Systematics and biogeography of Appalachian Anillini, and a taxonomic review of the species of South Carolina (Coleoptera, Carabidae, Trechinae, Anillini) Author Harden, Curt W. 0000-0002-5226-8725 Department of Plant and Environmental Sciences, Clemson University, 277 Poole Agricultural Center, Clemson, South Carolina, 29634, USA Author Caterino, Michael S. 0000-0002-2597-5707 Department of Plant and Environmental Sciences, Clemson University, 277 Poole Agricultural Center, Clemson, South Carolina, 29634, USA text ZooKeys 2024 2024-08-08 1209 69 197 journal article 10.3897/zookeys.1209.125897 CEE78803-61EB-40CC-8D63-46142E6383A1 Serranillus dunavani (Jeannel, 1963) Figs 2 A , 10 G , 11 A, C , 13 A , 14 B, D , 15 A – E , 16 , 17 A , 21 S , 25 A – C Anillinus dunavani Jeannel, 1963: 76; Barr 1995: 245 . Serranillus dunavani (Jeannel): Sokolov et al. 2004: 188 . Material examined. Holotype male ( USNM ) : point mounted, not dissected, labeled: “ Rocky Bottom , Pickens Co. S. C. , 25 Aug. 1932 , D. Dunavan Co. ” “ Sassafras Mtn. 3,500 ft ” “ In leaf mold ” “ TYPE [red paper] ” “ Type No. 69542 USNM [red paper] ” “ Anillinus dunavani n. sp. R. Jeannel det. 19 / ”. 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. SEM micrographs of mouthparts of Appalachian Anillini genera A mentum of Serranillus dunavani (Jeannel) B mentum of Anillinus choestoea C mandibles of S. dunavani , dorsal aspect D mandibles of A. choestoea , dorsal aspect. Scale bars: 0.1 mm. Other material ( n = 177, CMNH , CUAC , CWHc , NCSU , OSUC , USNM ). USA • North Carolina • 1 ♂ ; Clay Co . ; Nantahala National Forest , Shooting Creek Bald ; 35.0679 , - 83.6466 ; 11 May 2020 ; C. W. Harden and M. S. Caterino leg.; CWH- 381, CUAC 000066851 ; • 1 ♂ , 4 ♀ ; same data as previous; CWHc ; • 1 ♂ , 1 ♀ ; Clay Co. ; Nantahala National Forest , Shooting Creek Bald ; 35.0681 , - 83.6464 ; 11 May 2020 ; C. W. Harden leg.; CWHc ; • 2 ♀ ; Henderson Co. ; 0.3 miles southwest of Bat Cave ; 35.4470 , - 82.2910 ; 22 July 1967 ; S. Peck and A. Fiske leg.; CMNH ; • 2 ♂ , 1 ♀ ; Macon Co. ; 1 mile northwest of Highlands ; 35.061 , - 83.217 ; 16 Aug. 1981 ; Q. D. Wheeler leg.; OSUC ; • 1 ♂ , 1 ♀ ; Macon Co. ; Nantahala National Forest , Cowee Bald ; 35.3269 , - 83.3350 ; 15 Sep. 2020 ; • 1 ♂ ; Montgomery Co. ; Uwarrie National Forest , 2 miles south of Eldorado , Route 109 ; 35.4340 , - 80.0190 ; 27 Aug. 1990 ; W. Reeves leg.; LSAM 0153884 ; NCSU ; • 2 ♂ , 1 ♀ ; Polk Co. ; Green River Game Lands ; 35.2940 , - 82.2610 ; 18 Mar. 2018 ; M. S. Caterino leg.; CUAC 000107846 , CUAC 000107847 , CUAC 000107856 ; • 1 ♀ ; Polk Co. ; Lower Bradley Falls Trail ; 35.3580 , - 82.2878 ; 19 Mar. 2018 ; M. S. Caterino leg.; CUAC 000107772 ; • 2 ♂ , 2 ♀ ; Polk Co. ; Melrose Falls ; 35.2217 , - 82.2985 ; 10 Aug. 2021 ; M. S. Caterino leg. ; • 1 unsexed; Swain Co. ; Great Smoky Mountains National Park , Deep Creek area , loop trail at Sunkota Ridge trail ; 35.4750 , - 83.4200 ; 25 Jul. 2002 ; C. E. Carlton leg.; NCSU _ ENT 00293747 ; NCSU ; • 1 unsexed; Swain Co. ; Great Smoky Mountains National Park , Lakeshore Trail ; 35.4520 , - 83.5410 ; 18 Jul. 2018 ; A. K. Tishechkin leg.; NCSU _ ENT 00293744 ; NCSU ; • 1 ♂ , 1 ♀ ; Transylvania Co. ; Sassafras Mountain ; 35.0656 , - 82.7776 ; 11 Jun. 2020 ; CUAC ; • 1 ♂ , 2 ♀ ; Transylvania Co. ; 1 mile south of Rosman , NC 178; 35.1290 , - 82.8230 ; [no date]; J. F. and S. Cornell leg.; NCSU ; • South Carolina ; • 1 ♂ ; Abbeville Co. ; Sumter National Forest , Long Cane Creek ; 34.1345 , - 82.3230 ; 4 Feb. 2022 ; M. Ferro leg.; CWHc ; • 1 ♀ ; Abbeville Co. ; Sumter National Forest , off Cedar Springs Road ; 34.1086 , - 82.3390 ; 25 Sep. 2020 ; C. W. Harden leg.; CWH- 267, CUAC 000066830 ; • 1 ♂ ; Greenville Co. ; Highway 97 , River Falls Lodge ; 35.1221 , - 82.5405 ; 18 Mar. 2017 ; M. Ferro leg.; CUAC 000049982 ; • 1 ♂ , 11 ♀ ; Greenville Co. ; Ashmore Heritage Preserve ; 35.0867 , - 82.5788 ; 14 Mar. 2020 ; C. W. Harden and L. M. Thompson leg.; CWHc ; • 1 ♂ ; Greenville Co. ; Ashmore Heritage Preserve ; 35.0933 , - 82.5930 ; 29 Jun. 2015 ; S. Myers leg.; MSC- 2463, CUAC 000185783 ; • 1 ♂ ; Greenville Co. ; Chestnut Ridge Heritage Preserve ; 35.1506 , - 82.2779 ; 8 Apr. 2018 ; M. Caterino and L. Vasquez leg.; CUAC 000108123 ; • 1 ♀ ; Greenville Co. ; Chestnut Ridge Heritage Preserve ; 35.1507 , - 82.2821 ; 20 Oct. 2021 ; C. W. Harden leg.; CWHc ; • n = 11; Kershaw Co. ; English Swamp , Wateree Floodland Memorial Forest ; 34.0911 , - 80.6578 ; 27 Feb. 2010 ; J. F. Cornell , S. Cornell and B. Gregory leg.; NCSU ; • 2 ♀ ; Oconee Co. ; Coon Branch Natural Area ; 35.0200 , - 83.0000 ; 21 Jun. 2018 ; D. Chandler leg.; CUAC 000109930 and CUAC 000109964 ; • 4 ♂ , 2 ♀ ; Oconee Co. ; Coon Branch Natural Area ; 35.0170 , - 82.9970 ; 18 Oct. 2020 ; C. W. Harden leg.; CWHc ; • 1 ♂ ; Oconee Co. ; Devils Fork State Park ; 34.9390 , - 82.8798 ; 26 Apr. 2015 ; S. Myers leg.; MSC- 2454, CUAC 000185791 ; • 1 ♂ , 2 ♀ ; Oconee Co. ; East Fork [ Chattooga River ]; 34.9843 , - 83.0981 ; 4 May 2015 ; M. S. Caterino and S. Myers leg.; CUAC 000110650 , CUAC 000110651 , CUAC 000110653 ; • 1 ♀ ; Oconee Co. ; East Fork trail , Ellicott Rock Wilderness ; 34.9913 , - 83.0980 ; 29 Jun. 2015 ; S. Myers leg.; SSM 250, CUAC 000185792 ; • 1 unsexed; Oconee Co. ; East Fork trail ; 34.9838 , - 83.0979 ; 4 May. 2015 ; S. Myers leg.; SSM 415, CUAC 000185793 ; • 2 ♂ , 1 ♀ ; Oconee Co. ; Hill above parking lot near Stumphouse Tunnel ; 34.8097 , - 83.1235 ; 5 Jul. 2019 ; C. W. Harden leg.; CWHc ; • 1 ♂ ; Oconee Co. ; Martin Creek Landing ; 34.6389 , - 82.8663 ; 26 May 2022 ; C. W. Harden leg.; CWH- 448, CUAC 000066843 ; • 1 ♀ ; Oconee Co. ; Oconee State Park , 6.7 miles north-northwest of Walhalla ; 34.8690 , - 83.1050 ; 1 Jul. 1983 ; J. Pakaluk leg.; NCSU ; • 2 ♂ , 5 ♀ ; Oconee Co. ; Oconee State Park ; 34.8690 , - 83.1050 ; 21 Jul. 1967 ; S. Peck and A. Fiske leg.; CMNH ; • 1 ♂ , 1 ♀ ; Oconee Co. ; Sumter National Forest , Yellow Branch Falls ; 34.8067 , - 83.1283 ; 12 Oct. 2017 ; M. S. Caterino leg.; CUAC 000107624 and CUAC 000107625 ; • 3 ♂ , 1 ♀ ; Oconee Co. ; Sumter National Forest , Forest service road 725 ; 34.8395 , - 83.1677 ; 13 Sep. 2020 ; A. Deczynski leg.; CWHc ; • 1 ♂ ; Oconee Co. ; Sumter National Forest , Indian Camp Branch ; 34.9898 , - 83.0724 ; 4 May 2015 ; M. S. Caterino and S. Myers leg.; CUAC 000010470 ; • 2 ♂ , 2 ♀ ; Oconee Co. ; Sumter National Forest , Tater Hill trail; 34.9612 , - 83.0116 ; 24 Feb. 2018 ; M. Caterino leg.; CUAC ; • 1 ♂ ; Pickens Co. ; Chimneytop Gap ; 35.0620 , - 82.8008 ; 10 Jun. 2015 ; S. Myers leg.; MSC- 2457, CUAC 000185780 ; • 1 ♂ ; Pickens Co. ; Chimneytop Gap ; 35.0632 , - 82.7964 ; 23 Mar. 2023 ; C. W. Harden leg.; CWHc ; • 1 ♂ , 1 ♀ ; Pickens Co. ; Chimneytop Gap ; 35.0630 , - 82.7968 ; 23 Mar. 2023 ; C. W. Harden leg.; CWHc ; • 1 ♂ ; Pickens Co. ; Clemson Experimental Forest ; 34.7384 , - 82.8436 ; 20 Feb. 2023 ; E. Recuero leg.; CWHc ; • 1 ♀ ; Pickens Co. ; Clemson Experimental Forest , Waldrop Stone area ; 34.7390 , - 82.8216 ; 2 Feb. 2021 ; C. W. Harden leg.; CWHc ; • 1 ♂ ; Pickens Co. ; Clemson Experimental Forest , Wildcat Creek ; 34.7561 , - 82.8551 ; 20 Jun. 2018 ; C. W. Harden leg.; CWHc ; • 3 ♂ , 1 ♀ ; Pickens Co. ; Clemson ; 34.6820 , - 82.8330 ; 4 Oct. 1966 ; J. A. Payne leg.; CMNH ; • 1 ♀ ; Pickens Co. ; Clemson ; 34.6820 , - 82.8330 ; 19 May 1962 ; J. A. Payne leg.; CMNH ; • 4 ♂ , 2 ♀ ; Pickens Co. ; Clemson ; 34.6820 , - 82.8330 ; 20 Jun. 1962 ; J. A. Payne leg.; CMNH ; • 1 ♂ ; Pickens Co. ; Eastatoe Creek Heritage Preserve ; 35.1577 , - 82.4910 ; 30 Mar. 2015 ; S. Myers leg.; MSC- 2455, CUAC 000185778 ; • 1 ♂ ; Pickens Co. ; Eastatoe Creek Heritage Preserve ; [incorrect coordinates]; 30 Mar. 2015 ; S. Myers leg.; MSC- 2461, CUAC 000185781 ; • 1 ♀ ; Pickens Co. ; Eastatoe Heritage Preserve ; 35.0462 , - 82.8178 ; 6 Apr. 2014 ; M. Caterino and K. Caterino leg.; CUAC 000002709 ; • 2 ♂ , 1 ♀ ; Pickens Co. ; near Chimneytop Gap ; 35.0639 , - 82.7996 ; 11 Sep. 2019 ; M. S. Caterino leg.; CUAC ; • 4 ♀ ; Pickens Co. ; Nine Times Preserve , 34.9460 , - 82.8023 ; 31 Sep. 2015 ; S. Myers leg.; SSM 184 to SSM 187, CUAC 000185770 , CUAC 000185773 to CUAC 000185775 ; • 2 ♂ , 1 ♀ ; Pickens Co. ; Nine Times Preserve ; 34.9465 , - 82.7992 ; 27 Mar. 2015 ; S. Myers leg.; SSM 188 to SSM 190; CUAC 000185769 , CUAC 000185776 , CUAC 000185777 ; • 3 ♂ , 1 ♀ ; Pickens Co. ; Nine Times Preserve ; 34.9464 , - 82.8027 ; 27 Mar. 2015 ; M. S. Caterino and S. Myers leg.; CUAC 000173553 , CUAC 000173556 , CUAC 000173557 , CUAC 000173559 ; • 1 larva , 1 ♂ ; Pickens Co. ; Sassafras Mountain ; 35.0640 , - 82.7767 ; 20 Oct. 2020 ; F. Etzler and P. Wooden leg.; CWH- 311 and CWH- 312, CUAC 000185899 and CUAC 000066867 ; • 8 ♂ , 9 ♀ ; same data as previous; CUAC ; • 2 ♀ ; Pickens Co. ; Sassafras Mountain ; 35.0645 , - 82.7774 ; 10 Jun. 2015 ; S. Myers leg.; SSM 67 and SSM 68, CUAC 000185771 and CUAC 000185785 ; • 2 ♀ ; Pickens Co. ; Sassafras Mountain ; 35.0634 , - 82.7760 ; 10 Jun. 2015 ; S. Myers leg.; SSM 69 and SSM 70, CUAC 000185784 and CUAC 000185790 ; • 3 ♂ , 1 ♀ ; Pickens Co. ; Sassafras Mountain ; 35.0579 , - 82.7705 ; 10 Jun. 2015 ; S. Myers leg.; SSM 71 to SSM 74, CUAC 000185786 to CUAC 000185789 ; • 3 ♂ , 5 ♀ ; Pickens Co. ; Sassafras Mountain ; 35.0647 , - 82.7774 ; 11 Jun. 2020 ; F. Etzler leg.; CUAC ; • 5 ♂ , 4 ♀ ; Pickens Co. ; Sassafras Mountain ; 35.0650 , - 82.7750 ; 21 Jul. 1967 ; S. Peck and A. Fiske leg.; CMNH . SEM micrographs of male left metafemora of Anillinus species. Scale bar: 0.1 mm. Dorsal habitus of late-instar larvae A Serranillus dunavani (Jeannel) B Anillinus jancae . Scale bars: 1 mm. 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. GenBank accession numbers for specimens from Sassafras Mountain: OR 839609 , OR 839610 , OR 839814 , OR 839815 , OR 839816 , OR 839817 , OR 839818 , OR 838294 , OR 853396 , OR 839363 , OR 839819 , OR 837946 , OR 838111 , OR 838295 . Literature records. Besides the type locality, no specific records have been previously published for S. dunavani . Sokolov and Carlton (2010) cite a molecular voucher of S. dunavani from “ GSMNP, Twentymile Trail. ” Sokolov and Carlton (2008) state that they studied material of S. dunavani only from “ GSMNP ”. Their fig. 7 shows the range of S. dunavani covering over 30 counties in Tennessee , North Carolina , and South Carolina , citing “ original data ” as the source for the figure. However, these data are not apparently available and we have not seen any material of S. dunavani from Tennessee nor from most of the North Carolina counties that are shaded in that figure. Diagnosis. This is the smallest Serranillus species , with most specimens having an ABL of 2 mm or less. The median lobe of the aedeagus is distinctive in being elongate with a recurved apex and lacking large, sclerotized spines in the internal sac (Fig. 15 E ). Serranillus dunavani (Jeannel) A habitus B last abdominal ventrite, ventral aspect C left paramere, left lateral aspect D right paramere, right lateral aspect E median lobe, right dorsolateral aspect. Scale bars: 1 mm ( A ); 0.1 mm ( B – E ). Redescription. Habitus Small for the genus ( ABL = 1.79–2.03 mm ), convex and ovoid, robust (Fig. 15 A ). Integument Microsculpture indistinct on dorsal surfaces of head and pronotum, giving a shiny appearance. Head Relatively large ( HW / PW = 0.73–0.75), antennomeres IV – X moniliform, slightly clavate. Frontoclypeal horn present, conspicuous. Ocular tubercles variable, present at least as a slight bulge, sometimes developed as a conspicuous hornlike projection below lateral carina. In some specimens, dorsad to lateral carina above tubercle on either side is a pale circular mark, more evident in teneral individuals. Two pairs of supraorbital setae present. Mentum with median pair of setae on tooth. Pronotum Relatively short ( PL / ABL = 0.23–0.24) and broad ( PW / EW = 0.83–0.84), moderately constricted posteriorly ( PbW / PW = 0.72–0.78), sides evenly convergent to obtuse posterior angles. Elytra Moderately convex and ovoid, relatively broad ( EW / ABL = 0.37–0.39), with weak traces of four striae. Legs Male protarsomeres 1 and 2 expanded and dentate on inner margin, with thick white adhesive setae ventrally (Fig. 10 G ). Male metatrochanters and posterior face of metafemora with coarse papillate microsculpture, metafemora slightly swollen. Female legs unmodified. Abdominal ventrites Last abdominal ventrite in males modified, apex protruding and bearing three blunt lobes, the inner slightly more prominent than the outer two (Fig. 15 B ). Male genitalia . Ring sclerite relatively large ( RL / ABL = 0.30), narrow and subtriangular, with lightly-sclerotized expansions laterally; apex narrow and bent ventrally. Median lobe (Fig. 15 E ) complex, strongly asymmetrical (Fig. 17 A ), twisted dorsally from plane of basal lobes, heavily sclerotized on all faces. In strict right lateral aspect appearing as a blunt club-shaped organ without discernible apex. In dorsal aspect, nearly straight for basal 2 / 3, without basal bend, with ventral margin abruptly curved ventrally with apical sinuation. Left side of median lobe with a small, deep channel at base, with associated carina running obliquely for ~ 1 / 2 the length of median lobe. Apex variable: in topotype specimens, it is small, rounded, and deflected ventrally; in Clay Co., North Carolina , it is larger and more angularly produced ventrally. Small setae present on ventral surface of median lobe in some individuals. Basal lobes strongly asymmetrical, right lobe reduced to a narrow strap, left lobe large and cup-shaped. Flagellum long, thick at base and filamentous for most of its length beyond, evenly curved. Internal sac covered in small scales. Short scroll-like sclerite present at left side of ostium (behind flagellum in right lateral aspect), rolled into a tube; in lateral view appearing as a lightly sclerotized plate, rolled shape apparent in more posterior or anterior aspects. Right paramere minute and asetose, difficult to see in some aspects (Fig. 15 D ). Left paramere conchoidal, with enlarged base that is subequal in size to the rest of the paramere; ventral margin asetose (Fig. 15 C ). Female genitalia . Spermatheca small, curved, gradually enlarged apically, with swollen base (Fig. 21 S ). Spermathecal gland present, elongate. Spermathecal duct long and heavily coiled. Distribution. This is the most widely distributed anilline in South Carolina (Fig. 16 ), with specimens known from Oconee Co. to Greenville Co. along the North Carolina border, south at least to Abbeville Co. and Kershaw Co. in the outer Piedmont and Sandhills physiographic regions, respectively. In North Carolina , the species has been found as far east as Montgomery Co., west to Clay Co., and north to Swain Co. Distribution map of Serranillus species that occur in South Carolina. The locality for Serranillus sp. “ South Carolina, Coon Branch ” is the same as the single South Carolina S. jeanneli occurrence, and is not shown. Data are from Harden (2024) . Multiple aspects of median lobes of South Carolina Serranillus species , progressively rotated dorsally from right lateral aspect (upper left) to left lateroventral aspect (lower right) A Serranillus dunavani B Serranillus jeanneli C Serranillus monadnock , holotype (Kings Mountain, SC) D Serranillus monadnock , paratype (Little Mountain, SC). Photographs not to same scale. Sympatry. In South Carolina , this species has been collected in association with the following eight anilline species: A. castaneus sp. nov. (Greenville Co.), A. sp. “ South Carolina , Chestnut Ridge ” (Greenville Co.), A. cherokee (Oconee Co.) , A. murrayae (Greenville Co., Pickens Co., Oconee Co.) , A. dentatus sp. nov. (Abbeville Co.), Anillinus sp. “ South Carolina , Wateree ”, S. jeanneli , and S. sp. “ South Carolina , Coon Branch ”. Natural history. Specimens have been obtained by Berlese extraction of sifted soil, litter, and coarse woody debris, and have been hand collected from under embedded rocks, bear dung, and pig carcasses. Two late instar larvae were collected in a sifted litter sample taken near the summit of Sassafras Mountain in October, in a mixed forest of pines and hardwoods. In a study of the soil and litter fauna of several South Carolina Piedmont forests, DuRant and Fox (1966) reported “ Anillus carolinae (Horn) ” as the most abundant species of Coleoptera captured in a single litter sample; these specimens were probably S. dunavani .