Anillinus jancae, Harden & Caterino, 2024

Harden, Curt W. & Caterino, Michael S., 2024, Systematics and biogeography of Appalachian Anillini, and a taxonomic review of the species of South Carolina (Coleoptera, Carabidae, Trechinae, Anillini), ZooKeys 1209, pp. 69-197 : 69-197

Anillinus jancae sp. nov.

Figs 10 A, D , 12 E , 13 B , 14 A, C, E , 21 N , 23 D – F , 24 M , 25 C , 26 A – C , 27 A , 43 C

Type material.

Holotype male ( USNM): point mounted, with genitalia in Euparal on microslide pinned beneath specimen. Original label: “ USA: SC, Abbeville Co. Sumter NF Long Cane Creek . 34.1350, - 82.3239. 15. March- 16. July. 2020. C. W Harden. Buried pipe trap baited with cheese. Deep clay soil. Oak, hickory, beech. LCC- 01-0716. ” “ CLEMSON-ENT CUAC 000168362 ” “ HOLOTYPE Anillinus jancae Harden & Caterino [orange cardstock] ”

Paratypes (n = 7, CUAC). USA • South Carolina • Abbeville Co. • Sumter National Forest, Long Cane Creek ; • 1 ♂; same data as holotype; CUAC 000168363 , CWH- 204 ; • 1 ♀; same data as holotype; CUAC 000168364 , CWH- 205 ; • 5 ♀; same data as holotype; CUAC 000168365 to CUAC 000168369 .

GenBank accession numbers for paratypes: OR 853233, OR 839266, OR 838005.

Other material

(n = 2). USA • South Carolina • Abbeville Co. • Sumter National Forest, Long Cane Creek • 1 ♀; 34.1362, - 82.3235; 21 March 2021; C. W. Harden leg.; Underside of rock. CWHc ; • 1 ♀; Near Cedar springs Rd. ; 34.10914, - 82.33948; 25 September 2020; C. W. Harden leg.; Underside of large rock; CUAC, CUAC 000168370 , CWH- 241 .

Diagnosis.

Largest species of Anillinus in SC, with distinctive habitus (Fig. 26 A ), dorsoventrally flattened and not as compact as typical Anillinus ; head relatively small (HW / PW 0.71–0.74); pronotum cordate, with sides subparallel at base, hind angles of pronotum prominent and rectangular or acute and extending posteriorly past hind margin. Protibiae of both sexes with a deep semicircular outer notch apically (Fig. 10 A ). Males with unique combination of secondary sexual characters: profemora with ventral spine (Fig. 27 A ), hind femora swollen and tuberculate, hind tibiae bowed inward and with inner surface scalloped, first abdominal ventrite bearing a short fin-like carina medially on posterior margin (Fig. 26 B, C ). Females with short spermatheca (Fig. 21 N ).

Description.

Habitus Body dorsoventrally flattened and large (ABL = 2.32–2.64 mm). Males (2.63–2.64 mm) larger than females (2.32–2.36 mm). Less compact than most Anillinus , with relatively narrow forebody (HW / PW 0.71–0.74, PW / EW = 0.76–0.83]). Pronotum with sinuate sides and long, parallel-sided posterior angles. Integument Dorsal surfaces of forebody mostly covered with irregularly isodiametric mesh of microsculpture, microsculpture absent from at least a small area on pronotal disc on each side of midline and from center of vertex, females more extensively microsculptured than males. Elytra with coarse mesh of isodiametric microsculpture, sculpticels small. Head HW / PW = 0.71–0.74. Antennomeres I – IV [♂] or I – III [♀] longer than wide, V – X [♂] or VI – X [♀] moniliform. Frontoclypeal horn small, nearly absent in some specimens. Two pairs of supraorbital setae present. Mentum with median pair of setae posterior to bead of mentum tooth, which is relatively small and obtusely triangular, blunt. Pronotum Cordate, base strongly constricted in females (average Pbw / PW = 0.71, n = 3), moderately so in males (average Pbw / PW = 0.75, n = 2). Average PW / EW 0.81 (females), 0.77 (males). Relatively short (PL / ABL = 0.22 in all specimens measured, n = 5). Sides sinuate before long hind angles which are parallel sided and rectangular or slightly acute and projected posteriorly. Elytra Parallel sided and flat, relatively long (EL / ABL = 0.54–0.57); each with five striae, inner two or three more strongly impressed than outer two; small subapical plica evident; fused inner margins carinate at apex. Legs Profemora of males modified, swollen, and bearing a peg-like spine on posterior margin at proximal 1 / 3 (Fig. 27 A ). Protibiae of both sexes with deep semicircular notch on outer edge of apex; males with inner margin of protibiae extended as a bifid spine (Fig. 10 A ). Protarsi of males with first protarsomere greatly enlarged and produced on inner surface as a blunt lobe, with adhesive setae ventrally, second protarsomere unmodified and without ventral adhesive setae (Fig. 10 D ). Mesotrochanters of males unmodified. Male metatrochanters flattened, elongate, and covered in small tubercles; metafemora of males swollen and bearing several tubercles along posterior margin and a blunt angulate protrusion in distal 1 / 3 (Fig. 12 E ). Metatibiae of males concave along inner margin. Tarsomeres of middle and hind legs relatively short in both sexes. Abdominal ventrites Males with keel-like median carina at posterior margin of second visible ventrite (Fig. 26 C ); females without modifications. Male genitalia Ring sclerite large (RL / ABL = 0.28), oval; margins thickened; narrowed end with margin slightly deflexed; sides with small flattened lateral expansions medially. Median lobe (Fig. 23 F ) heavily sclerotized, with several parallel, diagonal sulci across surface in proximal 1 / 3 on left side, appearing as dark lines; strongly bicarinate ventrally, carinae forming a channel along most of ventral surface; band of many hairlike setae present across ventral surface and left side medially; apex expanded, with sides strongly curved ventrally, appearing bill-like in head-on view; in right lateral view, apex appears hatchet shaped, and thickened portions of curved cuticle appear as dark linear structures. Internal sac with long flagellum abruptly narrowed and filamentous past bulb-like base; small sclerotized straplike structure present ventrally at ostial opening. Right paramere (Fig. 23 E ) relatively short, with blunt apical margin bearing four or six long, thick setae that surpass the apex of the median lobe when paramere is attached. Left paramere (Fig. 23 D ) bluntly rounded at apex, with four poriferous canals and no setae on apical margin. Female genitalia Spermatheca short and small, abruptly enlarged distally, with short curved stem (Fig. 21 N ). Spermathecal duct longer than spermatheca, with a few loose coils.

Distribution.

Known for sure only from a single hillside on the southeast side of Long Cane Creek in Abbeville Co, within Sumter National Forest (Figs 25 C , 43 C ).

Sympatry.

Anillinus chandleri and A. dentatus have been collected with this species. Serranillus dunavani is known from nearby.

Natural history.

Members of this species are endogean in habit. Most of the type series was collected from a buried pipe trap operated from March to July. The trap was set deeply in a layer of pure red clay, well below the shallow organic soil horizon. A late-instar larva was found underneath a deeply embedded rock in early February, indicating either overwintering of the egg or larval stage, or winter breeding and oviposition. A female adult was collected underneath a large, embedded stone. This female was kept alive for several months in a container with soil from the same locality packed into the bottom. The female quickly found a way around and underneath the packed clay soil on the bottom of the container, and apparently spent the remainder of its life in an inverted position on the underside of the clay. Given this behavior, and the microhabitat in which the trap that collected most of the type series was set, the habitat of this species is probably the series of crevices formed naturally in clay soils. Although clay is typically thought to be impervious, it forms naturally into aggregates called “ peds ” ( Sherwood and Garst 2016), creating a series of crevices through which air and water (and small invertebrates) can pass. This habitat is widely distributed in the southeastern U. S., and further targeted sampling of it using buried traps is likely to discover many more species of anillines and other subterranean arthropods.

Species status justification.

The overall habitus, male secondary sexual characters, great length of setae on the apex of the right paramere, and characters of the median lobe are all unique within the genus. The consistent placement of this species in a clade with the geographically distant A. albrittonorum support its distinction from other Anillinus species.

Derivation of species name.

This remarkable species is named in honor of Janet C. Ciegler, in recognition of her contributions to the study of Coleoptera in South Carolina and the southeastern United States. Her many identification guides have made the study of beetles more accessible to amateur and professional entomologists alike. The specific name is a genitive noun derived from the shortened first name (“ Jan ”) and first letter of the surname.

Notes.

The female from the Cedar Springs Road site ( CUAC 000168370 ) is from the opposite side of Long Cane Creek, and differs from the type series in several respects. The specimen is smaller (ABL = 1.88 mm) and more compact, resembling A. dentatus in habitus. The head also has three supraorbital setae on each side, versus two in the type series. The spermatheca agrees with the females of the type series. DNA sequence data indicate that this female could represent a different species. The uncorrected p-distance of the barcoding region of COI of this specimen is 3.7 % and 3.8 % divergent from the two paratypes with sequenced barcodes. Its 28 S sequence differs from that of the paratype male at only two sites – one substitution and a 2 - bp insertion. CAD 4 sequences of these same two specimens differ in 15 nucleotides. Male specimens from the Cedar Springs Road site would help resolve the situation. We note that members of A. dentatus have been collected from the same two sites, and show no differences in COI or 28 S, so Long Cane Creek is likely not a barrier to dispersal in that species.

The male holotype of A. jancae is the only specimen in the “ quadrisetose clade ” to have more than four apical setae on the right paramere. Considering that the right paramere of the other known male of A. jancae is quadrisetose, we interpret the extra setae on the right paramere of the holotype to represent an unusual variant.

Our description of Anillinus jancae represents the first documented example of modified profemora in the genus. However, we found that the male profemora of the previously described Anillinus lescheni are also modified, though quite different in form, having a large, triangular tooth distally (Fig. 27 C ). Two undescribed species with modified male profemora are known, from North Carolina (Fig. 27 B ) and Alabama (Fig. 27 D ). The profemora of these two undescribed species are similar in form to A. jancae and A. lescheni , respectively. In the case of the North Carolina species, from which we have DNA sequence data, the similarities are the result of convergence.

‘ valentinei group’

Figures

Figure 10. SEM micrographs of male protarsi A Anillinus jancae, left protarsus, dorsal aspect B Anillinus mica, left protarsus, dorsal aspect C Anillinus montrex left protarsus, dorsal aspect D A. jancae, right protarsus, ventral aspect E Anillinus choestoea, right protarsus, ventral aspect F Anillinus dentatus, right protarsus, ventral aspect G Serranillus dunavani (Jeannel), right protarsus, ventral aspect H Anillinus murrayae Sokolov & Carlton, right protarsus, ventral aspect I A. montrex, right protarsus, ventral aspect. Scale bars: 0.1 mm.

Figure 12. SEM micrographs of male left metafemora of Anillinus species. Scale bar: 0.1 mm.

Figure 13. Dorsal habitus of late-instar larvae A Serranillus dunavani (Jeannel) B Anillinus jancae. Scale bars: 1 mm.

Figure 14. Details of late-instar larvae of Appalachian Anillini A head of Anillinus jancae sp. nov., dorsal aspect, left antenna and right mandible omitted B left mandible of Serranillus dunavani (Jeannel), dorsal aspect C right maxilla of A. jancae, dorsal aspect D right maxilla of S. dunavani, dorsal aspect E urogomphi of A. jancae, dorsal aspect. Scale bar: 0.1 mm.

Figure 21. Spermathecae of Anillini species A, B Anillinus chandleri C Anillinus castaneus D Anillinus murrayae E Anillinus simplex F Anillinus dentatus G Anillinus sp. “ South Carolina, Coon Branch ” H Anillinus micamicus I Anillinus mica J Anillinus choestoea K Anillinus seneca L Anillinus montrex M Anillinus arenicollis N Anillinus jancae O Anillinus cf. nantahala P Anillinus cherokee Q Anillinus loweae R Anillinus merritti S Serranillus dunavani T Serranillus jeanneli. Scale bar: 0.1 mm.

Figure 23. Male genitalia of Anillinus dentatus (A – C) and Anillinus jancae (D – F). Median lobes in right lateral (C) and right dorsolateral (F) aspects. Left parameres (A, D) and right parameres (B, E) in left and right lateral aspects, respectively. Scale bars: 0.1 mm.

Figure 24. Male ring sclerite, ventral aspect. A Anillinus chandleri B Anillinus castaneus C Anillinus sp. “ South Carolina, Chestnut Ridge ” D Anillinus murrayae E Anillinus simplex F Anillinus dentatus G Anillinus mica H Anillinus micamicus I Anillinus choestoea J Anillinus seneca K Anillinus arenicollis L Anillinus montrex M Anillinus jancae N Anillinus loweae O Anillinus cherokee P Anillinus merritti. Scale bar: 0.5 mm.

Figure 25. Anillini habitat in South Carolina A Waldrop Stone Falls, Pickens Co. (Anillinus mica, Anillinus micamicus, Anillinus murrayae, Serranillus dunavani) B Lower Whitewater River gorge, Oconee Co. (Anillinus cherokee, A. murrayae, Anillinus sp. “ South Carolina, Coon Branch ”, S. dunavani, Serranillus jeanneli, Serranillus sp. “ South Carolina, Coon Branch ” C Long Cane Creek, Abbeville Co. (Anillinus chandleri, Anillinus dentatus, Anillinus jancae, S. dunavani). The holotypes of A. mica and A. micamicus were collected under embedded rocks uphill of the wooden staircase in A. The holotype of A. jancae was collected in a pipe trap set on a hill just out of view in C.

Figure 26. Anillinus jancae A habitus B abdominal ventrites, left lateral aspect C SEM micrograph of abdominal ventrites, ventral aspect. Black arrows point to abdominal keel. Scale bar: 1 mm.

Figure 27. Modified profemora of Anillinus species. A Anillinus jancae B Anillinus sp. “ North Carolina, Orange Co. sp. 2 ” C Anillinus lescheni D Anillinus sp. “ Alabama, Aladdin Cave sp. 1 ”. White arrows point to protrusions on profemur and / or protrochanter. Scale bars: 0.1 mm.

Figure 43. Topology and distribution maps of “ quadrisetose clade ” species groups of Anillinus A collapsed maximum likelihood tree of “ quadrisetose clade ” Anillinus species, from 6 - gene concatenated core matrix, SBS values shown below nodes B distribution map of all known occurrences of Anillinus erwini and the valentinei group C distribution map of all known occurrences of the “ ESP + LSL ” clade and albrittonorum group D distribution map of all known occurrences of the “ ESP clade ” E distribution map of all known occurrences of the langdoni group and Anillinus sp. “ Tennessee, Kings Saltpeter Cave ” F distribution map of all known occurrences of the steevesi group G distribution map of all known occurrences of the loweae group. Data for distribution maps come from Harden (2024).