Apheloria uwharrie Marek, Means, Hennen & Tingley, 2025

Marek, Paul E., Means, Jackson C., Hennen, Derek A. & Tingley, Carol, 2025, Revision of the millipede genus Apheloria Chamberlin, 1921 (Polydesmida, Xystodesmidae, Apheloriini), Zootaxa 5701 (3), pp. 315-350 : 328-331

Apheloria uwharrie Marek, Means, Hennen & Tingley , new species

Vernacular name: “The Uwharrie cherry millipede”

Figs 9 , 15

Apheloria “species”— Shelley 2000: 193.

Apheloria “Myrtle Beach”— Means et al. 2021b: 7.

Material examined:Type specimens— ♂ holotype ( VTEC, MPE03855 ), 1 ♀ paratype ( VTEC, MPE03857 ), 2 ♂, 2 ♀ paratypes ( VMNH, MPE03856 , 3858 ; VTEC, 3859 , 3875 ) from South Carolina, Horry Co., Myrtle Beach: Forested area near the intersection of Robert M Grissom Parkway and 29th Avenue North ( 33.711699°N, - 78.882573°W, Elev. 8 m), 30 March 2018 (Coll: G. Schiermeyer) ( vidi) . Non type specimens— North Carolina, Stanly Co. , 2 ♂ ( VTEC, MPE01977 , MPE01979 ), 1 ♀ ( VMNH, MPE01978 ) Uwharrie Mountains , Morrow Mountain State Park , forest by old cabin & small parking lot ( 35.371667°N, - 80.096111°W, Elev. 140 m), 7 August 2016, 16:00 ( Coll: J. Means). Randolph Co. , 3 ♂ ( VTEC, MPE05400 , 5402 , 5403 ) Uwharrie National Forest , Birkhead Mountain Trail, near Tot Hill ( 35.6307°N, - 79.9064°W, Elev. 240 m), 27 August 2022 ( Coll: C. Tingley). Montgomery Co. , 1 ♂ ( VTEC, MPE05401 ) Uwharrie National Forest , Wood Run Trailhead Parking ( 35.3108°N, - 80.0433°W, Elev. 167 m), 22 August 2022 ( Coll: C. Tingley). Material examined are archived in the Virginia Tech Data Repository at: https://doi.org/10.7294/29829209

Diagnosis: Apheloria uwharrie is distinct from other apheloriine species based on the following combination of characters: Color. Tergites with three yellow (Uwharrie Mountains, Fig. 15A ) or red spots (Myrtle Beach) or red stripes (Myrtle Beach, Fig. 15B ). ♂ Gonopods. Gonopodal acropodite smoothly circular, O-shaped ( Fig. 9A )— without elbow as in A. virginiensis ( Fig. 7A , elb). Not smoothly oval-shaped as in Rudiloria , nor D-shaped as in Sigmoria . Acropodite narrow, about one-half width of tibia on leg pair 9; of uniform width throughout. Acropodite tapered to L-shaped acuminate apex and abruptly twisted ( Fig. 9B )—not gradually tapered to curved J-shaped apex as in A. polychroma ( Fig. 8B ). Acropodite shaft without cingulum nor preapical teeth nor projections as in Appalachioria , Brachoria . Prefemur with a long, scythe-like prefemoral process ( Fig. 9B , pfp)—not short, scythe-like as in A. polychroma ( Fig. 8B , pfp). With acute angle at prefemur-acropodite junction ( Fig. 9B ), not with distinct bend tubercle at junction as in A. corrugata , A. virginiensis , A. montana ( Fig. 4B , 7B , 13B , bt).

Diagnosis notes: A millipede with red spots or stripes and circular acropodites occurring in Myrtle Beach, South Carolina, unequivocally diagnoses A. uwharrie from all other species. Similarly, a millipede with yellow spots and circular acropodites occurring in the Uwharrie Mountains unequivocally diagnoses A. uwharrie from all other species. However, A. virginiensis has similar coloration and occurs in the coastal plain and piedmont of North Carolina and Virginia, and northeast of A. uwharrie . Although there are no A. virginiensis with red stripes, there are A. virginiensis with yellow or red spots.

Description. Based on holotype ( ♂) MPE03855

Measurements: BL = 49.18, CW = 6.70, IW = 4.25, ISW = 1.27, B11W = 9.30. Head: Antenna length— extendable posteriorly to anterior margin of 3rd tergite; relative antennomere lengths 4>2>3>5>6>7>1. Antenna with 4 sensillum types; all sensillum shafts smooth, without barbules. Four apical cones ( AS) in square pattern on 8th antennomere. Chaetiform sensilla (CS) on antennomeres 2–7.Antennomere 1 smooth without CS; antennomeres 2–4 with sparse CS vestiture; antennomeres 5–7 with dense CS vestiture. Trichoid sensilla (TS) on antennomeres 1– 7; encircling apical rims. Spiniform basiconic sensilla (Bs 2) in clusters of 15 on apicodorsal surface of antennomeres 5, 6; Bs 2 1/10 length of CS. Tergites: Collum with straight cephalic edge, tapering laterally. Collum with carina present on anterolateral margins. Caudolateral corners of paranota rounded on body rings 1–4; body rings 5–19 with corners squared projecting caudally. Caudolateral corners of paranota 8–19 with small posterior-projecting tab. Paranota dorsal surface glossy, loosely wrinkled, appearing leathery. Ozopores opening dorsolaterally. Pore formula normal for order: 5, 7, 9, 10, 12, 13, 15–19. Paranota with anterodorsal area scooped out. Gonapophyses apically cylindrical. Pleural tubercle absent, with faint swelling between paranotal base and spiracle. Sterna without posteriorly projecting spines, with slightly anteriorly curved caudal margin. Sterna 2–9 sparsely setose (about 10 setae); sterna 10–18 lacking setae. Gonopods: Acropodite bent posteroventrally at base, curved medially in smooth O-shaped form ( Fig. 9A ). Distal-most arc of acropodite oriented laterally. O-shaped acropodite nearly closed between prefemoral process and apex ( Fig. 9A ). Right, left acropodites not crossed midlength, acropodites stacked. Gonocoxae with rounded protuberance apically, telopodites arising subapically. Telopodites—Prefemur with sharp prefemoral process, one-ninth length of acropodite, with apex tapered to sharp curved point, scythe-shaped, width at base 1/4 its length ( Fig. 9A pfp). Acropodite bent 90° posteroventrally at prefemur ( Fig. 9B ); prostatic groove bent 90° from cannula to acropodite base ( Fig. 9B ). Not with distinct bend tubercle at prefemur-acropodite junction. Gonopodal acropodite narrow, one-half width of tibia on leg pair 9; tapered to curved acuminate apex. Acropodite with ventrobasal surface facing laterally. Acropodite without spines on dorsal surface. Acropodite basal and apical ventral surfaces not coplanar, apical surface facing ventrolaterally; anterior twist faint ( Fig. 9B ). Acropodite ventral surface flat, smooth, no bumps nor swellings.Acropodite without cingulum.Acropodite shaft uniform width to region with setae, tapered to acuminate apex. Acropodite elliptical in cross-section, acuminate apex thinner, transparent. Acropodite margin smooth, rounded, lacking sharp edge; marginal teeth absent. Acropodite with apical 1/9 recurved, L-shaped, projecting cephalically ( Fig. 9B ). Acropodite base with setae about 1/3 its total length ( Fig. 9B ).

Paratype ( ♀) MPE 03857—Somatic measurements: BL = 56.54, CW = 6.64, IW = 4.83, ISW = 1.49, B10W = 10.07. Cyphopods: Cyphopod receptacle at its greatest width equal to prefemur length. Receptacle heart-shaped, pointed base facing medially. Cyphopodal valves symmetrical, anterior valve more convex. Cyphopods with valve suture facing laterally, suture without mid-length ramp-like swelling.

Variation. There are four color morphs of Apheloria uwharrie : (1) three-spotted, with yellow paranotal, metatergal and collum spots, and legs ( Fig. 15A ); (2) striped, with red metatergal—paranotal stripes, anterior collum spot, and legs ( Fig. 15B ); (3) striped/three-spotted superimposition of striped and three-spotted red morphs ( Fig. 15C ); and (4) three-spotted, with yellow paranotal spots and legs, and faint orange or missing metatergal and collum spots, appearing nearly two-spotted ( Fig. 15D ). There is a geographical assortment of color morphs, and A. uwharrie from the Uwharrie Mountains display the yellow spotted morph and those from Myrtle Beach have the red striped or spotted morphs.

There is typical sexual size dimorphism between males and females, where females are larger, and negligible variation of measurements within individuals of the same sex. Somatic measurements: ♂ (n = 5) BL = 46.19–55.63 (51.06/4.01). CW = 6.30–7.47 (6.95/0.50). IW = 4.02–4.86 (4.44/0.37). ISW = 1.26–1.33 (1.30/0.03). B10W = 8.72–10.24 (9.53/0.60). ♀ (n = 4) BL = 50.34–56.54 (53.38/3.15). CW = 6.64–7.82 (7.21/0.48). IW = 4.71–5.60 (4.97/0.42). ISW = 1.42–1.72 (1.56/0.13). B10W = 10.06–10.59 (10.20/0.26).

Ecology. Apheloria uwharrie individuals were often encountered in more xeric habitats than is typical for members of the tribe. Individuals from Uwharrie National Forest were encountered in dry hardwood slopes and ridges, and those from Myrtle Beach were found in an oak, pine, magnolia forest in dry leaf litter.An individual from Morrow Mountain State Park in the Uwharrie Mountains was discovered in a damp, deciduous forest composed of live oak, huckleberry, and maple.

Apheloria polychroma and A. corrugata are well-known mimics with sympatric xystodesmid species, such as Brachoria , Rudiloria , and Appalachioria , but A. uwharrie is not known to mimic others. There is a resemblance between A. uwharrie and syntopic family members such as with Sigmoria species and a Pleuroloma species. Sigmoria and Pleuroloma species have a shared yellow spotted coloration in the Uwharrie Mountains, and appear striped red in Myrtle Beach. These resemblances are geographically clustered with yellow spotted species occurring north of the Carolinas’ border and red striped species to the south ( Shelley and Whitehead 1986). This resemblance is likely a result of Müllerian mimicry, or for camouflage in red light-dominated forest ecosystems ( Endler 1993). Due to resemblance in color and overlap in distribution, A. uwharrie may be confused in the field with two species of Sigmoria [ Sigmoria latior ( Brölemann, 1900) and Sigmoria simplex ( Shelley, 1977) ] and a species of Pleuroloma , Pleuroloma pinicola Shelley, 1980 . Apheloria uwharrie can be distinguished from these taxa by gonopod morphology, specifically by the presence of a circular acropodite ( Fig. 9A ).

Distribution. Known from two areas: the Uwharrie Mountains, NC, and Myrtle Beach, SC (Montgomery, Randolph, and Stanly cos., NC; and Horry Co., SC; Fig. 11 ). Several localities intervening these were cited by Shelley (2000, 2007), and may be A. uwharrie . These counties are as follows: Richmond and Union cos., NC; and Chesterfield, Kershaw, and Georgetown cos., SC. The location from Georgetown Co., SC, is close to the type locality of A. uwharrie . The outline of the distribution of A. uwharrie in Figure 11 reflects these probable locations.

Etymology. The specific name is a noun in apposition, and derived from the northernmost localities of A. uwharrie in the Uwharrie Mountains, North Carolina. The Uwharrie Mountains are inselbergs formed by erosion of ancient terrains and are a rich repository of unusual and rare biological, geological and archaeological heritage ( Lewis 2023).

Figures

FIGURE 4. Apheloria corrugata (Wood, 1864), male gonopods (specimen # MPE00770). A: Anterior view; B: Medial view. pfp: prefemoral process. bt: bend tubercle.

FIGURE 7. Apheloria virginiensis (Drury, 1770), male gonopods (specimen # SPC000312). A: Anterior view; B: Medial view. pfp: prefemoral process. bt: bend tubercle. elb: elbow.

FIGURE 8. Apheloria polychroma Marek, Means & Hennen, 2018, male gonopods (specimen # MMC0311). A: Anterior view; B: Medial view. pfp: prefemoral process.

FIGURE 9. Apheloria uwharrie Marek, Means, Hennen & Tingley, sp. nov., male gonopods (specimen # MPE03855). A: Anterior view; B: Medial view. pfp: prefemoral process.

FIGURE 11. Distribution of species of the genus Apheloria Chamberlin, 1921. Colored dots indicate specimens used in the phylogenetic analysis. Outlines bound colored dots plus historical occurrence records from the periphery of the genus’ geographical range from Shelley & McAllister (2007), R.L Hoffman’s notes, and vetted research-grade observations from iNaturalist. Black dots denote unexamined material referable to Apheloria sp. and are undetermined.

FIGURE 13. Apheloria montana (Bollman, 1887), male gonopods (specimen # SPC000134). A: Anterior view; B: Medial view. pfp: prefemoral process. bt: bend tubercle.

FIGURE 15. Apheloria uwharrie Marek, Means, Hennen & Tingley, sp. nov., color morphs. A: Three-spotted, with yellow paranotal, metatergal and collum spots, and legs; B: Striped, with red metatergal—paranotal stripes, anterior collum spot, and legs; C: Striped/three-spotted superimposition of striped and three-spotted red morphs. D. Three-spotted yellow individual with a faint, orange or missing metatergal spots that appears nearly two-spotted.