter, M.Caterino, EEtzler?; 2 2: “USA: NC: Haywood Co., 35.6414°N, 83.1958°W, SmokyMtsNP, Balsam Mt. Tr., 4752’, xi.5.2020, M.Caterino & EFEtzler, Sifted litter”; 3 9: “USA: NC: Haywood Co., 35.6425°N, 83.2007°W, SmokyMtsNP, Balsam Mt.Tr., 5167’, xi.5.2020, M.Caterino & EEtzler, Sifted litter’; 9 2, 3 2: “USA: NC: Haywood Co., 35.6453°N, 83.2025°W, SmokyMtsNP, Balsam Mt.Tr., 5086’, xi.5.2020, M.Caterino & FEtzler, Sifted litter”. Other material. (229 adults, 7 larvae) GA: Rabun Co., Chattahoochee NF, Rabun Cliffs, 4082 ft., 11-May-2021 (7 9, 3 @); Towns Co., Chattahoochee NE, Brasstown Bald, 4495 ft., 17-Nov-2020 (1 9, 1 3); NC: Buncombe Co. Co., Pisgah National Forest, Big Butt Trail, 5190 ft., 19-Mar-2016 (1 9, 2 3); Cherokee Co., Nantahala National Forest, Hickory Branch trail, 4156 ft., 26-Jul-2015 (1 2); Clay Co., Nanta- hala National Forest, Riley Knob, 4330 ft., 11-May-2020 (1 2); Clay Co., Nantahala National Forest, Shooting Creek Bald, 4809 ft., 11-May-2020 (1 2); Clay Co., Nan- tahala National Forest, Tusquitee Bald, 4656—5015ft, 1-Sep-2020 (4 9, 2 4); Clay Co., Nantahala National Forest, Chunky Gal Trail, 4014 ft., 1-Sep-2020 (2 9, 1 4); Graham Co., Nantahala National Forest, Teyahalee Bald, 4060—4663ft., 12-Apr-2022 (2 9,3 4); Graham Co., Nantahala National Forest, Cherohala Skyway — Wright Ck., 4702 ft., 4-May-2020 (2 9, 3 3); Graham Co., Nantahala National Forest, Huckle- berry Knob, 5491-5522 ft., 4-May-2020 & 13-Oct-2020 (10 9, 5 4); Graham Co., Nantahala National Forest, jct. Indian & Santeetlah Cks., 2770-2833 ft., 24-Jun-2015 (11 2, 10 3); Graham Co., Nantahala National Forest, Joyce Kilmer Forest, 2696— 2942 ft., 20-Jul-2015 (2 2, 5 4); Haywood Co., Blue Ridge Parkway National Park, Mt. Hardy, 6110 ft., 8-Sep-2020 (9 9, 6 3); Haywood Co., Pisgah National Forest, Mountains to Sea Trail, 5540 ft., 8-Sep-2020 (1 2); Haywood Co., Pisgah National Forest, Black Balsam Knob, 6072 ft., 7-May-2018 (1 2); Haywood Co., Blue Ridge Parkway National Park, Richland Balsam Mt., 6207 ft., 11-Sep-2019 (1 9); Haywood Co., Blue Ridge Parkway National Park, Pisgah Mt., 5245 ft., 10-Aug-2021 (1 9, 1 3); Jackson Co., Sumter National Forest, Ellicott Rock Wilderness, Bad Creek trail, 2397 ft., 3-Jun-2015 (1 9, 2 3); Jackson Co., Nantahala National Forest, Whiteside Mt., 4740 ft., 22-Jun-2022 (1 2); Jackson Co., Cashiers, Hwy 64, 3700 ft., 1-Feb-2020 & 16-Feb-2020 (6 9, 2 4); Jackson Co., Nantahala National Forest, Toxaway Mt., 4770 ft., 5-Aug-2020 (2 9, 1 3); Jackson Co., Blue Ridge Parkway National Park, along Blue Ridge Pkwy, 5572 ft., 11-Sep-2019 (2 2); Jackson Co., Balsam Mountain Pre- serve, Doubletop Mountain, 4839 ft., 17-Jun-2015 (3 9, 2 @); Jackson Co., Balsam Mountain Preserve, Sugarloaf Mountain, 4484 ft., 15-Jun-2015 (2 2); Jackson Co., Balsam Mountain Preserve, Boar ridge, 4040 ft., 16-Jun-2015 (4 2, 1 3); Jackson Co., Balsam Mountain Preserve, Dark ridge, 3290 ft., 20-Jun-2015 (2 9, 7 &); Jackson Co., Blue Ridge Parkway National Park, Waterrock Knob, 6281 ft., 29-May-2018 (1 9, Appalachian Euconnus (Cladoconnus) 141 1 3); Macon Co., E Highlands, Hwy 64, 3880-3990 ft., 1-Mar-2020 (6 9, 2 3); Macon Co., Nantahala National Forest, Jones Gap, 4447 ft., 16-Jul-2015 (1 2); Ma- con Co., Nantahala National Forest, Jones Knob, 4237 ft., 28-Jul-2015 (1 2); Macon Co., nr. Wayah Bald, 5280 ft., 16-Mar-2016 (2 2); Macon Co., Nantahala National Forest, Copper Ridge Bald, 5144 ft., 9-Jul-2019 (1 9, 1 3); Macon Co., Nantahala National Forest, Cowee Bald, 4839-494 2ft., 9-Jul-2019 (5 9, 5 4); Macon Co., Hwy. 64, nr. Dry Falls, 16-May-1986 (1 3); Madison Co., Pisgah National Forest, Camp Creek Bald, 4741 ft., 1-Mar-2022 (1 4); McDowell, Pisgah National Forest, Snooks Nose Trail, 2219 ft., 25-Aug-2015 (1 9, 2 3); Polk, Melrose Falls (lower), 1103 ft., 10-Aug-2021 (1 9, 3 ); Polk, Green River Game Lands, Lower Bradley Falls Tr., 1620 ft., 19-Mar-2018 (3 9, 1 3); Polk, Green River Game Lands, Green River Cove Tr., 1070 ft., 18-Mar-2018 (1 Q); Polk, Green River Game Lands, 1740, 18-Mar- 2018 Polk, Green River Game Lands, 1740 ft., 18-Mar-2018 (1 9, 1 4); Buncombe Co. Co., Blue Ridge Parkway National Park, Bull Gap, 3100 ft., 1-May-1990 (1 3); Swain Co., Great Smoky Mountains National Park, Clingmans Dome, 6264—6500 ft., 4-Jun-2018 & 14-Sep-2021 (4 4); Swain Co., Great Smoky Mountains National Park, Lakeshore Tr., Payne Ck., 1816 ft., 12-Apr-2022 (1 9, 2 3); Transylvania Co., Pisgah National Forest, Hwy 215, 1 mi. S. Blue Ridge Pkwy, 5122 ft., 7-May-2018 (1 2); Yancey Co., Pisgah National Forest, Woody Ridge Tr., 5086-5301 ft., 15-Jun-2020 & 19-Oct-2021 (1 9, 2 4); SC: Greenville Co., Chestnut Ridge Heritage Preserve, 1090 ft., 8-Apr-2018 (1 2); Pickens Co., Eastatoe Creek Heritage Preserve, 1421 ft., 30-Apr-2015 (1 3); Pickens Co., Sassafras Mountain summit, 3347 ft., 10-Jun-2015 (1 4); Oconee Co., Sumter National Forest, Ellicott Rock Wilderness, 2113-2679 ft., 3-Jun-2015 & 4-May-2015 (5 9, 4 3); Oconee Co., Sumter National Forest, Ri- ley Moore Falls, 900 ft., 3-Mar-2018 (2 2); Oconee Co., Coon Branch Nat. Area, 1950 ft., 28-Feb-2016 (3 9, 1 @); TN: Unicoi, Cherokee Co. National Forest, Big Bald, 5346-5430 ft., 5-Aug-2020 & 21-May-2021 (5 2); Blount Co., Great Smoky Mountains National Park, Whiteoak Sink, 1724 ft., 27-Oct-2021 (1 3); Sevier Co., Great Smoky Mountains National Park, Alum Cave Bluff Trail, 5196 ft., 25-Jun-2019 (3 9, 1 4); Sevier Co., Great Smoky Mountains National Park, Appalachian Trail nr. Newfound Gap, 5456 ft., 4-Jun-2018 (4 2, 1 &); Sevier Co., Great Smoky Mountains National Park, Off Hwy 441, 4575 ft., 12-Mar-2020 (2 2). LARVAE: NC: Swain Co., Great Smoky Mountains National Park, Clingmans Dome, 14-Sep-2021; Hay- wood Co., Great Smoky Mountains National Park, Big Cataloochee Mt., 5-Nov-2020; Haywood Co., Great Smoky Mountains National Park, Balsam Mt. Trail, 5-Nov-2020; Jackson Co., Blue Ridge Parkway National Park, Browning Knob, 22-Sep-2020; TN: Sevier Co., Great Smoky Mountains National Park, Alum Cave Bluff Tr., 28-Sep-2021; GA: Rabun Co., Chattahoochee NE, Rabun Cliffs, 25-Nov-2019. Description. Males winged, females lacking fully developed flight wings; large, elon- gate, generally dark but elytra reddish at humeri and often along elytral suture (Fig. 1C), female generally darker (Fig. 1D); male eyes large, protuberant, with ~ 30 large om- matidia (Fig. 1A, E); female eyes smaller, flush with side of head, comprising ~ 12 om- matidia (Fig. 1B, F); scape and pedicel similar in length, ca. as long as width of eye, 142 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Figure |. Habitus and character photos of Euconnus megalops A male, lateral view B female, lateral view C male, dorsal view D female, dorsal view (BgBld.B.349) E SEM of male head, lateral F SEM of female head, lateral. antennomeres 3—6 uniformly shorter, ca. as long as wide, male antennomere VII weakly asymmetrical, slightly narrowed anterad, male antennomeres VIII-XI enlarged and elon- gate (longer than wide) but none carinate on anterior edges (Fig. 1E); female with anten- nal club tetramerous, antennomeres shorter than width (Fig. 1F). Frons and vertex with long, erect setae, each ca. as long as scape; vertex narrowed to a broadly rounded point; neck ca. one-half maximum head width; frons shallowly depressed between antennal bases; epistoma deeply depressed below antennae; labrum with anterior margin evenly rounded, with comb of short setae at middle. Pronotum densely setose at sides and anteriorly, more sparsely posterad, ‘bell-shaped’, widest just beyond middle, narrowed anterad and posterad, posterior margin slightly widening; pronotum depressed along Appalachian Euconnus (Cladoconnus) 143 A B Cc D Figure 2. Aedeagus A Euconnus megalops B Euconnus vexillus C Euconnus cumberlandus D Euconnus vetustus. posterior margin with very short median longitudinal carina and stronger lateral longitu- dinal carinae, a distinct fovea on either side between these carinae; a weak sublateral ca- rina merging with lateral one at posterior pronotal corner, a distinct fovea between them (apparently the anterior-most of the two shown for E. (C.) motschulskii in Jatoszynski [2018]), and an additional shallow fovea below, on the side; prosternum rather short in front of procoxae, lacking median carina, with dense fringe of setae along anterior mar- gin; elytron sparsely covered with fine setae, with pair of deep foveae at base, longitudi- nally weakly depressed behind these; elytral apices rounded; mesoscutellar shield hidden; mesoventrite with strong carina, setose along its crest, separating mesocoxae, extending anterad to separate apices of procoxae, posteriorly merging with metaventrite immedi- ately behind mesocoxae; mesepimera densely setose, produced laterad mesocoxa; meta- coxae narrowly separated by metaventral process; abdominal ventrites unmodified; legs generally slender, setose; protibia with only weakly expanded adhesive setae along inner margin of apical third. Aedeagus (Fig. 2A) with median lobe broadly truncate, parameres thin, rather short, straight, tapered to apex, each with three or four setae extending to or just beyond apex of median lobe; compressor plate asymmetrical, with apex obliquely truncate to weakly lobed off center; endophallus with asymmetrical armature; upper armature comprising two long curved sclerites, one longer, nearly reaching apex of me- dian lobe at rest, abruptly curved near apex with thin inner blade bridging the apex, its inner edge concave, the other upper sclerite curving opposite, thinner, bluntly rounded at apex; lower armature comprising three separate processes: one basal, short, strongly curved, and with a bifid or trifid apex; one longer, medial, broadly hooked; the third lateral and more slender, elongate, apex acute and variously straight or bending mediad. Distribution. This species is the most abundant and widespread of the Appala- chian Cladoconnus species, occurring from Brasstown Bald in the southwest, northeast to Celo Knob in the Black Mts. It also exhibits the widest elevational range of the species, from ca. 900 ft in upstate South Carolina, all the way to the top of Clingmans Dome at 6500 ft. 144 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) tx E. megalops Figure 3. Map of collecting records for Euconnus (Cladoconnus) megalops. Remarks. This species’ morphology is relatively invariant across its broad range. The upper sclerites of the endophallic armature vary slightly in apical curvature and shape, but without obvious geographic trends. Similarly, none of these variants cor- respond to a geographically dispersed, divergent genetic subclade within the broader species, a peculiar result that merits further investigation. This species name refers to the conspicuously enlarged eyes of the males. Euconnus vexillus sp. nov. https://zoobank.org/47 ED 1 80B-4423-46AC-8CA4-3BB1FEDCDB19 Figs 2B, 4, 5 Type material. Holotype &, deposited in FMNH: “USA:SC: Greenville Co. 35.1523°N, 82.2814°W, Chestnut Ridge Heritage Preserve, vi.05.2015, S. Myers, Hardwood lit- ter” / “[QR code] CLEMSON ENT CUAC000026944”. Paratypes (10, CUAC, FMNH) — 2 6, 2 9: same data as type; 1 @, 1 9: “USA:SC: Greenville Co. 35.1518°N, 82.2839°W, Chestnut Ridge Heritage Preserve, vi.05.2015, S. Myers, Hardwood Appalachian Euconnus (Cladoconnus) 145 litter”; 1 9: “USA:SC: Greenville Co. 35.1501°N, 82.2820°W, Chestnut Ridge Herit- age Preserve, vi.05.2015, S. Myers, Hardwood litter”; 2 9: “USA:SC: Greenville Co. 35.1406°N, 82.2790°W, Chestnut Ridge Heritage Preserve, vi.05.2015, S. Myers, Hardwood litter”; 1 9: “USA:SC: Greenville Co. 35.1506°N, 82.2799°W Chestnut Ridge Heritage Preserve, iv.08.2018, M. Caterino & L. Vasquez-Vélez, sifted litter”. Other material. (26) WV: Mercer Co., Camp Creek State Forest, 23-Jul-1971, leaf litter (4 4, 11 Q); ‘Black Mts’, x.1901 (3 3); NC: Caldwell Co., Grandfather Mt. State Park, Nuwati Trail, 4020 ft., 17-May-2021 (1 4); McDowell Co., Pisgah Na- tional Forest, Mackey Mountain Trail, 3433 ft., 25-Aug-2015 (2 3); McDowell Co., Pisgah National Forest, Snooks Nose Trail, 1998 ft., 25-Aug-2015 (3 9, 1 3); Polk Co., Green River Game Lands, 1740 ft., 18-Mar-2018 (1 <). Diagnostic description. This species is very similar to the preceding species and can best be distinguished by male genitalic and antennal characters. Like the preced- ing, males are winged, while females appear not to be. A few noteworthy external dif- ferences can also be cited: males with distinct carinae on antennomeres VIII and IX (Fig. 4E); female antennae less distinctly tetramerous, with antennomere VIII interme- diate in size between antennomeres VII and IX (Fig. 4D); body darker, stout, with only faintly rufescent elytral humeri, most distinctly lighter along posterior half of elytral suture; male eyes smaller, less protuberant, with only ~ 25 ommatidia; median basal carina of pronotum slightly better developed to base (Fig. 4F); aedeagus (Fig. 2B) with median lobe broadly truncate at apex; parameres short, tapered, bearing three apical setae; compressor plate strongly asymmetrical, rather short; endophallic armature with strong pair of upper processes: the left long and strongly hooked apically, with a thin inner laminar blade, its inner edge concave; the right shorter, more strongly curved toward middle of longer process; lower endophallic armature consisting of three hook- like processes of varying lengths, the lateral-most rather short and strongly curved, the medial-most of intermediate length, the one between them the longest, its tip just vis- ible between bases of upper processes (in ‘dorsal’ view). Distribution. This species has been found at several widely scattered localities east and northeast of the Asheville Depression, from the headwaters of the French Broad near the southern end of its range, at Chestnut Ridge, South Carolina, to southern West Virginia in the northeast. Most of its occurrences are at middle elevations, from 1090 to 4020 ft, most below 2000. Remarks. ‘This species is similar and, judging by male genitalia, closely related to E. megalops. Both have similar opposing hooked upper endophallic sclerites, though the shorter right one (Fig. 2B) of E. vexillus is distinctive. Moreover, the lower trio of endophallic sclerites are quite distinct in £. vexillus, with three hooks projecting distad to varying lengths. In males, the carinae of antennomeres VII and IX will immediately distinguish this species from FE. megalops, and the only weakly tetramerous club of females can be used to distinguish most of those, although they are similar to darker females of E. adversus. This species name means ‘standard (or flag)-bearer’, referring to its possession of Cladoconnus-typical carinae, borne proudly on its antennae. 146 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Figure 4. Habitus and character photos of Euconnus vexillus A male, dorsal view B female, dorsal view C SEM of mouthparts, ventral view D SEM of female head, lateral E SEM of male antenna, lateral F SEM of elytra and pronotum G SEM of meta- and mesoventrites. Appalachian Euconnus (Cladoconnus) 147 O E. adversus vx E. vetustus O E. vexillus Figure 5. Map of collecting records for Euconnus (Cladoconnus) vexillus (squares), E. (C.) vetustus (stars), and £. (C.) adversus (circles). Euconnus cumberlandus sp. nov. https://zoobank.org/8E94B9B6-C01A-4460-AC19-4E34FFFE452D Fics 2G, -67 Type material. Holotype 3, deposited in FMNH: “Fall Creek Falls S.Pk., Van Buren Co., TENN, 13.X.1962” / “Rhododendron duff, H.R.Steeves, leg.” / “Caterino DNA Voucher Extraction MSC12284”. Paratypes (9, FANH, CUAC, CMNC);: 2 3,29: same data as type; 3 9, 2 4: “Fall Creek Falls $.Pk., Fall Creek Falls Pit, Bledsoe Co., TENN., 14.X.1961” / “Rhododendron duff, H.R.Steeves, leg.”. Other material. (12, CMNC, CUAC, FMNH) TN: Grundy Co., Savage Gulf State Natural Area, 1150ft., 4-May-2021, Litter — bottom of canyon (2 9); Pickett Co., Jamestown, Jordan Motel, 16-Jun-1962, forest floor near falls (1 2); GA: Dade 148 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Co., Cloudland Canyon State Park, 16-May-1972, rhododendron litter (2 6, 5 ); Dade Co., Cloudland Canyon State Park, 7-Aug-1962, forest floor (1 Coe Diagnostic description. This species is very similar to the preceding species and can best be distinguished by male genitalic characters. Like the preceding, males are winged, while females appear not to be. The antennal carinae of male antennomeres VIII and IX are present but rather weakly developed (Fig. 6), with neither quite span- ning the entire length of the antennomere itself; female antennae less distinctly tetram- erous, with antennomere VIII intermediate in size between antennomeres VII and IX; body darker, stout, with only faintly rufescent elytral humeri, most distinctly lighter along posterior half of elytral suture; male eyes smaller, less protuberant, with only ~ 25 ommatidia; aedeagus (Fig. 2C) with median lobe broad, apex weakly rounded, slightly narrowed subapically; parameres short, tapered, bearing three apical setae; compres- sor plate asymmetrical, produced to subacute tip on right side; endophallic armature with strong pair of upper processes: the left long and strongly hooked apically; the right shorter, strongly bifid, its apices extending beneath left process; lower endophal- lic armature consisting of four hook-like processes of varying lengths, two longer and moderately to strongly hooked, two shorter ones borne on a single sclerite, apices directed distad. Distribution. This species is known from northwestern Georgia to north- central Tennessee. Two of the Tennessee localities, including the northernmost, are represented only by females, and they can be assigned only tentatively to this species. Figure 6. Antennal carina of E. cumberlandus. Appalachian Euconnus (Cladoconnus) 149 O E. cumberlandus vx E. astrus Figure 7. Map of collecting records for Euconnus (Cladoconnus) cumberlandus (circles) and E. (C.) astrus (stars). Remarks. This species is similar to and apparently closely related to E. megalops and FE. vexillus. The only sequenced specimen of this species is a female from Savage Gulf, TN, a locality not yet represented by males. Sequences from other localities, in- cluding the type locality would help confirm the species’ unity as circumscribed here. Euconnus vetustus sp. nov. https://zoobank.org/417B9BE8-BE69-4792-926B-3B4FA19CD3DD Figs 2D, 5, 8 Type material. Holotype 3, deposited in FMNH: “USA: NC: Mitchell Co., 36.1038°N, 82.0809°W, PisgahNE vi.8.2020 Grassy Ridge Bald, 6083’, M. Caterino, deciduous shrub litter” / “[QR code] CLEMSON-ENT CUAC000004025” / “Ca- terino DNA Voucher Extraction MSC4491, Morphosp. GrB.A.336”. Paratypes (4) —2 9,2 4: same data as type. Other material. (7) — NC: Graham Co., Nantahala National Forest, Huckleberry Knob, 5511 ft., 13-Oct-2020 (1 4); Haywood Co., Pisgah National Forest, Black Balsam Knob, 6072 ft., 7-May-2018 (1 4); Yancey Co., Pisgah National Forest, Dev- ils Gap, 3813 ft., 24-Aug-2015 (2 3); ‘Black Mts’ (1 2); TN: Unicoi Co., Cherokee 150 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) National Forest, Big Bald, 5430-5464 ft., 5-Aug-2020 & 21-May-2021 (1 9, 2 3); Sevier Co., Great Smoky Mountains National Park, Off Hwy 441, 4575 ft., 12-Mar- 2020 (1 9). Diagnostic description. This species is similar to the preceding species being dark in color and wing-dimorphic, and can best be distinguished by male genitalic charac- ters. External differences, however, include smaller body size, entirely dark coloration (Fig. 8A); head, especially crest of vertex, rounder; frons only weakly depressed be- tween antennal bases; male and female eyes do not differ appreciably in size, both hav- ing ~ 20 ommatidia; the protibia of both sexes is shorter, widened apically, and bears conspicuously modified setae on the apical half (Fig. 8C); male antennal club tetram- erous, with the club segments slightly longer than wide (Fig. 8A), no antennomeres bearing carinae; female antenna shorter, with club 4-segmented, club segments ca. as long as wide (Fig. 8B); aedeagus (Fig. 2D) with median lobe narrowed to narrowly rounded apex; rather straight parameres bear two apical setae extending just to apex of median lobe, these setae finely bent inward at apex; compressor plate symmetrical, narrow, reaching nearly to apex of median lobe; upper endophallic armature with one short curved hook, not extending beyond shoulders of tegmen, its apex blunt and dis- tinctly fringed (at higher magnification); immediately internal to it is a rather broadly rounded thin plate (the apical margin of which is strongly sclerotized, appearing as if maybe a second opposing hook); lower endophallic armature with a long bifid process extending from deep in the basal bulb to beyond its shoulders, the inner blade thin, curving weakly upward, the outer blade longer, reaching to near apex of compressor plate, its apex finely fimbriate. Distribution. This species occurs over a relatively broad but disparate range, span- ning the Asheville Depression, from the Unicois (Huckleberry Knob) in the southwest to the Roan Highlands and Big Bald in the northeast. So far it has not been found in the Black Mts or on Grandfather Mountain. Known occurrences are all at higher eleva- tions, from 4575 to 6100 ft. Remarks. Individuals from across the broad range of this species show consider- able genetic variation, and the male genitalia do show slight variation. Northern ex- amples (Big Bald, Roan Highlands) exhibit a more strongly hooked upper endophallic process. A few specimens, from scattered localities (e.g., Smokies, Balsam Mts., and Roan Highlands), have the bi-arcuate mandibles described for European Cladoconnus. But this character varies as well; specimens from Big Bald, NC, have mandibles that are less distinctly arcuate, as well as longer and more slender. Male genitalia have been re- examined with this in mind, but no corresponding differences emerge. Further genetic work and longer series of males may justify separating some of these. Females from Devil’s Gap, NC are light in color (perhaps teneral) but match in other characters, including DNA. The name of this species comes from the Latin for ‘old’, referring to the possibility the species has inhabited the area for a long time, as judging by its broad distribution, only distant relation to the rest of the species described here, and deep genetic diver- gences among populations. Appalachian Euconnus (Cladoconnus) 151 Figure 8. Habitus and character photos of Euconnus vetustus A male habitus, dorsal (BBK.A.020) B male habitus, lateral (GRB.A.336) C SEM of protibial setae. Euconnus falcatus sp. nov. https://zoobank.org/868CC7AB-B90A-4337-AEOE-F8329719CAAC Figs 9, 10A, B, 11 Type material. Holotype 3, deposited in FMNH: “USA: NC: Swain Co., 35.5566°N, 83.4966°W, SmokyMts NP, 6272’, Clingmans Dome, sifted litter, ix.14.2021, M.Caterino & E.Recuero” / “[QR code] CLEMSON-ENT CUAC000156725” / “Caterino DNA Voucher Extraction MSC7899, Morphosp. CD.B.307”. Paratypes (40, CUAC, CNCI, FMNH, UNHC) —- 1 @: “USA: NC: Swain Co., 35.5613°N, 83.5006°W, SmokyMts NP, 6364’, ClingmansDome, W. slope, sifted litter, vi.5.2018, M.Caterino” / “[QR code] CLEMSON-ENT CUAC000079152” / “Caterino DNA Voucher Extraction MSC2834 Morphosp. CD.A.021”; 1 9, 3 4: “USA: NC: Swain Co., 35.5824°N, 83.3979°W, SmokyMtsNPoffHwy 441, 4610’, iii.12.2020, M.Caterino & EEtzler, sifted litter’; 1 2: “USA: TN: Swain Co., 35,.6237°N, 83.4163°W, SmokyMtsNPoftHwy 441, 4575’, iii.12.2020, M.Caterino & FEtzler, sifted litter” / “[QR code] CLEMSON-ENT CUAC000110837” / “Caterino DNA Voucher Extraction MSC4193 Morphosp. Hwy.A.006”; 1 2, 8 G: “USA: TN: Sevier Co., 35.6160°N, 83.4149°W, SmokyMts NP, 5456’, App.Tr. E of Newfound Gap, sifted litter, vi.5.2018, M.Caterino”; 2 9, 9 @: “USA: TN: Sevier Co., 35.6308°N, 83.3904°W, SmokyMts NP, 6190’, Mt. Kephart, vi.5.2018, M.Caterino”; 1 S: “USA: TN: Sevier Co., 35.6311°N, 83.3895°W, SmokyMts NP, 6183’, Mt. Kephart,sifted 152 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) litter, vi.5.2018, M.Caterino” / “[QR code] CLEMSON-ENT CUAC000079110” / “Caterino DNA Voucher Extraction MSC2790 Morphosp. MK.A.009”; 1 3: “USA: TN: Sevier Co., 35.6311°N, 83. 3893°W, SmokyMts NP, 6187’, Mt. Kephart, sift- ed litter, ix.14.2021, M.Caterino & E.Recuero” / “[QR code] CLEMSON-ENT CUAC000156767” / “Caterino DNA Voucher Extraction MSC7941 Morphosp. MK.B.482”; 4 9, 4 3: “USA: TN: Sevier Co., 35.6425°N, 83.4427°W, SmokyMts NP, 5196’, Alum Cave Tr., vi.25.2019, M.Caterino & M.Ferro, sifted mixed litter”; 1 : “USA: TN: Sevier Co., 35.6529°N, 83.4378°W, SmokyMts NP, 6467’, Mt LeCon- te, vi.25.2019 M.Caterino & M.Ferro, sifted conifer litter” / “[QR code] CLEMSON- ENT CUAC000079302” / “Caterino DNA Voucher Extraction MSC3483 Morphosp. MLc.009”; 1 9, 1 3: “USA: TN: Sevier Co., 35.6541°N, 83.4364°W, SmokyMts NP 6588’, Mt LeConte, ix.28.2021, M.Caterino, sifted litter’; 1 4: “NC Gr.Sm.Mts.N.P, Clingmans Dome, 1950—2020m 2.VI.86 A.Smetana” (CMNC); 1 ¢: “TENN:Great Smoky Mts.St.Pk.: Newfound Gap” / “X.24.1969 W.Shear + E.Coyle leg.” / “FM(HD) 69-62 Spruce litter” (FMNH). Other material. (36, CUAC, UNHC, CNCI) — NC: Haywood Co., Pisgah Na- tional Forest, Black Balsam Knob, 6033-6072ft, 7-May-2018 & 20-Oct-2020 (3 @); Haywood Co., Pisgah National Forest, Mountains to Sea Trail, 5540ft., 8-Sep-2020 (1 3); Haywood Co., Blue Ridge Parkway National Park, Mt. Pisgah, 5420ft., 13-Sep- 2022 (1 9, 1 4); Jackson Co., Blue Ridge Parkway National Park, Mt. Lyn Lowry, 6097 ft., 15-Jun-2021, sifted litter (1 9); Jackson Co., Blue Ridge Parkway National Park, Mt. Lyn Lowry, 6205 ft., 22-Sep-2020, sifted litter (1 9); Jackson Co., Blue Ridge Parkway, National Park, along Blue Ridge Pkwy, 5572ft., 11-Sep-2019 (1 9, 1 4); North Carolina, Jackson Co., Blue Ridge Parkway National Park, Browning Knob, 6140-6221, 22-Sep-2020 & 29-May-2018 (3 9, 1 3); Jackson Co., Blue Ridge Parkway National Park, Waterrock Knob, 6059, 29-May-2018 (4 9, 4 3); Jack- son Co., Blue Ridge Parkway National Park, Waterrock Knob, 6281, 29-May-2018; Jackson Co., Balsam Mountain Preserve, Doubletop Mountain, 5396-5480, 7-Feb- 2015 & 15-Jun-2015 (4 4); Jackson Co., Nantahala National Forest, Toxaway Mt., 4750, 5-Aug-2020 (1 &); Jackson Co., Wolf Mt. overlook, 26-May-1986 (3 3); Jack- son Co., Blue Ridge Parkway National Park, 5572, 11-Sep-2019 (1 2); Macon Co., Nantahala National Forest, Copper Ridge Bald, 5144, 9-Jul-2019 (1 @); Macon Co., Hwy. 64, nr. Dry Falls, 16-May-1986 (2 9, 1 4); Macon Co., Coweeta Hydrological Lab, Shope Fork, 3200, 28-May-1983 (1 3). Description. Body rufescent (Fig. 9A-D), weakly translucent; head and elytra sparsely setose, pronotum densely setose, especially at sides; head with frons weakly depressed between antennal bases (Fig. 9H), vertex convex, neck just over one-half maximum head width; eyes of male moderately large, protuberant, comprising ca. 15 distinct ommatidia (Fig. 9H), situated immediately behind antennal insertions, width slightly more than that of tempora behind; eyes of female smaller (Fig. 9G), less prominent, comprising < 8 ommatidia, width only one-third that of tempora; antennae inserted under blunt frontal shelf; antennae of male with scape and pedicel Appalachian Euconnus (Cladoconnus) hs Figure 9. Habitus and character photos of Euconnus falcatus A male, lateral view B female, lateral view C male, dorsal view D female, dorsal view E SEM, male antenna F SEM, pronotum and elytra G SEM, female head, lateral view H SEM, male head, anterior view. 154 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) similar in length, ca. twice length of antennomeres III-VI, antennomere VII slightly larger, asymmetrical, slightly produced on anterior margin, antennomeres VIII-XI dis- tinctly larger, antennomeres VIII and IX with longitudinal carinae on anterior margin (Fig. 9E), that of VIII more or less parallel to antennal axis, that of IX more strongly produced at apex, antennomeres VIII-X similar in length, terminal antennomere ca. twice as long; female antennae shorter, club composed of only three short, wide an- tennomeres (Fig. 9D, G); pronotum weakly bell-shaped, with sides arcuate (Fig. 9F), widest ca. one-fourth behind straight anterior margin, narrowed posteriorly, similar in width to anterior, with short longitudinal carinae on either side, transversely de- pressed between, with small submedian foveae on either side of low median ridge (median basal carina absent), and two small foveae outside lateral carina, pronotal sides very densely bristled; mesoscutellar shield hidden; elytra broadly rounded, wid- est just anterad middle, each with two small basal foveae, sparse setae randomly ar- ranged; flight wings poorly developed or absent in both sexes; abdominal ventrites without secondary sexual differences; aedeagus (Fig. 10A) broadly rounded basally, apex of median lobe narrowed, pinched before apex, subtruncate; parameres thin, close to median lobe, with two long apical setae extending beyond median lobe apex; dorsal diaphragm crescent-shaped; compressor plate thin, narrowly rounded at apex, slightly asymmetrical; endophallus with two asymmetrical primary sclerites, an upper one almost linear, weakly sinuous to acute apex, the lower one strongly sickle-shaped, bearing a single conspicuous and sometimes an additional small secondary processes on its inner surface. Distribution. Euconnus falcatus exhibits a wide distribution, especially consider- ing its flightlessness, including the Great Smoky Mountains, continuing along the high Blue Ridge Parkway corridor into the Plott Balsams and Great Balsams, as well as slightly lower parts of the Nantahala and Cowee Mountains. In elevational range it occurs from 3200 ft at Coweeta to its highest occurrences on Clingmans Dome and Mt. LeConte above 6500 ft. Remarks. A more complete description of this species is provided to serve as a general description for several subsequent species that differ in few or no obvious ex- ternal characters, including E. cataloochee, E. kilmeri, E. draco, E. tusquitee, E. attritus, E. cultellus, E. adversus, and E. astrus. The modified male antennomeres differ signifi- cantly in a couple of these (E. adversus and E. astrus), but only slightly or not at all in the others. Euconnus falcatus can only be unambiguously distinguished from the others by male genitalic characters. Specifically, the lower endophallic sclerite is long, sickle- shaped, and bears a small secondary hook near its midpoint. The apex of the median lobe is more narrowly knobbed than most others, and the apex of the compressor plate is only subtly asymmetrical. There appear to be some differences in exact shapes of endophallic sclerites among localities (e.g., Fig. 10B). Some of this variation may be real, but some also appears to result from varying degree of extroversion of the lower hooked sclerite relative to the upper one. This species name refers to the distinctively ‘sickle-shaped’ hook of the endophallic armature. Appalachian Euconnus (Cladoconnus) 155 Figure 10. Aedeagus. A Euconnus falcatus B compressor plate and upper endophallic armature of Eucon- nus falcatus (Plott Balsams variant) C Euconnus cataloochee D Euconnus kilmeri. Euconnus cataloochee sp. nov. https://zoobank.org/FD84FEAD-72A4-4FBD-B0OD6-B3A76AA6B697 Figs 10C, 11 Type material. Holotype 3, deposited in FMNH: “USA: NC: Haywood Co., 35.6721°N, 83.1760°W, Smoky Mts NP, 6150’, Big Cataloochee Mt., xi.5.2020, sift- ed litter, M.Caterino & EEtzler” / “[QR code] CLEMSON-ENT CUAC000135173” / “Caterino DNA Voucher Extraction MSC6485 Morphosp. BCat.B.315”. Paratypes (32 - CUAC, FMNH, CNCI, UNHC):7 &, 7 6: same data as type; 5 9,5 6: “USA: NC: Haywood Co., 35.6453°N, 83.2025°W, SmokyMtsNP, Balsam Mt.Tr., 5086’, xi.5.2020, M.Caterino & EEtzler, Sifted litter’; 1 9, 1 @: USA: NC: Haywood Co., 35.6686°N, 83.1749°W, Smoky Mts NP, 5725’, Big Cataloochee Mt., vii.14.2020, sifted litter, M.Caterino, EEtzler’; 1 6: “USA: NC: Haywood Co., 35.6724°N, 83.1761°W, Smoky Mts NP, 6130’, Big Cataloochee Mt., vii.14.2020, sifted litter, M.Caterino, FEtzler”; 3 2, 2 @: “USA: NC: Haywood Co., 35.6675°N, 83.1805°W, Smoky Mts NP, 5586’, MtSterlingIr@Lost BottomCk, vii.14.2020 M.Caterino& EEtzler, sifted litter”. Diagnostic description. This species exhibits no obvious external differences from the preceding species and can only be distinguished by male genitalic characters. An- tennomere VIII of male slightly more strongly produced at inner basal corner, slightly oblique to antennal axis; aedeagus (Fig. 10C) with median lobe slightly narrowed, pinched near apex, bluntly rounded; parameres curving slightly inward at tips, each with three apical setae; compressor plate weakly asymmetrical and narrowly rounded; endophallus with two asymmetrical primary sclerites, one long, sinuate, apically bifid, with lateral hook curving beneath a median apically directed spear; lower endophallic sclerite originating nearer base, broad at base, subdivided into two widely spaced apical hooks, curving the same direction as inner hook of upper sclerite. 156 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Distribution. This is a very restricted species, only known from a couple localities on or very near Big Cataloochee Mountain in the Smokies, and is only known above 5100 ft. Remarks. The bifid apex of the upper endophallic sclerite is similar to that of E. cultellus, but the subapical (lower) tip is more strongly developed, almost appear- ing as a separate sclerite. The two well-developed hooks on a shorter overall lower endophallic sclerite distinguish it from £. falcatus. Sequence differences from the latter are minimal to non-existent, suggesting either very recent ancestry or introgression, as they do seem to be sympatric in the central Great Smoky Mountains, the only locality known for £. cataloochee. This species is named for its type locality, Big Cataloochee Mt., as it is known from only a small area near its summit. The name apparently comes from a Cher- okee word referring to the prominent wooded ridges in this region of the Great Smoky Mountains. Euconnus kilmeri sp. nov. https://zoobank.org/435F225A-A9C8-4F63-AEE1-262A9852E969 Figs 10D, 11 Type material. Holotype 3, deposited in FMNH: “USA:NC: Graham Co., 35.3433°N, 83.96207°W, Joyce Kilmer, VII.20.2015 S. Myers, Sifted litter” / “[QR code] CLEMSON-ENT CUAC000011399” / “Caterino DNA Voucher Extraction MSC11827”; Paratypes (0). Diagnostic description. This species exhibits no obvious external differences from the preceding two species, and can best be distinguished by male genitalic characters; antennal carinae slightly weaker than preceding; aedeagus (Fig. 10D) with median lobe rather short, narrowed to bluntly rounded apex, only very weakly knobbed; parameres with apices obliquely truncate, bearing three long, curved setae; compressor plate strongly asymmetrical, apex narrowly knobbed and displaced to one side; endophallus with dominant upper sclerite sickle-shaped, with long straight inner edge, outer edge strongly curved, broadest just beyond middle, bearing a secondary inner tooth directed mediad and inward; lower endophallic sclerite shorter, bearing two widely separated distally pointing spikes, and an obliquely directed subapical spike on one side. Distribution. Euconnus kilmeri is only known from a single locality within the Joyce Kilmer Memorial Forest in far western North Carolina. This site sits at an eleva- tion of 2800ft. Remarks. ‘The aedeagus of £. kilmeri is quite distinct in the shape of the strongly asymmetrical compressor plate, the long, straight inner edge of the upper endophallic sclerite, and in the trifid lower sclerite, with two apically pointing spines similar in length. This species is named to honor the American poet Joyce Kilmer “I think that I shall never see, a poem as lovely as a tree...” for whom the type locality stands as a proper monument to his appreciation for nature. Appalachian Euconnus (Cladoconnus) 157 E. tusquitee F. cataloochee E. falcatus F. draco FE. kilmeri x E. atiritus Figure I 1. Map of collecting records for Euconnus (Cladoconnus) falcatus complex. E. (C.) falcatus (circles), E. (C.) cataloochee (stars), E. (C.) tusquitee (triangules), E. (C.) draco (hexagons), E. (C.) kilmeri (plus sign), and E. (C.) attritus (x). A 5 O O + Euconnus draco sp. nov. https://zoobank.org/4C7 107D3-59AB-4C24-BBFF-A5GEDD2568AB Figs 11 12a Type material. Holotype 3, deposited in FMNH: “USA:NC: Graham Co., 35.3216°N, 83.9929°W, NantahalaNkK v.4.2020 5522’, Huckleberry Knob, M.Caterino & FEtzler, sifted spruce litter” / “[QR code] CLEMSON-ENT CUAC000135352” / “Caterino DNA Voucher Extraction MSC6877 Morphosp.HKnb.B.329”. Paratypes (6, CUAC, FMNH)-—3 3: same dataas type; 1 2: “USA:NC: Graham Co., 35.3210°N,83.9934°W, NantahalaNE, v.4.2020, 5491’, Huckleberry Knob, M.Caterino&EEtzler, sifted de- ciduous litter” / “[QR code] CLEMSON-ENT CUAC000003924” / “Caterino DNA Voucher Extraction MSC4308 Morphosp.HKnb.A.048”; 1 2, 1 ¢: “USA:NC: Gra- ham Co., 35.3171°N,83.9833°W, NantahalaNE v.4.2020, 4702’, Cherohala Skyway, M.Caterino & EEtzler, sifted deciduous litter” Diagnostic description. This species exhibits no obvious external differences from the preceding three, and can best be distinguished by male genitalic characters; its male antennomere IX has the apical corner quite dentate; aedeagus (Fig. 12A) with apex 158 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) of median lobe narrowly knobbed and bluntly rounded; parameres weakly curving inward to apices, each bearing three long apical setae; compressor plate weakly asym- metrical, broadly rounded, nearly reaching to apex of median lobe; endophallus with two asymmetrical primary sclerites, upper sclerite long and narrowly sickle-shaped., its apex straight and acute; lower sclerite with three inwardly directed hooks, the apical- most long, thin, curved, median hook thicker, more strongly bent inward, basal-most thick, short, close to median. Distribution. Euconnus draco is also basically a single site endemic, known only from the vicinity of Huckleberry Knob, at 5500ft in the Unicoi Mts. The second, Cherohala Skyway locality is only 1km away, and just slightly lower at 4700ft. Remarks. The long, trifid lower endophallic sclerite of E. draco, with the apical hook long and thin, best distinguishes this species. The shape of the median lobe, com- pressor plate, and upper endophallic sclerite are otherwise quite similar to E. falcatus. This name of this species derives from the Latin ‘dragon’, as its type locality is near the popular motoring route “Tail of the Dragor’. Euconnus tusquitee sp. nov. https://zoobank.org/7F5099B6-92EE-4C41-B3FE-7C6203F5554D Figs 11, 12B Type material. Holotype 3, deposited in FMNH: “USA:NC: Clay Co., 35.1419°N, 83.7269°W, NantahalaNF, Tusquitee Bald, vii.6.2021, 5270’, Quercus/ Tsuga lit- ter M.Caterino & E.Recuero” / “[QR code] CLEMSON-ENT CUAC000171987” / “Caterino DNA Voucher Extraction MSC11727 Morphosp.TsqBld_171987”. Paratypes (1) — DNA only, voucher lost in extraction: same data as type, DNA Extract 7816, MorphospeciesTsqB.A.223. Diagnostic description. This species exhibits no obvious external differences from the preceding species, and can best be distinguished by male genitalic characters; carina of male antennomere VIII slightly oblique, ~ 20 degrees off the long axis of the anten- na; aedeagus (Fig. 12B) with apex of median lobe narrowed to near truncate apex, not knobbed; parameres weakly curved, bearing three apical setae; compressor plate short, strongly asymmetrical, sinuate with blunt median and acute lateral lobes; endophal- lus with dominant upper asymmetrical sclerite long, broad to middle, with narrow, curving apical portion nearly reaching apex of median lobe, bearing an inner, medially directed secondary tooth just beyond middle; lower endophallic sclerite shorter, reach- ing just to middle of upper, curving strongly opposite. Distribution. Another single-site endemic, Euconnus tusquitee is only known from Tusquitee Bald in the western Nantahala Mts, at an elevation of 5270 ft. Remarks. ‘The strongly asymmetrical and unevenly bilobed apex of the compressor plate is the best character for recognizing E. tusquitee. The strongly opposing hook on the lower endophallic sclerite is also distinctive. Its sickle-shaped upper endophallic sclerite is longer than, but otherwise similar to that of E. kilmeri. Appalachian Euconnus (Cladoconnus) 159 Figure 12. Aedeagus. A Euconnus draco B Euconnus tusquitee C Euconnus attritus. DNA of two specimens of this species was extracted. The first one was lost in the extraction process, though its DNA and images on the Flickr site (morphospecies code TsqB.A.223) remain. Unfortunately, neither extract sequenced well, so the placement of this species among the others of the £. falcatus complex remains uncertain. This species name refers to its type and only known locality, Tusquitee Bald. Euconnus attritus sp. nov. https://zoobank.org/37927727-6073-4A5B-8B 1 C-928 1D47B3699 Figs 11, 126715 Type material. Holotype &, deposited in FMNH: “USA:NC: Mitchell Co., 36.0931°N, 82.1453°W, Roan High Bluff, 6225’ viii.15.2018, M.Caterino, sifted Abies litter” / “LQR code] CLEMSON-ENT CUAC000003146” / “Caterino DNA Voucher Ex- traction MSC3360 Morphosp.RHB.A.012”; Paratypes (62, CUAC, FMNH, CNCI, UNHC) — 3 9, same data as type; 18 2, 16 gd: “USA:NC: Mitchell Co., 36.0933°N, 82.1447°W, Roan High Bluff, 6251’ viii.15.2018, M.Caterino, sifted Abies litter”; 1 Q@, 1G: “USA:NC: Mitchell Co., 36.0999°N, 82.1345°W, Roan High Bluff, 6140’, viii. 15.2018, M.Caterino, Rhododendron litter”; 2 2, 12 4: “USA: NC: Mitchell Co., 36.1041°N, 82.1223°W, PisgahNE, vi.8.2020, Roan High Knob, 6276’, M. Caterino, conifer litter’; 5 9, 4 6: “USA: NC: Mitchell Co., 36.1045°N, 82.1224°W, PisgahNE, vi.8.2020, Roan High Knob, 6286’, M. Caterino, conifer litter”. Other material. (52) — TN: Unicoi Co., Cherokee National Forest, Big Bald, 5237-5346ft, 5-Aug-2020 (1 2,5 4); NC: Avery Co., Grandfather Mt., 5240-537 0ft., 160 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) 21-Apr-2022 (4 9, 5 4); Caldwell Co. Grandfather Mt. State Park, Calloway Peak, 6-Oct-2020 & 17-May-2021, 5775—-5915ft. (12 9, 13 4); Caldwell Co. Grandfather Mt. State Park, Nuwati Tr., 4190ft. (1 @); Yancey Co., Pisgah National Forest, Woody Ridge Tr., 5086-5387ft., 15-Jun-2020 (5 9, 4 4); Yancey Co., Pisgah National Forest, Celo Knob, 6300ft., 19-Oct-2021 (1 3'); Madison Co., Pisgah National Forest, Camp Creek Bald, 4741 ft., 1-Mar-2022 (1 <4). Diagnostic description. This species is extremely similar in external morphology to many of the preceding, and is also best distinguished by male genitalic characters. However, it does exhibit a few unusual characters. The male antennomeres VIII-IX are slightly enlarged, but lack carinae on their inner/anterior edges (Figs. 13A, E); the female’s antennal club is shorter, and essentially trimerous (Fig. 13D), though anten- nomere VIII is slightly enlarged relative to VI; males and females flightless; median Figure 13. Habitus and character photos of Euconnus attritus A male, lateral view B male, dorsal view C female, lateral view D female, dorsal view E SEM of male antenna F SEM of venter of head. Appalachian Euconnus (Cladoconnus) 161 lobe of aedeagus (Fig. 12C) relatively long, nearly as long as basal bulb, narrowed to bluntly rounded, but not knobbed apex; shoulders of aedeagus sloped; parameres weakly curved, apices tapered, bearing three (rarely two) long straight setae; compres- sor plate symmetrical, parallel-sided to near apex, then narrowed to subacute apex, nearly as long as median lobe; endophallus with single long, upper sclerite, strongly sinuate, apically acute, bearing a sharp secondary process near its midpoint; the right side of the compressor plate sclerotized in a linear band, appearing as a second similar linear sclerite; actual second upper sclerite is much shorter, bent strongly at thickened midpoint, tapering to a thin, acute tip; lower endophallic sclerite a similar-length, trifid claw-like process on a sinuate stem originating near basal orifice. Distribution. This species is known only from northeast of the Asheville depres- sion, although there it is moderately widespread, occurring in all the major ranges: the Black Mts, Roan Highlands, Grandfather Mountain, Big Bald, and Camp Creek Bald. Its known localities span an elevation range of 3800-6300 ft. There is a CUAC speci- men labelled “Toxaway Mountain’ (in the more southerly Nantahala Mts.) that is al- most certainly mislabeled, as that site was visited the same day as the Big Bald locality. Remarks. Of the pale species of Cladoconnus, this species is only sympatric with E. adversus (it is also sympatric with the larger and darker FE. vetustus and E. vexillus). Males of E. adversus have very conspicuous antennal carinae, which are completely lacking in E£. attritus. Females, however, will be indistinguishable. Males of E. attritus have very distinctive endophallic sclerites, particularly the long, sinuate left upper sclerite, with its secondary median spike. The name of this species suggests that the subgenus-typical male antennal carinae are ‘worn away . Euconnus adversus sp. nov. https://zoobank.org/E8D7FF15-E9B6-48EE-9191-E1C4E7ABE41B Figs 11, 14, 15A, B Type material. Holotype 3, deposited in FMNH: “USA: NC: Yancey Co., 35.8524°N, 82.2485°W, PisgahNF, CeloKnob, x.19.2021, 6300’, M.Caterino, E.Recuero, & A.Haberski, sifted litter” / “[QR code] CLEMSON-ENT CUAC000157519” / “Ca- terino DNA Voucher Extraction MSC9225 Morphosp.CK.B.370”. Paratypes (4, CUAC) - 1 3: “USA: NC: Yancey Co., 35.8447°N, 82.2369°W, Pisgah NE, Woody Ridge Tr., vi.15.2020, 5244’, M.Caterino, F.Etzler, sifted litter” / “[QR code] CLEM- SON-ENT CUAC000137633” / “Caterino DNA Voucher Extraction MSC5528 Mor- phosp.WR.A.046”; 1 4: “USA: NC: Caldwell Co., 36.1184°N, 81.7909°W, Grandfa- ther Mt.SP, 4020’, Nuwati Tr., v.17.2021, A.Haberski, P. Wooden, sifted litter” / “[QR code] CLEMSON-ENT CUAC000135057” / “Caterino DNA Voucher Extraction MSC6364 Morphosp.NT.A.009”; 1 3: “USA: NC: Buncombe Co.Co. 35.7955°N, 82.3392°W, Big Butt Tr.,iii.19.2016, S.Myers, L.Vasquez-Velez, sifted litter” / “[QR code] CLEMSON-ENT CUAC00026846”; 1 3: “USA: TN: Unicoi Co., 35.9950°N, 162 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) 82.48972°W, Cherokee NF, Big Bald, v.21.2021, CW. Harden, A.Haberski, P Wooden, sifted litter” / “[QR code] CLEMSON-ENT CUAC000135395” / “Caterino DNA Voucher Extraction MSC6920 Morphosp.BgBld.A.026”. Other material. (19): NC: Jackson Co., Balsam Mountain Preserve, Sugarloaf Mountain 4491 ft, 7-Feb-2015, Sifting litter, oak litter in old stump depression; Ashe Co., Mt. Jefferson State Park, SE Reservoir, 4-Jul-1960; SC: Greenville Co., Chestnut Ridge Heritage Preserve, 1140 & 1220 ft., 5-June-2015; Pickens Co., Clemson Ex- perimental Forest, Seed Orchard Rd., 700ft, 12-Jul-2016; Oconee Co., Ellicott Rock Wilderness, East Fork Chattooga River, 2110ft, 4-May-2015; Oconee Co., Ellicott Rock Wilderness, Indian Camp Creek, 2822ft, 4-May-2015; Greenville Co., Mtn. Bridges Wilderness, 2230ft., 10-Mar-2018; GA: Rabun Co., NE Pine Mt., Chattooga R., 1800ft, 5-Jun-1981 (CUAC, CNCI, FMNH). Diagnostic description. This species is generally very similar to the preceding, and can best be distinguished by male genitalic characters; it and the following, how- ever, exhibit the most prominent antennal carinae among American Cladoconnus, those on antennomeres VIII and IX both being strong and oblique (Fig. 14C), that of VIII most produced at base and that of IX most produced at apex; antennomere VII also exhibits some expansion along its inner margin; males may be wing-poly- morphic, winged individuals appearing larger and darker in body color; female wings not observed; aedeagus (Figs. 15A, B) with median lobe evenly tapered and narrowly rounded at apex; parameres thin, each bearing three short terminal setae, the setae not reaching apex of median lobe; compressor plate short, asymmetrical, truncate on one side (‘left’ as drawn), produced on other; upper endophallic armature com- prising two dominant curved sclerites, one tapered to narrowly subacute apex, the shorter one much broader, more weakly curved inward to a bluntly truncate apex that meets the apex of the other; the lower endophallic armature consisting of a shorter, deeply bifurcate (or trifurcate) process, the apices slightly varied in curvature, gener- ally directed distad. Distribution. This species is widespread but rare, found at scattered lower eleva- tion sites across northwestern South Carolina, to higher elevations in the Nantahalas, Blacks, Grandfather Mountain, and Mount Jefferson in northernmost North Carolina. It has a broad elevational range as well, from just 700 ft up to the highest peaks in the region at 6500 ft. A single male labelled as from the Florida panhandle is almost certainly mislabeled. Collected by Stewart Peck on 8 June 1981, it was collected just 3 days after he collected another specimen of this species in Rabun County, Georgia. The ‘Florida’ specimen probably belongs to the Georgia series. Remarks. ‘The strongly modified antennomeres of male £. adversus will distinguish them immediately from anything sympatric (though not the more western E. astrus, below). There is considerable variation site-to-site in the detailed shapes of the upper and lower endophallic armature. In males from the Balsam Mt. Preserve (NC), the innermost endophallic sclerite is deeply trifucate, whereas in those from the Chest- nut Ridge Heritage Preserve (SC) the right tip of the lower endophallic sclerite is curved inward (compare Fig. 15B and Fig. 15A, respectively). None of these southern Appalachian Euconnus (Cladoconnus) 163 Figure 14. Habitus and character photos of Euconnus adversus A male, lateral view B male, dorsal view (NT.A.009) C SEM of male antenna. localities, however, are represented by sequence data, so before recognizing these vari- ants taxonomically, better representation for molecular data would be advisable. The name of this species refers to the seemingly ‘opposable’ carinae of male anten- nomeres VIII and IX. Euconnus astrus sp. nov. https://zoobank.org/ DO2BE2CE-A8EE-44C8-995F-5C69F95FB1 17 Figs 7, 15C Type material. Holotype ¢, deposited in CMNC: “ALA., Jackson Co., 5mi.N.W.Princeton, 19.V.1972, S.Peck. Ber.240” / “Caterino DNA Voucher Ex- traction MSC12286 Cladoconnus(AL)”; Paratypes (3): 1 3, same data as type; (FMNH); 1 3, 1 Q: “nr. Jess Elliot Cave, Jackson Co., ALA. 8.IV.1961” / “Hollow Tree, V.D.Patrick, H.R.Steeves, leg.” Other material. (4) — AL: Marshall Co., Grant, 25-May-1958 (1 @); Colbert Co., Maud, nr. McCluskey Cave, 26-Mar-1962 (3 9, 1 @). Diagnostic description. This species is externally identical to E. adversus, above, and can only be distinguished by male genitalic characters. Both species share very prominent antennal carinae (e.g., Fig. 14C), those on antennomeres VIII and IX both strong and 164 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Figure |5. Aedeagus A Euconnus adversus (type locality - Celo Knob, NC) B Euconnus adversus (non- type locality —- Chestnut Ridge, SC) C Euconnus astrus D Euconnus cultellus. oblique, that of VII most produced at base and that of [IX most produced at apex. They similarly appear to be wing-dimorphic, with winged males and wingless females; aedea- gus (Fig. 15C) with median lobe evenly tapered and narrowly rounded at apex; para- meres thin, each bearing three short terminal setae, the setae not reaching apex of median lobe; compressor plate short, asymmetrical, truncate on one side (left as drawn), nar- rowly and unevenly produced on other; left upper endophallic sclerite strongly hooked, with prominent, blunt median tooth on inner margin; right upper endophallic sclerite a short curved spine, the tip pointing laterad; lower endophallic armature comprising a deeply bifurcate process, with two long, slightly sinuate spikes pointed distad. Distribution. This species is only definitely known from northeastern Alabama, where it has been found in a few caves, as well as a few free-living situations. A series from Colbert County, in northwestern Alabama, comprises only females. Males from this locality would be interesting to examine. Remarks. ‘This species appears closely related to E. adversus, but differs substantial- ly in genitalic characters, with the right upper endophallic sclerite quite different, and the lower endophallic sclerites much more elongate. The species name means ‘starry’, referring to the nearby NASA rocket science and spacecamp facilities. Euconnus cultellus sp. nov. https://zoobank.org/796C1879-2BB4-45 15-815D-FDBDAF6C43AD Figs LSD s2lGe V7 Type material. Holotype 3, deposited in FMNH: “USA: GA: Rabun Co., 34.9658°N, 83.2997°W, ChattahoocheeNE, Rabun Bald, 4663’, v.11.2021, sifted litter, M.Caterino & A.Haberski” / “[QR code] CLEMSON-ENT CUAC000146083” / “Caterino DNA Voucher Extraction MSC12039”. Paratypes (8) — 5 2,5 3: same data as type. Appalachian Euconnus (Cladoconnus) 165 Figure 16. Habitus and character photos of Euconnus cultellus A male, dorsal view B female, dorsal view C SEM of male antenna. Other material. (28) GA: Clay Co., Chattahoochee National Forest, Brasstown Bald, 4590ft, 19-Sep-2015 (3 3); NC: Cherokee Co., Nantahala National Forest, Hickory Branch trail, 3923ft., 26-Jul-2015 (1 4); Cherokee Co., London Bald Tr., 4108ft, 26-Jul-2015 (2 2); Graham Co., Nantahala National Forest, Teyahalee Bald, 4591ft., 12-Apr-2022 (1 3); Macon Co., Nantahala National Forest, Cowee Bald, 4942ft., 9-Jul-2019 (2 3’); Macon Co., Nantahala National Forest, E. Highlands, Hwy 64, 3880ft., 1-Mar-2020 (1 9 1 4); Macon Co., Nantahala National Forest, Cop- per Ridge Bald, 5032ft., 15-Sep-2020 (1 @); Swain Co., Nantahala National Forest, Miller Cove app trail, 2323ft., 20-Jul-2015 (2 9, 6 4); SC: Oconee Co., Buzzards Roost Heritage Preserve, 1250ft., 16-Jan-2015 (1 3); Oconee Co., Chau-Ram Coun- try Park, 850ft., 15-Oct-2015 (1 9, 1 4); Oconee Co., Sumter National Forest, Yellow Branch Falls, 1560ft., 12-Oct-2017 (2 2, 1 3); Oconee Co., Sumter National Forest, Chattooga river, 1580ft., 2-Apr-2015 (1 ); Clay Co., Nantahala National Forest, Tusquitee Bald, 5015ft., 1-Sep-2020 (1 3). Diagnostic description. This species exhibits few obvious external differences from the preceding ‘falcatus complex’, and can best be distinguished by male geni- talic characters; body color sometimes darker (Fig. 16A); carina of male antennomere VIII (Fig. 16C) more strongly inclined, produced further at the base, than in most of the previous (except E. adversus and E. astrus), and carina of male antennomere IX slightly oblique, ~ 20 degrees off the long axis of the antenna; males at least sometimes 166 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) * E. cultellus Figure 17. Map of collecting records for Euconnus (Cladoconnus) cultellus. winged, females apparently wingless; aedeagus (Fig. 15D) with apex of median lobe knobbed, bluntly truncate; parameres short, curved, bearing 3 setae at their apices; compressor plate more or less symmetrical, narrow, parallel-sided from near base, sinu- ately tapering to subacute apex; endophallic armature with upper blade long, nearly straight, widened beyond middle, curving and tapered to bifid, acute apices (a second tooth projecting straight behind the uppermost); a second upper sclerite bends laterad strongly from the base, curving distad and tapering to long thin apex beneath ‘right’ edge of compressor plate; lower sclerite with two strong hooks, the basalmost broad and blunt, the apicalmost sinuately curving behind apex of upper endophallic sclerite. Distribution. Euconnus cultellus occurs across a relatively limited portion of west- ern North Carolina, in the Cowee and Nantahala Mts, northeast Georgia, and upstate South Carolina. Its distribution seems to be limited on the east by the Little Tennessee River system, not (yet) found east of the Tuckasegee tributary, and not having been found in the Great Smoky Mountains, or on any of the spruce-fir peaks sampled. That limit aside, it has a broad elevational range, occurring from 850—5000ft. Remarks. The bifid apex of the upper endophallic sclerite is distinct from all other Cladoconnus species except E. cataloochee. The distinctive broad basal hook of the lower endophallic sclerite differentiates FE. cultellus from all others, as does the deeply curved, apically slender, tapering right arm of the upper armature. This species is named for the ‘cutting’ edge of the males finely serrate antennal carinae. Appalachian Euconnus (Cladoconnus) 167 Phylogeny Phylogenetic analysis of available COI sequences does not support monophyly of American Cladoconnus relative to all other Euconnus (Fig. 18). Most of the new species fall out in one large clade. But Euconnus vetustus is resolved as a separate clade outside all the others. This is a dark, flightless species that lacks male antennal modifications. It is also an unusually widespread species, known from several localities on both sides of the Asheville depression. Neither lineage falls out near the two European Cladoconnus, E. (C.) denticornis Miller & Kunze and £. (C.) carinthiacus Ganglbauer for which COI sequences are available, with those forming a sister lineage to most of the rest of Euconnus (within which the Cladoconnus vetustus lineage falls out). However, it must be acknowledged that all these lineages (the three distinct ‘Cladoconnus lineages) are highly divergent (> 20% K2P distance), and COI struggles to resolve these deeper relationships. It is worth noting that two American Euconnus sensu stricto species fall out near some European members of that subgenus, and far from any of the newly described Cladoconnus — these new species had initially been considered as potentially belonging there. Overall, these deeper results are based on a very sparse sampling of global Euconnus diversity, and only a small fragment of a single mitochondrial gene, so can only be given so much credence. Within the main American Cladoconnus lineage, the species are divided into two main clades, one containing Euconnus megalops, E. cumberlandus, E. vexillus, E. adversus, and E. kilmeri. Euconnus astrus is not represented by molecular data, but it may be ex- pected to be a member of this group, likely as sister to E. adversus. Euconnus megalops is probably not yet adequately resolved, as one sample considered to belong there (the Brasstown Bald locality) falls well outside the main group, as sister to a Euconnus megalops + E. vexillus lineage. But this male’s genitalia do not differ obviously from the rest of E. megalops. Euconnus vexillus, on the other hand, exhibits distinct male genitalia and male antennal characters, so is clearly a distinct species. More sequences and more genitalia from outlying localities will be necessary to resolving this uncertainty. There is also considerable genetic diversity within the widespread species E. megalops. But it shows relatively little geographic structure, with populations from both sides of the Asheville Basin somewhat intermingled. There is one subclade within this that shows a higher degree of genetic variation, from mostly southwestern localities (Huckleberry Knob, Tusquitee Bald, etc.). But these show no obvious morphological coherence. The other large clade has Euconnus cultellus as sister to Euconnus attritus and sev- eral very closely related species of the Euconnus falcatus complex. Euconnus cultellus occurs in only the southern part of the region, from scattered localities in the Nanta- hala Mountains just into far western South Carolina. There is considerable genitalic variation site-to-site, but their sequences, while showing some diversity, don’t suggest significant differentiation. Euconnus attritus occurs only in the northeastern part of the region, including the Blacks, Grandfather Mt., and the Roan Highlands, showing potentially meaningful COI differences among the localities. The Euconnus falcatus complex, on the other hand, includes three quite distinctive genitalic forms that show surprisingly little COI differentiation. One, Euconnus cataloochee, is even scattered 168 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Brachycepsis Nc-Jk_11399 Euconnus kilmeri force ----- Euconnus astrus?? ' NC-BMP_26255 So NSC OHHP. 25642 nT_A.009 | Euconnus (C.) adversus BgBid_A_026 Ck_B_370 WR_A_046 TN_145978 Euconnus cumberlandus BBid B 334 megalops?? NT_A_ 00} SC-ChestnutHP_26905 . NC-3B GmR_107838 =| _Euconnus (C.) vexillus SC-CRHP_26945 NG-Mackey Mt_11946 A NC-Melrose_ 160199 NC-Melrose_171982 SC-Eastatoe_171981 NC-M2STr 111147 MHy_B_320 NC-Whiteside_171831 CB_019 CB_B_351 CrB_017 CrB_018 TsqB_B_326 NC-Cherohala_109850 HKnb_A_011 HKnb_B 314 TN-WhiteOak_157337 BCat_B 316 Euconnus (C.) megalops BCat_A_009 ( ) 9 p BCat_A_039 NC-CmpCkBld_ 160280 WR_A_029 BoB At 6-LARVA BgBid_B_349 era wit -LARVA BM GA-RabBlid_ 146083 GA-BBld_20004 SC-Buzz_71962 Sc-ChauRam_110358 SC-YellBrnch_107623 CrB_B_321 Euconnus (C.) cultellus NC-Teyahalee_ 166075 NC-MillCove_109165 NC-TsqB_B_327 NC-HickBrnch_11636 NC-Highlands_108294 SC-Chattooga_171975 RHB 13 BgBld_B_348 Euconnus (C.) attritus NC-CmpCkBld_160266 GrM_B_305 NG-GrM_ 166260 NC-GrM_166308 Hkte-Bss9 | Eueonnus (C.) draco LLB 307 LL_A_157 NC-BMP_26304 BMT_B_303 * NC-BMP_25813 CrB_014 SC-Tox_ 143418 BCat_A_008 * Euconnus (C.) falcatus CD_B 307 Hwy_A 006 & MKB 482 * Euconnus (C.) cataloochee BCat_A 062 * BCat_B 315% MLc_009 CD_A021 BBK_A019 NC-M2STr_111154 NC-Mt.Pisgah BBK_B 303 Euconnus (C.) denticornis Euconnus (C.) carinthiacus Euconnus sp. Euconnus (Ss. str.) sp.1 Euconnus (s. str.) sp.2 Euconnus sp. Euconnus sp. Euconnus sp. Euconnus (s. str.) hirticollis Euconnus (s. str.) fimetarius Euconnus sp. Euconnus (s. str.) rutilipennis Euconnus sp. Euconnus (Napochus) sp. Euconnus (Napochus) sp. Euconnus (Neonapochus) maeklini Euconnus (Napochus) spp. Euconnus (Psomophus) sp. Euconnus sp. Euconnus sp. Euconnus (Tetramelus) pubicollis HKnb_B 313 BBK_A_020 GRB_A_336 Hwy_A 029 BgBld A 018 Euconnus (C.) vetustus MNC-DevGap_26607 TN-BgBld_142207 BgBid_B_332 Figure 18. Phylogeny of American species of Euconnus (Cladoconnus). Hypothesized position of E. astrus, for which no sequence data is available, is indicated with a dotted line. Appalachian Euconnus (Cladoconnus) 169 within another both genetically and geographically (they are marked by asterisks in Fig. 18). This non-monophyly is peculiar and requires further genetic data to resolve. The genitalic differences between FE. falcatus and E. cataloochee are marked and consist- ent. [wo individuals representing E. draco resolve as sisters to the remainder of this clade, but are not themselves monophyletic (despite coming from the same locality, Huckleberry Knob). Euconnus tusquitee would also be expected to be very similar to these, but it has not yet yielded clean sequence data. These results together indicate either rapid speciation, over a time frame insufficient for coalescence of species-specific haplotypes, or a significant slow-down in mitochondrial evolution. Again, additional markers will be necessary to resolve this question. Discussion The discovery of a diverse, previously unreported radiation of beetles in the southern Appalachians is surprising, even considering their small size and cryptic habits. The region has been popular with collectors and researchers for many years, and while new species are still encountered commonly, it is rare for major lineages to have escaped de- tection. Scydmaeninae have perhaps received less attention from taxonomists over the years than more prominent groups, and one could imagine other ‘hidden’ radiations in other similarly neglected arthropod taxa. While only European members of Cladoconnus have previously been sequenced, it is worth considering whether the relationships of the Appalachian species lie among the Asian species. There aren't any particular morphological characters that suggest such a relationship. But the existence in Japan and Korea of species that are sexually dimorphic in the possession of flight wings, as observed in several Appalachian spe- cies, might be informative. Then again, that begs the question of how a lineage with flightless females may have reached the area to begin with. But the deep genetic diver- gences, and somewhat incongruous distributions of some of the species (in particular, E. vetustus ignoring the biogeographically significant Asheville Depression) together suggest that Cladoconnus has been resident in the southeastern US for a very long time. Given that, however, it is surprising that their overall distribution is still somewhat limited. The species found to the west, in parts of the Cumberland Plateau, extend somewhat beyond Appalachia proper. But none have yet been found to occur in higher elevations of Virginia. Some older lineages of Appalachian arthropods find relatives in the Ouachita Mts of Arkansas (flightless Lathrobium, for example — Watrous and Haberski, pers. comm.; Arianops Brendel — Carlton 1990; Anillinus Casey — Allen 1990, Carlton and Robison 1998). It might be worth examining such collections more closely for overlooked representatives. Our ‘barcode-everything’ approach revealed several larvae of Cladoconnus, all, as definitely associated by DNA placement, of E. (C.) megalops (Fig. 19). Of the sev- en extracted and sequenced, vouchers of six survived the process more or less intact. No larvae of the larger genus Euconnus have yet been formally described (Newton 1991; Jatoszynski. pers. comm,), and I do not do so here. However, these represent a 170 Michael S. Caterino / ZooKeys 1137: 133-175 (2022) Figure 19. Larvae of Euconnus (Cladoconnus) megalops A dorsal view (specimen from Rabun Cliffs, GA) B lateral view (specimen from Woody Ridge Trail, NC). potentially valuable source for future character data for reconstructing scydmaenine relationships. Generally, the larvae resemble other described Glandulariini, such as Stenichnus Thomson (Wheeler and Pakaluk 1983; Newton 1991; Jatoszyriski and Kil- ian 2012), prognathous, elongate, setose, slightly flattened, with a rather broad abdo- men, and a large antennomere II with a simple, domelike apical cap. While the bulk of our records seem to suggest an exclusively high-elevation re- stricted group, with more than two-thirds of the available records coming from above 4500 feet (Fig. 20), the ranges of many of the species extend considerably lower, even below 1000 feet. These lower records tend to be from deep riparian canyons, and the higher humidity of such locations is probably part of the explanation. However, there is also a clear sampling bias here, in that recent fieldwork by our lab has targeted high elevation sites. Still, scanning our own older, lower elevation samples, and attempting to borrow specimens from other collections where lower elevation samples were bet- ter represented turned up relatively few additional specimens. It may be that many of these beetles have been ignored or overlooked in the abundant morass of litter micro- coleoptera. But there have been a number of dedicated devotees over the years, includ- ing Sean O’Keefe, Donald Chandler, Christopher Carlton, Walter Suter, Thomas Barr, and others, so the apparent predilection of Cladoconnus for higher elevations seems likely to be real. Activity patterns of the species present similar questions. The majority of records come from late spring and early summer samples (Fig. 21), and such a pattern would conform to many species in the region. Our own sampling has had some of this bias, although for the past several years, during which most of these beetles were collected, similar emphasis was given to fall sampling, in September and October, and there is no indication of a secondary peak of specimen records, so there is probably some mean- ingful signal there, as well. The range in genitalic diversity, especially among the apparently closely related species in the E. falcatus complex, is remarkable. In those species known from multiple Appalachian Euconnus (Cladoconnus) 171 elevation (ft.) 200 180 160 140 120 100 80 60 40 ai = , === [0, 1000] (1000, 2000] (2000, 3000] (3000, 4000] (4000,5000] (5000,6000] (6000, 7000] Figure 20. Histogram of record counts (y-axis) by elevation (x-axis, range in feet). month of activity 40 | = * a a, 0 | (1, 2] (2, 3] (3, 4] (4, 5] (5,6) (6,7) (7,8} (8, 9] (9, 10) (10, 11] Figure 21. Histogram of record counts (y-axis) by month (x-axis). localities, it seems to be the rule that variation in specific conformation of endophallic sclerites can be seen. I’ve taken a generally conservative approach in considering many to be widespread, variable species for now. But closer study could well split some of these more finely. Me? Michael S. Caterino / ZooKeys 1137: 133-175 (2022) It would be fascinating to understand the in situ mechanisms of these species in- credibly complex genitalia. It is tempting to hypothesize explanations of sperm com- petition and perhaps rival sperm removal, or of active female choice spurring an arms- race of male elaborations for maintaining hold and proper position. The dimorphic antennomeres of at least some of these species might also point to some similar sorts of intersexual dynamics. Unfortunately, for the present such a discussion would be pure speculation. Simply understanding the physical mechanisms that deploy the endophal- lic sclerites during intromission would constitute a major undertaking. Yet it might repay the effort, as similar dynamics may pertain to the diversification of a wide variety of ‘dark taxa’, within Staphylinoidea at least, where such structures are commonplace. For now, it must suffice to call attention to these remarkable creatures, and hope that future workers with a more applicable skillset take up the challenge. Acknowledgements This study was funded by the U.S. National Science Foundation (Award DEB- 1916263 to MSC) and the Clemson University Experiment Station (SC-1700596 to MSC). I also acknowledge the support of the John and Suzanne Morse Endowment for Arthropod Biodiversity. For permissions and assistance with field work I am grateful to the North Carolina State Parks, Great Smoky Mountains National Park, Blue Ridge Parkway National Park, Monica Martin, Frank Etzler, Ernesto Recuero, Curt Harden, Anthony Deczynski, Patricia Wooden, Adam Haberski, Roy Kucuk, Laura Vasqez- Velez, Laary Cushman, Paul Marek, Michael Ferro, and Will Kuhn. For other advice and specimens I thank Pawel Jaloszynski, Peter Hlavac, Donald Chandler, Alfred New- ton, Anthony Davies, Zachary Falin, and Christopher Carlton; and for assistance in the lab I thank Mary Atieh, Caroline Dukes, Caroline McCluskey, Grace Holliday, Grace Arnold, Hannah Skinner, and Anthony Villanueva. Finally, thank you to Regi- nald Webster for comments that helped improve the manuscript. This paper represents Technical Contribution No. 7105 of the Clemson University Experiment Station. References Allen RT (1990) Insect endemism in the interior highlands of North America. 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Bulletin of the American Museum of Natural History 290: 1-95. https://doi.org/10.1206/0003-0090(2005)290<0001:SBOT GA>2.0.CO;2 Wheeler QD, Pakaluk J (1983) Descriptions of larval Stenichnus (Cyrtoscydmus): S. turbatus and S. conjux, with notes on their natural history (Coleoptera: Scydmaenidae). Proceedings of the Entomological Society of Washington 85: 86-97. Appalachian Euconnus (Cladoconnus) 175 Supplementary material | Specimen data for all material examined Authors: Michael S. Caterino Data type: occurrence Explanation note: Specimen level data for all Cladoconnus specimens examined in this paper. Columns include type status, repository, voucher numbers (for Clemson University Arthropod Collection), morphospecies designation (which can be searched on Flickr for additional images), DNA extraction number (extractions in Caterino Lab/CUAC collection), GenBank accession numbers, and collection event and geographical information. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODDbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1137.97068.suppl1 Supplementary material 2 Phylogenetic character data (COI barcoding region) Authors: Michael S. Caterino Data type: DNA sequence Explanation note: Nexus file containing all sequences analyzed in this paper, includ- ing Brachycepsis outgroups, new Cladoconnus species, new sequences for Nearctic Euconnus (Napochus) and Euconnus s. str. spp., and previously published Euconnus sequences from GenBank and BOLD. Line names contain their BOLD and/or GenBank Accession numbers for previously published sequences. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODDbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1137.97068.suppl2