Anillinus dentatus, 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 dentatus sp. nov.

Figs 2 C , 12 F , 21 F , 22 A – C , 23 A – C , 24 F , 25 C , 43 B

Type material.

Holotype male ( USNM): point mounted, with abdominal ventrites and genitalia in Euparal on microslide pinned beneath specimen. Original labels: “ USA: SC, Abbeville Co. Sumter NF Long Cane Crk at end of FS rd 530 . 34.1133, - 82.3300. 5. ii. 2022. CW Harden & K Ivanov. Beneath embedded mossy rock in waterlogged soil. ” “ [QR Code] CLEMSON-ENT CUAC 000163530 ” “ Harden DNA Voucher CWH- 420 Anillinus ‘ dentate’ M Ext. 6 - February- 2022 [green-bordered cardstock] ” “ HOLOTYPE Anillinus dentatus Harden & Caterino [orange cardstock] ”

GenBank: OR 853208, OR 839248.

Paratypes (n = 21; CMNH, NCSU, VMNH, OSAC, CUAC). USA • South Carolina • 1 ♂; same data as holotype; CUAC 000163531 , CWH- 421; CUAC ; • 5 ♀; same data as holotype; CUAC 000163539 to CUAC 000163541 , CWH- 422 to CWH- 428; CUAC ; • 1 ♀; Abbeville Co.; Sumter National Forest, near Secession St bridge ; 34.134, - 82.324; 25 Jan. 2020; C. W. Harden leg.; Underside of embedded rock; CUAC 000163526 , CWH- 113; CUAC ; • 1 ♂; Abbeville Co.; Sumter National Forest ; 34.1345, - 82.3230; 5 Feb. 2022; M. Ferro leg.; Sift subcort CWD; CUAC 000168286 ; • 2 ♂; Abbeville Co.; Sumter National Forest, near Secession Street bridge ; 34.1366, - 82.3252; 5. February. 2022; CW Harden leg.; beneath embedded mossy rock CUAC 000163524 and CUAC 000163529 , CWH- 417 and CWH- 418; CUAC ; • 1 ♀; same data as previous; CUAC 000163542 , CWH- 419; CUAC ; • 1 ♂; Abbeville Co.; Sumter National Forest, Long Cane Creek ; 34.13519, - 82.32503; 12 Jan. 2020; C. W. Harden & L. M. Thompson leg.; Soilwash flotation Berlese, clay-rich soil from ferny hill above floodplain; CUAC 000163525 , CWH- 067; CUAC ; • 2 ♂; Abbeville Co.; Sumter National Forest, Long Cane Creek ; 34.1350, - 82.3239; 15 Mar. – 16 Jul. 2020; C. W. Harden leg.; Buried pipe trap baited with cheese, deep clay soil, LCC- 05-0716; CUAC 000163527 , CUAC 000163528 ; • 7 ♀; same data as for proceeding; CUAC 000163532 to CUAC 000163536 ; • 3 ♂, 3 ♀; Abbeville Co.; Sumter National Forest, Long Cane Creek area ; 34.1370, - 82.3226; 5 May 2023; C. W. Harden leg.; Under small rocks in ditch.; OSAC, VMNH, ADGc .

GenBank accession numbers for paratypes: OR 853105, OR 853210, OR 839250, OR 839420, OR 837801, OR 837962, OR 838150, OR 838114, OR 853209, OR 839249, OR 839458, OR 839458, OR 837972, OR 839247, OR 839246.

Other material

(n = 5). USA • South Carolina • 2 ♂, 1 ♀; Abbeville Co.; Sumter National Forest, Long Cane Creek area ; 34.1370, - 82.3226; 5 May 2023; C. W. Harden leg.; Under small rocks in ditch.; CWHc ; • 1 ♂; Abbeville Co.; Sumter National Forest, Long Cane Cr near Secession St bridge; 34.134, - 82.324; 25 Jan. 2020; C. W. Harden leg.; Underside of embedded rock; CWHc ; • 1 ♀; Abbeville Co.; Sumter National Forest, Long Cane Creek ; 34.1349, - 82.3241; 21 Mar. 2021; C. W. Harden and L. M. Thompson leg.; On underside of rock; CWHc .

Diagnosis.

Males of this species are easily recognized by the dentate mesotrochanters, a unique character in the genus. Females are most likely to be confused with A. jancae sp. nov., described below. Females of A. jancae are larger (ABL = 1.88 mm or greater), and the spermatheca is smaller (Fig. 21 N ).

Description.

Habitus Body small (ABL = 1.49–1.69 mm, average = 1.59 ± 0.05 mm), flattened dorsoventrally and relatively narrow (avg. EW / ABL = 0.35) (Fig. 22 A ). Average ABL of males (1.63 mm, n = 5) greater than females (1.57 mm, n = 11). Integument Dorsal surfaces of forebody completely microsculptured with coarse mesh of irregular isodiametric cells. Head HW / PW = 0.73–0.83. Antennomeres IV – X moniliform, slightly clavate. Frontoclypeal horn absent or barely suggested, inconspicuous in lateral view (Fig. 22 C ). Three pairs of supraorbital setae present. Mentum with median pair of setae posterior to the bead of the subtriangular mentum tooth, which is relatively large and blunt. Pronotum Relatively short (PL / ABL = 0.22–0.23), width variable (PW / EW 0.78–0.85); subcordate, strongly to moderately narrowed basally (bPW / PW = 0.70–0.80), anterior angles slightly prominent; sides typically slightly sinuate or evenly convergent towards obtuse hind angles; rarely, the sides are abruptly parallel sided just anterior to nearly right hind angles; pronotal sides with 3 or 4 basal serrulations. Elytra Parallel sided and flat, relative length variable (EL / ABL = 0.53–0.58); margins strongly serrulate; traces of five striae evident on disc of each elytron (Fig. 22 A ); without prominent subapical plica; fused inner margins slightly carinate at apex. Legs Profemora of males unmodified; protarsomere 1 of males moderately expanded, with inner margin spinose, bearing adhesive setae ventrally; protarsomere 2 not expanded, without adhesive setae ventrally (Fig. 10 F ). Mesotrochanters of males with spinose projection ventrally (Fig. 22 B ); mesotrochanters of females either evenly rounded or with blunt projection ventrally. Metafemora of males with prominent triangular toothlike projection on posterior margin (Fig. 12 F ). Tarsomeres of middle and hind legs of both sexes short and broad. Abdominal ventrites Unmodified in either sex. Male genitalia Ring sclerite (Fig. 24 F ) ~ 1 / 4 the total body length (RL / ABL = 0.25), sides asymmetric, margin of narrowed end slightly thickened but not deflexed in lateral view; sides without flattened lateral expansions medially. Median lobe of aedeagus (Fig. 23 C ) narrow and moderately curved, not twisted from plane of basal lobes; dorsal margin largely unsclerotized; ventral margin bladelike toward apex; apex rounded and slightly bent ventrally. Internal sac with flagellum elongate and filamentous distally, in repose coiled around itself in a circle and situated in basal 1 / 3 of median lobe. Right paramere (Fig. 23 B ) small and narrow, bearing four long setae on apex. Left paramere (Fig. 23 A ) with three or four small pores on ventral margin near apex, without setae. Female genitalia Spermatheca elongate, abruptly expanded distally; stem slightly sinuate, straight at proximal juncture with duct; duct long, tightly coiled in corkscrew pattern in situ (shown partially distended in Fig. 21 F ).

Distribution.

Known from two localities within a small area of Sumter National Forest in Abbeville Co., SC along Long Cane Creek (Figs 25 C , 43 B )

Sympatry.

Members of this species have been collected under rocks in association with adults of A. chandleri and a larva of A. jancae sp. nov., and with S. dunavani in a sample of sifted coarse woody debris. Adults of A. jancae sp. nov. have been collected nearby.

Natural history.

Members of this species are endogean, inhabiting deep clay soils in mesic deciduous woods. Specimens have been found on the underside of embedded rocks in January, February, March, and May, and were collected in buried pipe traps operating from March to July but not January to March. One male specimen was collected in early February from Berlese extraction of sifted coarse woody debris. The sifted material was primarily subcortical, but some may have possibly come from the underside of logs that were on the soil surface (M. Ferro, pers. comm., February 2022). This material was collected near Long Cane Creek following a period of heavy rainfall, when the water level was above normal. Most of the specimens examined have had Laboulbeniales fungi present on the dorsum of the elytra, usually near the apex.

Species status justification.

The combination of morphological characters is unique within the genus, and DNA sequence data indicate that A. dentatus is distantly related to all other Anillinus . No known species of Anillinus possess characters suggesting a close relationship to A. dentatus .

Derivation of species name.

A male adjective referring to the triangular projections on the male metafemora, from the Latin for “ toothed. ”

Notes.

The long, coiled form of the flagellum in A. dentatus is also seen in Anillinus barri Sokolov & Carlton , Anillinus erwini Sokolov & Carlton , Anillinus folkertsioides Sokolov , and Anillinus inexpectatus Sokolov. However , with the exception of A. folkertsioides , all of these species differ from A. dentatus in most external characters. Members of A. folkertsioides and a possibly undescribed sister species from Fort Payne, Alabama are similar to A. dentatus in having fully developed microsculpture on the forebody and a depressed habitus, but differ from A. dentatus in having numerous long hairlike setae on the right paramere and male legs without modifications. Two character-states seen in members of A. dentatus are unique among known members of the genus: the nearly straight stem of the spermatheca and the spinose mesotrochanters of males.

‘ albrittonorum group’

Figures

Figure 2. Habitus of eastern Nearctic Anillini. A Serranillus dunavani (South Carolina, Oconee Co.) B Anillinus moseleyae (Tennessee, Sevier Co.) C Anillinus dentatus (South Carolina, Abbeville Co.) D Serranillus jeanneli (South Carolina, Oconee Co.) E Anillinus folkertsioides (Alabama, Blount Co.) F Anillinus pecki (North Carolina, Avery Co.) G Anillinus sp. “ Georgia, Barnes Creek sp. 1 ” H Anillinus montrex (South Carolina, York Co.) I Anillinus sp. “ Tennessee, Kings Saltpeter Cave ” J Anillinus virginiae (Virginia, Bath Co.) K Anillinus erwini (Virginia, Grayson Co.) L Serranillus sp. (North Carolina, Graham Co.) M Anillinus sp. “ South Carolina, Waldrop Stone ” (South Carolina, Pickens Co.) N Anillinus sp. “ Alabama, Horseshoe Cave sp. 1 ” O Anillinus loweae (North Carolina, Haywood Co.) P Anillinus sp. “ Kentucky, Hestand sp. 1 ” Q Anillinus indianae (Indiana, Lawrence Co.) R Anillinus cf. barberi (West Virginia, Greenbrier Co.) S Anillinus hirsutus (Alabama, Madison Co.). Scale bar: 1 mm.

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 22. Anillinus dentatus A dorsal habitus B SEM micrograph of metaventrite, ventral aspect (white arrows = mesotrochanter spines) C head, left lateral aspect (black arrow = absence of frontoclypeal horn). Scale bars: 1 mm (A); 0.1 mm (B).

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 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).