Anillinus mica, 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 mica sp. nov.

Figs 10 B , 12 A , 21 I , 24 G , 25 A , 34 E , 35 G – I , 36

Type material.

Holotype male ( USNM): point mounted, with abdominal ventrites glued to point and genitalia in Euparal on microslide pinned beneath specimen. Original labels: “ USA: SC, Pickens Co. Clemson Experimental Forest Waldrop Stone Falls . 34.7393, - 82.8205. 8. x. 2021. CW Harden. Under rock on steep slope near falls. ” “ [QR code] CLEMSON-ENT CUAC 000163558 ” “ Harden DNA Voucher CWH- 403 Anillinus ‘ caterino’ M Ext. 19 - December- 2021 [green-bordered cardstock] ” “ HOLOTYPE Anillinus mica Harden & Caterino [orange cardstock] ”

GenBank: OR 853287, OR 839293, OR 838076.

Paratypes (n = 18, CUAC, USNM). USA • South Carolina • 2 ♂; Pickens Co.; Keowee WMA, near Todd Creek Falls ; 34.74986, - 82.81467; 6 Sep. 2020; C. W. Harden leg.; Under embedded rock on dry soil, mixed woods; CUAC 000163551 and CUAC 000163552 , CWH- 230 and CWH- 231 ; • 1 ♂; Pickens Co.; Central , ~ 3 mi N Clemson; 34.7251, - 82.8248; 2 Oct. 2020; C. W. Harden leg.; Buried pipe trap, sandy clay mica - rich soil; CUAC 000134435 ; USNM ; • 1 ♂; same locality as previous; 14 Aug. to 24 Oct. 2021; C. W. Harden leg.; Buried pipe trap, clay soil, trap 07; CUAC 000170072 ; USNM ; • 1 ♂; same locality as previous; 24 Oct. 2021 to 10 Apr. 2022; C. W. Harden leg.; Buried pipe trap, mica - rich sand / clay. Trap 05; CUAC 000170073 ; • 1 ♂, 1 ♀; Pickens Co.; 5 mi N Clemson ; 34.7252, - 82.8245; 8 Nov. 2016; M & P Caterino leg.; floated from soil; CUAC 000163553 and CUAC 000163554 , MSC- 2458 and MSC- 2466 ; • 1 ♂, 1 ♀; Pickens Co.; Clemson Experimental Forest, Issaqueena Lake Rd., West side ; 34.74136, - 82.86541; 12 Nov. 2020; C. W. Harden leg.; Underside of rock, clay soil; CUAC 000163555 and CUAC 000163556 , CWH- 333 and CWH- 334 ; • 1 ♂; Pickens Co.; Clemson Experimental Forest, Waldrop Stone area ; 34.73603, - 82.81725; 12 Nov 2020; C. W. Harden leg.; Under large rock at lake edge; CUAC 000163557 , CWH- 328 ; • 3 ♂, 3 ♀; Pickens Co.; Clemson Experimental Forest, Waldrop Stone Creek ; 34.7395, - 82.8270; 11 Sep. 2021; C. W. Harden leg.; Under embedded rock on stream bank; CUAC 000163559 to CUAC 000163564 , CWH- 409 to CWH- 414 ; • 1 ♀; Pickens Co.; Clemson Experimental Forest ; 34.7431, - 82.8481; 14 Oct. 2019; C. W. Harden leg.; Soilwash-flotation berlese, mesic oak-pine woods, dark sandy soil near stream; CWH- 057, CUAC 000168224 ; • 1 ♀; Pickens Co.; Clemson Experimental Forest ; 34.74265, - 82.84167; 23 Nov. 2019; C. W. Harden leg.; On underside of large embedded rock during rain, oak-hickory woods, clay soil; CUAC 000168225 , CWH- 060 .

GenBank accession numbers for paratypes: OR 839739, OR 839745, OR 839626, OR 853280, OR 839410, OR 853281, OR 839413, OR 853283, OR 838130, OR 853110, OR 853285, OR 839292, OR 839553, OR 837876, OR 838021, OR 838209, OR 838124, OR 853286, OR 839554, OR 837877, OR 838022, OR 838210, OR 853288, OR 839294, OR 838079, OR 853289, OR 839288, OR 839289, OR 839290.

Other material

(n = 5, CWHc, CUAC). USA • South Carolina • 1 ♂; Pickens Co.; Nine Times Preserve ; 34.946, - 82.806; 10 Nov. 2019; C. W. Harden leg.; On underside of embedded rock, steep rocky ditch, oak-hickory woods; CUAC 000168226 , CWH- 039; CUAC • 2 ♂; Pickens Co.; 3 miles N Clemson ; 34.7252, - 82.8247; 4 July to 19 December 2022; C. W. Harden leg; buried pipe trap, trap 07; CWHc ; • 1 ♀; Pickens Co.; Clemson Experimental Forest .; 34.7431, - 82.8481; 14 Oct. 2019; C. W. Harden leg.; Soilwash-flotation berlese, mesic oak-pine woods, dark sandy soil near stream; CWH- 056, CUAC 000163565 ; CUAC ; • 1 ♂; Pickens Co.; Clemson Experimental Forest, Pike Road, south of inlet ; 34.7116, - 82.828; 1 Jun 2023; C. W. Harden and J. R. LaBonte; under rock; CWHc .

Diagnosis.

Among members of the sinuaticollis group, males are recognized by the modified hind femora (Fig. 12 A ) bowed anteriorly and triangularly produced posteriorly, with a blunt tuberculate projection in the distal 1 / 3. The male genitalia (Fig. 35 G – I ) are similar to those of A. micamicus , but differ in the more rounded shape of the apex of the median lobe and the presence of two well-sclerotized strips along the ostium of the internal sac.

Description.

Habitus (Fig. 34 E ) ABL 1.60–1.89 mm, avg. 1.82 mm, n = 7), males larger than females (male mean ABL = 1.85 n = 5; female mean ABL = 1.74, n = 2). Body moderately depressed dorsoventrally and parallel-sided, relatively elongate (EW / ABL = 0.35–0.37). Integument Irregular isodiametric microsculpture weakly impressed at anterior angles of pronotum, absent from disk, which is slightly rugose; microsculpture distinct on dorsal surface of head except for sides at base of vertex, where it is stretched and weak or entirely absent. Head HW / PW = 0.74–0.80, wider in females than in males. Antennomeres IV – X moniliform, slightly clavate. Labrum slightly emarginate. Frontoclypeal horn present, well developed. Three pairs of supraorbital setae present. Pronotum Variable in length (PL / ABL = 0.23–0.29), broader in males (PW / EW = 0.82) than females (0.78–0.79). Moderately constricted basally (Pbw / PW = 0.75–0.80). Sides slightly sinuate before obtuse hind angles; 3 or 4 basal serrulations present. Elytra Moderately depressed and parallel sided, not markedly elongate (average EL / ABL = 0.55); humeri not sloped; 3 or 4 weakly impressed striae present; without prominent subapical plica. Legs Profemora of males not modified; protarsi of males with first tarsomere enlarged and spinose on inner margin (Fig. 10 B ), with adhesive setae ventrally, second tarsomere unmodified and without adhesive setae. Mesotrochanters of males unmodified. Metafemora of males modified: anterior margin arcuately swollen, posterior margin arcuate proximally towards broad, triangular tuberculate projection in distal 1 / 2; with several long setae along posterior margin (Fig. 12 A ). Female legs unmodified. Abdominal ventrites Unmodified in either sex. Male genitalia Ring sclerite relatively large (RL / ABL = 0.30), subtriangular and slightly asymmetrical in anterior 1 / 2. Median lobe (Fig. 35 I ) arcuate, not strongly twisted from plane of basal lobes; dorsal margin lightly sclerotized for ~ 1 / 2 of its length; ventral margin without setae, deeply arcuate medially, with shallow sinuation just before apex, which is broadly rounded. Internal sac with flagellum not rotated dorsally, relatively small, and abruptly curved past enlarged basal area; sides lightly sclerotized, “ open ” laterally; three groups of long, dark spines present near flagellum on left side of internal sac, variable in number; two sclerotized straps present at ostium, one on left side appearing as a broad, vertical strip and the other ventrally, appearing as a U-shape just beyond apex of flagellum in repose. Right paramere (Fig. 35 H ) partially membranous and enclosed in a feathery sheath, with four setae on apical margin. Left paramere (Fig. 35 G ) slightly quadrate, with apical margin relatively broad; four long setae present ventrally near apex. The male from the Nine Times Preserve has the dorsal margin of the median lobe nearly straight proximally, giving the organ a more triangular shape overall; the shape of the flagellum and additional sclerites are identical to those in males of other populations. Female genitalia. Bursa copulatrix with sclerotized folds. Spermatheca long, gradually enlarged distally, forming a loose coil proximally (Fig. 21 I ). Spermathecal duct short and slightly curved, not coiled.

Distribution.

Known only from Pickens Co., SC, from Nine Times Forest south to the historic course of the Twelve Mile River north of Clemson (Fig. 36 ).

Sympatry.

Collected under rocks in association with A. murrayae and Anillinus micamicus sp. nov. The ranges of Serranillus dunavani and Anillinus sp. “ South Carolina, Waldrop Stone ” overlap with this species, but have so far not been collected in association.

Natural history.

Members of this species are endogean in habit. Specimens examined were collected from beneath rocks, buried pipe traps and soil washing. Hand collected specimens were found in May, September, October, and November. Teneral specimens were hand collected in September, indicating that immature stages occur in the Spring and Summer.

Species status justification.

The combination of external characters is unique within the genus, and the male genitalia are distinct from those of any other described species. The closely related and morphologically similar species A. micamicus sp. nov. shows consistent morphological differences in the characters of the male hind legs and genitalia, and sampled individuals of the two species are reciprocally monophyletic in our molecular phylogeny despite occurring together in syntopy under the same rocks. This provides strong support for the hypothesis that the two species are reproductively isolated.

Derivation of species name.

A noun in apposition, named for the mineral mica , which is conspicuous in the soils at most localities where this species has been collected.

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 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 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 34. Dorsal habitus of Anillinus species in the elongatus group and sinuaticollis group A Anillinus montrex B Anillinus arenicollis C Anillinus choestoea sp. nov. D Anillinus seneca sp. nov. E Anillinus mica sp. nov. F Anillinus micamicus sp. nov. Abdomens removed in B – F for DNA extraction. Scale bars: 1 mm.

Figure 35. Male genitalia of Anillinus species in the sinuaticollis group A – C Anillinus choestoea sp. nov. D – F Anillinus seneca sp. nov. G – I Anillinus mica sp. nov. J – L Anillinus micamicus sp. nov. Median lobe of aedeagus in right dorsolateral (C, I, L) or right lateral (F) aspect. Left parameres (A, D, G, J) and right parameres (B, E, H, K) in left and right lateral aspects, respectively. Scale bar: 0.1 mm.

Figure 36. Distribution map of sinuaticollis - group species of Anillinus in South Carolina. Data are from Harden (2024). Blue lines show modern and pre-inundation courses of the Tugaloo River and Seneca River and their major tributaries. The confluence of the two rivers becomes the Savannah River.