taxonID	type	description	language	source
B36B3E40F851AF72FB98F9F5FAC4FC38.taxon	vernacular_names	Vernacular name: The cherry millipedes	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F851AF72FB98F9F5FAC4FC38.taxon	type_taxon	Type species: Fontaria montana Bollman, 1887, by original designation. Detailed taxonomic history in Appendix 1.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F851AF72FB98F9F5FAC4FC38.taxon	diagnosis	Genus diagnosis: Apheloria is distinct from other apheloriine genera based on the following combination of characters: Color. Tergites with 2 – 4 spots or stripes. Spot, stripe hues variable; usually yellow stripes, often with red spots on paranota (Fig. 2) — never with purple as in some species of Sigmoria. Those with yellow stripes often show a brick-red tinge on the inside margin of the paranotal spots. Gonopods. Gonopodal acropodite circular (Figs 3, 4 A) — not smoothly oval-shaped (0 - 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 gradually tapered to curved J- or L-shaped acuminate apex (Fig. 4 B). Acropodite shaft without cingulum nor preapical teeth nor projections as in Appalachioria, Brachoria. Prefemur with a scythe-like prefemoral process located medially (Fig. 4 B, pfp) — not absent, rounded, nor located marginally as in Appalachioria, Brachoria. Acropodite bent 90 ° posteroventrally at prefemur (Fig. 4 B); prostatic groove bent 90 ° from cannula to acropodite base. With acute angle or distinct protuberance on corner of bend, “ bend tubercle ” (Fig. 4 B, bt). Genus diagnosis notes: The genus diagnosis applies solely to the species of Apheloria besides A. whiteheadi, a geographically isolated species with very different looking acropodites. See diagnosis of A. whiteheadi below.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F851AF72FB98F9F5FAC4FC38.taxon	etymology	Genus etymology: Chamberlin did not provide an etymology of the genus name Apheloria when he named it in 1921. It is presumed that the name is from the Greek apo (ἀπό) meaning away or off, and the Greek helios, meaning sun (ἥΛΙΟΣ) (Brown 1954). Chamberlin may have selected the name because millipedes in the genus Apheloria, and most members of the family Xystodesmidae, flee from the sun to avoid desiccation. Similarly, the genus name appears similar to the word aphelion, which means a point on a circular orbit that is farthest from the sun. A circular orbit appears similar to the circular gonopods of the genus Apheloria. The circular acropodite is the basis of the single sentence description of the genus by Chamberlin (1921: 232), “ Erected for a group of species … in which the telopodite of the gonopod of male is a simple, coiled blade with a small spur at base. ” This latter explanation would be consistent with other genus names in the tribe Apheloriini Hoffman, 1980 by Chamberlin based on shape of the telopodite, such as Brachoria Chamberlin, 1939 and Sigmoria Chamberlin, 1939; and Rudiloria Causey, 1955.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F857AF71FB98FA2DFA22FE20.taxon	vernacular_names	Vernacular name: “ The pink and yellow cherry millipede ”	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F857AF71FB98FA2DFA22FE20.taxon	description	Figs 2 – 6, 10	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F857AF71FB98FA2DFA22FE20.taxon	materials_examined	Material examined: Type specimens — ♂ syntype (USNM) from New York, Oneida County, Trenton Falls (Coll: D. Mack), no other collection information provided (non vidi). Non type material examined in Appendix 2. Materials examined are archived in the Virginia Tech Data Repository at: https: // doi. org / 10.7294 / 29829209	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F857AF71FB98FA2DFA22FE20.taxon	diagnosis	Diagnosis: Apheloria corrugata is distinct from other apheloriine species based on the following combination of characters: Color. Tergites with two pink spots and yellow metatergal stripe, anterior collum stripe, and legs (Figs 2, 3, 5 A, 6). ♂ Gonopods. Gonopodal acropodite smoothly circular, O-shaped (Fig. 3, 4 B) — without elbow as in A. virginiensis (Fig. 7 A, elb). Not smoothly oval-shaped (0 - 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. 4 B) — not gradually tapered to curved J-shaped apex as in A. polychroma (Fig. 8 B). Acropodite shaft without cingulum nor preapical teeth nor projections as in Appalachioria, Brachoria. Prefemur with a long, scythe-like prefemoral process (Fig. 4 B, pfp) — not short, scythe-like as in A. polychroma (Fig. 8 A, pfp). With distinct bend tubercle at prefemur-acropodite junction (Fig. 4 B, bt), not with acute angle at junction as in A. polychroma, A. uwharrie (Figs 8 B, 9 B). Note about coloration: The pink-spotted yellow stripe morph unequivocally diagnoses A. corrugata from all other species of Xystodesmidae, except for in the Valley and Ridge Mountains of Virginia where Appalachioria calcaria (Keeton, 1959) is a mimic (Fig. 5). However, there are about 10 distinct color morphs of the species with some of these morphs appearing as if a product of the superimposition of distinct morphs atop one another, such as morphs 5 and 7 below (Figs 2, 10). Because color varies considerably intraspecifically, it should be cautiously used as a diagnostic character for this species.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F857AF71FB98FA2DFA22FE20.taxon	description	Variation. There are at least 10 color morphs of A. corrugata with a continuum of coloration between them (in order of decreasing frequency): (1) striped, with yellow metatergal stripes and pink paranotal spots, and yellow legs (Figs 2, 5 A, 6); (2) striped, with yellow metatergal — paranotal stripes, anterior collum stripe, and legs (Fig. 10 A); (3) two-spotted, with yellow paranotal spots, and yellow legs (Fig. 10 B); (4) three-spotted, with yellow paranotal, metatergal and collum spots, and yellow legs (Fig. 10 C); (5) striped / three-spotted superimposition of striped (morph 1) and three-spotted yellow morphs (morph 4) (Fig. 10 D); (6) striped, with pink metatergal — paranotal stripes, anterior collum stripe, and legs (Fig. 10 E); and (7) striped / three-spotted superimposition of striped (morph 6) and three-spotted yellow morphs (morph 4) (Fig. 10 F). Some that appear like morphs 2 or 4 have small brick red crescents next (anterior) to their yellow paranotal spots (Figs 10 A, C). 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 = 13) BL = 34.07 – 46.48 (39.53 / 3.14). CW = 6.00 – 8.19 (7.01 / 0.67). IW = 3.71 – 4.99 (4.40 / 0.35). ISW = 1.02 – 1.32 (1.16 / 0.09). B 10 W = 7.28 – 10.85 (8.94 / 1.12). ♀ (n = 14) BL = 37.85 – 57.23 (46.92 / 5.30). CW = 6.70 – 7.82 (7.14 / 0.29). IW = 4.52 – 5.64 (4.95 / 0.31). ISW = 1.20 – 1.57 (1.41 / 0.11). B 10 W = 9.06 – 10.64 (9.79 / 0.46).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F857AF71FB98FA2DFA22FE20.taxon	biology_ecology	Ecology. Apheloria corrugata individuals were typically encountered in mesic habitats such as broadleaf deciduous forests (Fig. 6). More seldom were they found in mixed forests, rhododendron groves, cedar glades, and hemlock forests. Syntopic tree species recorded with A. corrugata included sugar maple, tulip poplar, white oak, pine, sycamore, beech, hickory, and black walnut. Individuals were normally found beneath decomposing leaves and occasionally walking atop detritus on the forest floor. Individuals of A. corrugata are commonly encountered walking on the lawn of the first author’s home in Blacksburg, Virginia in spring, and they may also enter houses, particularly damp basements (W. Shear, pers. communication). Due to mimicry in color and overlap in distribution, A. corrugata may be confused in the field with A. polychroma, A. virginiensis, A. whiteheadi and the following species of Appalachioria, Brachoria, Cherokia Chamberlin, 1949 b, Pleuroloma Rafinesque, 1820, and Rudiloria Causey, 1955: Appalachioria bondi Marek, Means, Hennen, 2021; Appalachioria calcaria (Keeton, 1959) (Fig. 5 B); Appalachioria ethotela (Chamberlin, 1942); Appalachioria falcifera Keeton, 1959; Appalachioria hamata (Keeton, 1959); Appalachioria separanda Chamberlin, 1947; Appalachioria versicolor (Hoffman, 1963); Brachoria badbranchensis Marek, 2010; Brachoria blackmountainensis Marek, 2010; Brachoria campcreekensis Marek, 2010; Brachoria electa Causey, 1955; Brachoria flammipes Marek, 2010; Brachoria gracilipes (Chamberlin, 1947); Brachoria grapevinensis Marek, 2010; Brachoria hoffmani Keeton, 1959; Brachoria indianae (Bollman, 1888); Brachoria laminata Keeton, 1959; Brachoria ligula Keeton, 1959; Brachoria viridicolens (Hoffman, 1948); Cherokia georgiana (Bollman, 1889); Pleuroloma flavipes Rafinesque, 1820; Rudiloria guyandotta (Shear, 1972); Rudiloria mohicana Causey, 1955; Rudiloria rigida Shelley, 1986; and Rudiloria trimaculata (Wood, 1864). Apheloria corrugata can be distinguished from other species of Apheloria by its diagnosis, and from Appalachioria, Brachoria, Cherokia, Pleuroloma and Rudiloria species by gonopod morphology, specifically by the absence of a cingulum, or a mid-length transverse groove on the acropodite, and the presence of a circular acropodite (Figs 3, 4 A).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F857AF71FB98FA2DFA22FE20.taxon	distribution	Distribution. Apheloria corrugata has the greatest distributional area of the genus, and is known from Virginia north to Connecticut and Montreal; west to southeastern Ontario Province, southern Michigan, southeasternmost Wisconsin; south to Illinois, southeastern Iowa, southern Missouri; western Oklahoma, northern Arkansas, western Tennessee, and the far northwestern corner of Alabama (Fig. 11). Pleuroloma flavipes has the greatest distributional area of the family and extends further south (to northern Louisiana) and west (Shelley 1980, Shelley et al. 2003).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F854AF7CFB98F8A4FC48FE58.taxon	vernacular_names	Vernacular name: “ The mountain cherry millipede ”	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F854AF7CFB98F8A4FC48FE58.taxon	description	Figs 12, 13	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F854AF7CFB98F8A4FC48FE58.taxon	materials_examined	Material examined: Type specimens — ♂ holotype (USNM) from Tennessee, Cocke County, Wolf Creek (Coll: C. Branner), no other collection information provided (non vidi). Non type material examined in Appendix 2. Materials examined are archived in the Virginia Tech Data Repository at: https: // doi. org / 10.7294 / 29829209	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F854AF7CFB98F8A4FC48FE58.taxon	diagnosis	Diagnosis: Apheloria montana is distinct from other apheloriine species based on the following combination of characters: Color. Tergites with three yellow spots and yellow legs (Fig. 12). ♂ Gonopods. Gonopodal acropodite smoothly circular, O-shaped (Fig. 13 A) — without elbow as in A. virginiensis (Fig. 7 A, elb). Not smoothly oval-shaped (0 - 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. 13 B) — not gradually tapered to curved J-shaped apex as in A. polychroma (Fig. 8 B). Acropodite shaft without cingulum nor preapical teeth nor projections as in Appalachioria, Brachoria. Prefemur with a long, scythe-like prefemoral process (Fig. 13 B, pfp) — not short, scythe-like as in A. polychroma (Fig. 8 A, pfp). With distinct bend tubercle at prefemur-acropodite junction (Fig. 13 B, bt), not with acute angle at junction as in A. polychroma, A. uwharrie (Figs 8 B, 9 B). Note about coloration: The three-spotted yellow morph unequivocally diagnoses A. montana from all other species of Apheloria northeast of the Great Smoky Mountains National Park in the Blue Ridge Mountains of Tennessee and North Carolina (Fig. 12).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F854AF7CFB98F8A4FC48FE58.taxon	description	Variation. The three-spotted yellow morph is the single color morph of A. montana. Individuals from Little Switzerland, Mitchell Co., North Carolina, possess smaller orange spots and orange legs. 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 = 7) BL = 41.85 – 52.89 (46.56 / 3.71). CW = 6.57 – 7.75 (6.91 / 0.45). IW = 4.24 – 4.89 (4.50 / 0.27). ISW = 1.15 – 1.31 (1.25 / 0.06). B 10 W = 8.74 – 10.51 (9.49 / 0.72). ♀ (n = 7) BL = 46.69 – 54.87 (49.51 / 2.98). CW = 6.44 – 8.21 (7.11 / 0.71). IW = 4.08 – 5.26 (4.75 / 0.41). ISW = 1.10 – 1.41 (1.30 / 0.11). B 10 W = 8.42 – 11.20 (9.78 / 0.99).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F854AF7CFB98F8A4FC48FE58.taxon	biology_ecology	Ecology. Apheloria montana individuals were typically encountered in mesic habitats such as broadleaf deciduous forests. They were also found in drier habitats such as mixed forests, rhododendron groves, and oak and beech forests. Syntopic tree species recorded with A. montana included pine, maple, oak, tulip poplar, witch hazel, alder, hemlock, sweetgum, buckeye, and maple. Individuals were typically found beneath decomposing leaves on the forest floor. Due to mimicry in color and overlap in distribution, A. montana may be confused in the field with A. polychroma; Appalachioria eutypa (Chamberlin, 1939); Brachoria hendrixsoni Marek, 2010; C. georgiana; and P. flavipes. Apheloria montana can be distinguished from other species of Apheloria by its diagnosis, and from Appalachioria, Brachoria, Cherokia and Pleuroloma species by gonopod morphology, specifically by the absence of a cingulum, or a mid-length transverse groove on the acropodite, and the presence of a circular acropodite (Fig. 13 A).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F854AF7CFB98F8A4FC48FE58.taxon	distribution	Distribution. Known from western North Carolina in Buncombe, Madison, McDowell, and Mitchell cos., and eastern Tennessee in Greene, Unicoi, and Washington cos. (Fig. 11). Apheloria montana has the second smallest distributional area of the genus; A. whiteheadi has the smallest distribution.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F858AF7BFB98F99CFEF1FBC4.taxon	vernacular_names	Vernacular name: “ The colorful cherry millipede ”	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F858AF7BFB98F99CFEF1FBC4.taxon	description	Figs 8, 14	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F858AF7BFB98F99CFEF1FBC4.taxon	materials_examined	Material examined: Type specimens — ♂ holotype (FMNH-INS 60792), 1 ♀ paratype (FMNH-INS 71228), 3 ♂, 3 ♀ paratypes (VTEC, MMC 0309, 310, 313, 305, 306, 308), 1 ♂, 1 ♀ paratypes (VMNH, MMC 0314, 312) from Virginia, Lee County, The Cedars, The Cedars State Natural Area Preserve, CR- 738 (36.65624 ° N, - 83.20165 ° W, Elev. 436 m), 28 September 2006, 16: 00 (Colls: P. and B. Marek) (vidi). Non type material examined in Appendix 2. Materials examined are archived in the Virginia Tech Data Repository at: https: // doi. org / 10.7294 / 29829209	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F858AF7BFB98F99CFEF1FBC4.taxon	diagnosis	Diagnosis: Apheloria polychroma is distinct from other apheloriine species based on the following combination of characters: Color. Tergites usually with 4 yellow spots on a jet black background: 1 metatergal, 1 prozonal, 2 paranotal spots (Fig. 14). Collum usually uniformly covered in yellow (Fig. 10). ♂ Gonopods. Gonopodal acropodite smoothly circular, O-shaped (Fig. 8 A) — without elbow as in A. virginiensis (Fig. 7 A, elb). Not smoothly oval-shaped, 0 - shaped, as in Rudiloria. Acropodite narrow, one-half width of tibia on leg pair 9; of uniform width throughout. Acropodite gradually tapered to curved acuminate, J-shaped apex (Fig. 8 B) — not L-shaped and abruptly twisted as in A. corrugata, A. virginiensis, A. montana (Figs 4 B, 7 B, 13 B). Acropodite shaft without cingulum nor preapical teeth nor projections as in Appalachioria, Brachoria. Prefemur with a shorter, scythe-like prefemoral process, one-ninth length of acropodite (Fig. 8 A, pfp) — not long and scythe-like as in A. corrugata, A. virginiensis, A. montana (Figs 4 A, 7 A, 13 A). With acute angle at prefemur-acropodite junction (Fig. 8 B), not with distinct bend tubercle at junction as in A. corrugata, A. virginiensis, A. montana (Figs 4 B, 7 B, 13 B, bt). Note about coloration: When it is present, the 4 - spotted yellow color morph with a collum that is uniformly covered in yellow unequivocally diagnoses A. polychroma from all other species of Xystodesmidae (Fig. 14). However, this morph is geographically restricted to the Powell River Valley from Big Stone Gap, Virginia, in the north and Norris, Tennessee in the south. There are no fewer than six distinct color morphs of the species with a continuum of hues and patterns between them (“ Figs 2 A – T, 5 ” in Marek et al. 2018). There are often multiple color morphs of the species that are syntopic and co-occur in a small area (≥ 10 m 2). Because color varies considerably intraspecifically, it should be cautiously used as a diagnostic character for this species.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F858AF7BFB98F99CFEF1FBC4.taxon	description	Variation. There are at least six color morphs of Apheloria polychroma with a continuum of coloration between them: (1) four-spotted, with yellow to orange paranotal, metatergal, prozonal and collum spots, and legs (collum often uniformly covered in yellow, Fig. 14) and sometimes with red legs; (2) striped, with yellow metatergal — paranotal stripes, anterior collum stripe, and legs (“ Figs 2 G, H ” in Marek et al. 2018); (3) three-spotted, with creamwhite paranotal, metatergal and collum spots, and red legs (“ Figs 2 I, J ” in Marek et al. 2018); (4) three-spotted, with yellow paranotal, metatergal and collum spots, and yellow or red legs (“ Figs 2 K, L; 5 A, B ” in Marek et al. 2018); (5) striped / four-spotted superimposition of striped and four-spotted yellow morphs (“ Figs 2 M, N ” in Marek et al. 2018); and (6) two-spotted, with yellow paranotal spots, and yellow or red legs (“ Figs 2 Q – T ” in Marek et al. 2018). Some three-spotted yellow individuals have faint metatergal spots and appear nearly two-spotted (“ Fig. 2 S ” in Marek et al. 2018). The dorsal color of A. polychroma is always yellow to red and jet black; however, the pattern varies from two, three, or four spots, to metatergal stripes, and superimposition of the latter two patterns. 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 = 13) BL = 39.59 – 58.52 (50.24 / 6.18). CW = 6.76 – 9.22 (7.69 / 0.69). IW = 4.19 – 5.21 (4.78 / 0.28). ISW = 1.10 – 1.72 (1.27 / 0.18). B 10 W = 9.29 – 12.47 (10.43 / 0.81). ♀ (n = 10) BL = 37.78 – 53.47 (44.36 / 5.78). CW = 6.75 – 8.62 (7.66 / 0.53). IW = 4.61 – 5.69 (5.09 / 0.32). ISW = 1.20 – 1.52 (1.34 / 0.10). B 10 W = 8.90 – 12.70 (10.61 / 1.09).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F858AF7BFB98F99CFEF1FBC4.taxon	biology_ecology	Ecology. Apheloria polychroma individuals were typically encountered in mesic habitats such as broadleaf deciduous forests, and seldom in more xeric habitats such as glades — for example in The Cedars Natural Area Preserve in Lee County, Virginia that consists of karst overlain with a mixed deciduous and eastern red cedar forest (Juniperus virginiana L.). Individuals were normally found beneath decomposing leaves and occasionally walking atop detritus on the forest floor. These individuals were more often exposed to view than others of the family, bolder in behavior, and were more likely to writhe and flail when disturbed — compared to coiling into a ball like others. When handled, individuals would emit copious amounts of defense secretions and liquid feces. Due to mimicry in color and overlap in distribution, A. polychroma may be confused in the field with A. corrugata, A. montana, Appalachioria eutypa (Chamberlin, 1939); C. georgiana (Bollman, 1889); P. flavipes; and eight species of Brachoria: Brachoria cedra Keeton, 1959; Brachoria dentata Keeton, 1959; Brachoria hansonia Causey, 1950; Brachoria hoffmani Keeton, 1959; Brachoria insolita Keeton, 1959; Brachoria mendota Keeton, 1959; Brachoria sheari Marek, 2010; Brachoria splendida (Causey, 1942). Mimetic resemblance between A. polychroma and B. mendota at Natural Tunnel State Park (Scott Co., Virginia) is one of the most accurate in the millipede mimicry system (Marek & Bond 2009, Marek et al. 2018). The Pennington Gap mimic millipede, Brachoria dentata Keeton, 1959, mimics A. polychroma in color morph at six of the seven localities where they were found to co-occur. Five other species of Brachoria co-occur with A. polychroma at fewer localities. When syntopic with A. polychroma, though rare, some individuals of the species B. cedra, B. dentata, B. mendota, and B. sheari can have 4 yellow spots, including the distinctive prozonal spot, and appear almost indistinguishable from A. polychroma. However, these Brachoria species never possess a collum that is uniformly covered in yellow. Apheloria polychroma can be distinguished from other species of Apheloria by its diagnosis, and from Appalachioria, Brachoria, Cherokia, and Pleuroloma species by gonopod morphology, specifically by the absence of a cingulum, or a mid-length transverse groove on the acropodite, and the presence of a circular acropodite (Fig. 8 A).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F858AF7BFB98F99CFEF1FBC4.taxon	distribution	Distribution. Known from the mountains between southwestern Virginia, southeastern Kentucky, eastern Tennessee, and northwestern Georgia (Fig. 11). Not known to occur in North Carolina. The greatest diversity in coloration of A. polychroma is seen in southwestern Virginia where the taxon is syntopic with six species of the genus Brachoria.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F85EAF66FB98FB15FE9EFEEC.taxon	description	Figs 9, 15	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F85EAF66FB98FB15FE9EFEEC.taxon	materials_examined	Material examined: Type specimens — ♂ holotype (VTEC, MPE 03855), 1 ♀ paratype (VTEC, MPE 03857), 2 ♂, 2 ♀ paratypes (VMNH, MPE 03856, 3858; VTEC, 3859, 3875) from South Carolina, Horry Co., Myrtle Beach: Forested area near the intersection of Robert M Grissom Parkway and 29 th 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, MPE 01977, MPE 01979), 1 ♀ (VMNH, MPE 01978) 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, MPE 05400, 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, MPE 05401) 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	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F85EAF66FB98FB15FE9EFEEC.taxon	diagnosis	Diagnosis: Apheloria uwharrie is distinct from other apheloriine species based on the following combination of characters: Color. Tergites with three yellow (Uwharrie Mountains, Fig. 15 A) or red spots (Myrtle Beach) or red stripes (Myrtle Beach, Fig. 15 B). ♂ Gonopods. Gonopodal acropodite smoothly circular, O-shaped (Fig. 9 A) — without elbow as in A. virginiensis (Fig. 7 A, 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. 9 B) — not gradually tapered to curved J-shaped apex as in A. polychroma (Fig. 8 B). Acropodite shaft without cingulum nor preapical teeth nor projections as in Appalachioria, Brachoria. Prefemur with a long, scythe-like prefemoral process (Fig. 9 B, pfp) — not short, scythe-like as in A. polychroma (Fig. 8 B, pfp). With acute angle at prefemur-acropodite junction (Fig. 9 B), not with distinct bend tubercle at junction as in A. corrugata, A. virginiensis, A. montana (Fig. 4 B, 7 B, 13 B, 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.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F85EAF66FB98FB15FE9EFEEC.taxon	description	Description. Based on holotype (♂) MPE 03855 Measurements: BL = 49.18, CW = 6.70, IW = 4.25, ISW = 1.27, B 11 W = 9.30. Head: Antenna length — extendable posteriorly to anterior margin of 3 rd 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 8 th 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. 9 A). Distal-most arc of acropodite oriented laterally. O-shaped acropodite nearly closed between prefemoral process and apex (Fig. 9 A). 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. 9 A pfp). Acropodite bent 90 ° posteroventrally at prefemur (Fig. 9 B); prostatic groove bent 90 ° from cannula to acropodite base (Fig. 9 B). 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. 9 B). 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. 9 B). Acropodite base with setae about 1 / 3 its total length (Fig. 9 B). Paratype (♀) MPE 03857 — Somatic measurements: BL = 56.54, CW = 6.64, IW = 4.83, ISW = 1.49, B 10 W = 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. 15 A); (2) striped, with red metatergal — paranotal stripes, anterior collum spot, and legs (Fig. 15 B); (3) striped / three-spotted superimposition of striped and three-spotted red morphs (Fig. 15 C); and (4) three-spotted, with yellow paranotal spots and legs, and faint orange or missing metatergal and collum spots, appearing nearly two-spotted (Fig. 15 D). 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). B 10 W = 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). B 10 W = 10.06 – 10.59 (10.20 / 0.26).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F85EAF66FB98FB15FE9EFEEC.taxon	biology_ecology	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. 9 A).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F85EAF66FB98FB15FE9EFEEC.taxon	distribution	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.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F85EAF66FB98FB15FE9EFEEC.taxon	etymology	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).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F843AF67FB98FEF5FBB0F8A9.taxon	vernacular_names	Vernacular name: “ The Virginia cherry millipede ”	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F843AF67FB98FEF5FBB0F8A9.taxon	description	Figs 7, 16	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F843AF67FB98FEF5FBB0F8A9.taxon	materials_examined	Material examined: Type specimens — type material lost (BMNH?) from Virginia, Dinwiddie County (Coll: J. Greenway), no other collection information provided. ♂ Neotype (FSCA), 1 ♀, 2 ♂ paratypes (FSCA), 1 ♂ paratype (USNM), 1 ♀ paratype (VMNH) from Virginia, Dinwiddie County, McKenney, VA- 40, 1.3 km west VA- 644 (36.993611 ° N, - 77.739444 ° W), 8 July 2016 (Colls: R. Shelley, G. Phillips) (non vidi). Non type material examined in Appendix 2. Material examined are archived in the Virginia Tech Data Repository at: https: // doi. org / 10.7294 / 29829209	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F843AF67FB98FEF5FBB0F8A9.taxon	diagnosis	Diagnosis: Apheloria virginiensis is distinct from other apheloriine species based on the following combination of characters: Color. Tergites with three yellow spots and yellow legs (Fig. 16 A). ♂ Gonopods. Gonopodal acropodite circular but with elbow (Fig. 7 A, elb) — not uniformly circular as in A. polychroma (Fig. 8 A). Not smoothly oval-shaped, 0 - 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. 7 B) — not gradually tapered to curved J-shaped apex as in A. polychroma (Fig. 8 B). Acropodite shaft without cingulum nor preapical teeth nor projections as in Appalachioria, Brachoria. Prefemur with a long, scythe-like prefemoral process (Fig. 7, pfp) — not short, scythe-like as in A. polychroma (Fig. 8 B, pfp). With distinct bend tubercle at prefemur-acropodite junction (Fig. 7 B, bt), not with acute angle at junction as in A. polychroma, A. uwharrie (Figs 8 B, 9 B). Note about coloration. A millipede with two or three yellow spots and circular acropodites occurring in the coastal plain and piedmont of North Carolina, excepting the Uwharrie Mountains, unequivocally diagnoses A. virginiensis from all other species. However, there are at least five distinct color morphs of the species. Because color varies intraspecifically, caution should be exercised with using it as a diagnostic character for identification of this species.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F843AF67FB98FEF5FBB0F8A9.taxon	description	Variation. There are five color morphs of A. virginiensis with a continuum of coloration between them (in order of decreasing frequency): (1) two-spotted, with yellow paranotal spots, and yellow legs (Fig. 16 B); (2) three-spotted, with yellow paranotal, metatergal and collum spots, and yellow legs (Fig. 16 A); (3) three-spotted, with pink paranotal spots, yellow metatergal and collum spots, and yellow legs (Fig. 16 D); (4) two-spotted, with orange paranotal spots, and orange legs; (5) three-spotted, with pink paranotal, metatergal and collum spots, and pink legs (Fig. 16 C). The two-spotted morphs often have faint metatergal spots on posterior rings 15 or 17 – 19. Morph 3 often have pink paranotal spots connected by a faint and thin pink stripe. 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 = 10) BL = 26.73 – 58.86 (47.41 / 8.96). CW = 5.02 – 8.30 (7.11 / 0.87). IW = 3.73 – 4.96 (4.57 / 0.36). ISW = 1.08 – 1.52 (1.33 / 0.13). B 10 W = 6.77 – 10.93 (9.68 / 1.21). ♀ (n = 7) BL = 33.02 – 51.56 (44.54 / 7.24). CW = 6.41 – 7.38 (6.97 / 0.33). IW = 4.25 – 5.21 (4.77 / 0.31). ISW = 1.24 – 1.64 (1.43 / 0.15). B 10 W = 8.42 – 10.21 (9.39 / 0.73).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F843AF67FB98FEF5FBB0F8A9.taxon	biology_ecology	Ecology. Apheloria virginiensis individuals were typically encountered in mesic habitats such as broadleaf deciduous forests. They were also found in mixed forests, and in sandy soils. Syntopic tree species recorded with A. virginiensis included pine, birch, beech, maple, oak, sweet gum, walnut, oak, magnolia, hickory, rhododendron, tulip poplar, and cherry. Individuals were normally found beneath decomposing leaves and logs, and occasionally walking atop leaf litter or on trails at night. They were often encountered beside streams and low sandy woods. Apheloria polychroma and A. corrugata are well-known mimics with sympatric xystodesmid species, such as Brachoria, Rudiloria, and Appalachioria, but A. virginiensis is not well known to mimic others. Due to similarity in color and overlap in distribution, A. virginiensis may be confused in the field with A. corrugata and A. whiteheadi and the following syntopic taxa of its family: Sigmoria latior (Brölemann, 1900) and Pleuroloma flavipes Rafinesque, 1820. The latter two species share with A. virginiensis a two-spotted yellow coloration. This resemblance is likely a result of Müllerian mimicry, however similarity may be due to another convergent evolution. Apheloria virginiensis can be distinguished from other species of Apheloria by its diagnosis, and from Sigmoria and Pleuroloma species by gonopod morphology, specifically by the absence of a cingulum, or a mid-length transverse groove on the acropodite, and the presence of a circular acropodite (Fig. 7 A).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F843AF67FB98FEF5FBB0F8A9.taxon	distribution	Distribution. Known primarily from the coastal plain and piedmont of North Carolina (Fig. 11). Apheloria virginiensis is commonly encountered in parks and natural areas in the Research Triangle of North Carolina, such as William B. Umstead State Park. Apheloria virginiensis also occurs in the north of its distribution in the coastal plain, piedmont, and Blue Ridge Mountains of Virginia. The westernmost localities are Wilkes County, North Carolina, and the Blue Ridge Mountains in Floyd County, Virginia. In the piedmont of Virginia, in Prince Edward County, A. virginiensis occurs on the campus of Hampden-Sydney College, but 10 km northeast is replaced by A. corrugata in Farmville. Material provided to the first author by W. Shear collected from these two locales have confirmed this allopatry and northernmost limit of A. virginiensis at Hampden-Sydney College.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F841AF65FB98FF04FE30FB8B.taxon	vernacular_names	Vernacular name: “ The Laurel Creek millipede ”	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F841AF65FB98FF04FE30FB8B.taxon	description	Figs 17, 18	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F841AF65FB98FF04FE30FB8B.taxon	discussion	Detailed taxonomic history in Appendix 1.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F841AF65FB98FF04FE30FB8B.taxon	materials_examined	Material examined: Type specimens — ♂ holotype (VMNH), 1 ♂, 2 ♀ paratypes (VMNH) from Virginia, Patrick County, along Laurel Creek on Blue Ridge Parkway at mile 174.3, 20 May 1983, 16: 00 (Colls: R. Hoffman). (vidi). (1 paratype collected by R. Hoffman from same locality on 23 June 1984.) Non type material examined in Appendix 2. Material examined are archived in the Virginia Tech Data Repository at: https: // doi. org / 10.7294 / 29829209	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F841AF65FB98FF04FE30FB8B.taxon	diagnosis	Diagnosis: Apheloria whiteheadi is distinct from other apheloriine species based on the following combination of characters: Color. Tergites with yellow stripes and legs (Fig. 17). ♂ Gonopods. Gonopodal acropodite ρ-shaped (Fig. 18 A) — not uniformly circular as in other Apheloria species nor smoothly oval-shaped as in Rudiloria. Acropodite apex flange-like and abruptly twisted (Fig. 18 B) — not of uniform width throughout as in other Apheloria species. Acropodite shaft without cingulum nor preapical teeth nor projections as in Appalachioria, Brachoria. Prefemur with a short, blunt prefemoral process located marginally (Fig. 18 B, pfp) — not long, scythe-like as in other Apheloria species nor located medially (Figs 4 B, 7 B, 8 B, 9 B, 13 B, pfp). Acropodite not bent 90 ° posteroventrally at prefemur (Fig. 17 B), prostatic groove generally straight from cannula to acropodite base, without bend tubercle nor acute angle on corner of bend (Figs 4 B, 7 B, 13 B, bt). Note about coloration: The anterior and posterior metatergal stripes of the collum meet at the paranota and appear as a continuous yellow ring around the collum’s periphery.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F841AF65FB98FF04FE30FB8B.taxon	description	Variation. There is a single color morph of A. whiteheadi: striped, with yellow metatergal — paranotal stripes, anterior collum stripe, and legs (Fig. 17). 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 = 2) BL = 40.53 – 40.78 (40.66 / 0.18). CW = 5.54 – 5.59 (5.57 / 0.04). IW = 3.65 – 3.77 (3.71 / 0.08). ISW = 1.01 – 1.05 (1.03 / 0.03). B 10 W = 6.89 – 6.91 (6.90 / 0.01). ♀ (n = 10) BL = 31.42 – 45.96 (38.71 / 5.32). CW = 5.59 – 6.59 (5.89 / 0.30). IW = 3.89 – 4.48 (4.17 / 0.18). ISW = 1.06 – 1.22 (1.13 / 0.06). B 10 W = 7.09 – 8.39 (7.59 / 0.36).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F841AF65FB98FF04FE30FB8B.taxon	biology_ecology	Ecology. Apheloria whiteheadi individuals were typically encountered in rhododendron cove forests (Means and Marek, 2017). More seldom were they found in montane mixed oak forest (Means et al. 2021 c). Syntopic tree species recorded with A. whiteheadi included oak, maple, tulip poplar, witch hazel, pine, beech, paper birch, and sweet gum. Due to potential mimicry in color and overlap in distribution, A. whiteheadi may be confused in the field with A. corrugata, A. virginiensis, and P. flavipes. Apheloria whiteheadi can be distinguished from these taxa by its distinctive gonopod morphology, specifically by its ρ-shaped acropodite (Fig. 18 A), which is not uniformly circular as in other Apheloria species, nor sublinear in shape, with an acicular prefemoral process as in P. flavipes. Pleuroloma flavipes also possesses a distinctive claw-like bifurcated acropodite composed of a tibial process and a solenomere (Shelley, 1980).	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F841AF65FB98FF04FE30FB8B.taxon	distribution	Distribution. Apheloria whiteheadi has the smallest distributional area of the genus, and is known from Floyd and Patrick counties in Virginia (Fig. 11). The species is listed as a threatened species in Virginia and is afforded protection by state law.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F840AF62FB98FA96FA36FEEC.taxon	discussion	Remarks. The status of Fontaria luminosa Kenyon, 1893 has been uncertain for some time. See the following articles for a discussion of the matter: Hoffman (1999), Shelley & Whitehead (1986), Shelley & McAllister (2007), and Marek et al. (2014). It was described as a luminous or phosphorescent member of the genus Fontaria from Omaha, Nebraska (Bruner 1891, Kenyon 1893). There are other bioluminescent family members: the 11 species of the genus Motyxia in California (Marek & Moore 2015). However, Kenyon’s and Bruner’s description of the shape of its photic organs as round and dorsally situated do not correspond with Motyxia nor any bioluminescent millipede known. The entire exoskeleton is bioluminescent in these millipedes. The type specimens of Kenyon’s are lost, but Shelley did find material in the USNM that is labeled “ Fontaria luminosa (Type?) ”. Although this material is labeled from Omaha, which is the type locality mentioned by Kenyon, the single male specimen’s gonopods were missing (Shelley & Whitehead 1986). Despite searching the Omaha area, Shelley and Hoffman were unable to find any millipedes of the genus Apheloria. Pleuroloma flavipes is known to occur in the Omaha area and is commonly encountered in eastern Nebraska (Shelley 1980). For more than 130 years since its discovery, the only material of the family found in the area was that of P. flavipes. Based on this information, and to bring order to the classification, we here synonymize F. luminosa with P. flavipes. In the unlikely event there is a bioluminescent Apheloria from the region, then F. luminosa can be revived, but for the present time and based on much collections of P. flavipes in the area, the synonymy seems justifiable. The description of the bioluminescence (Bruner 1891) matches that of the glowworm beetle genus Phengodes (Phengodidae), whose adult females are larviform and predaceous on millipedes, and are sometimes themselves confused for millipedes (Ferreira et al. 2024). It may be that Bruner observed Phengodes females glowing and collected their millipede prey, thereby convincing himself that the millipedes were producing the bioluminescence.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F848AF6DFB98FE8CFB90FC9F.taxon	discussion	Remarks: Apheloria roanea was described from Roane County, which falls within the range of Apheloria polychroma and is outside the range of A. montana. Hoffman synonymized it under Apheloria montana, but the original description fits better with A. polychroma, potentially making A. polychroma a junior synonym. Unfortunately, fresh material from the type locality is not available, and the type and only known specimen of A. roanea cannot be located. Additionally the name Apheloria polychroma has now been used in a multitude of publications. For taxonomic stability we do not list A. polychroma as a junior synonym of A. roanea.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F84FAF6AFB98FF4CFF11FDFB.taxon	type_taxon	Type species: Pleuroloma flavipes Rafinesque, by monotypy.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
B36B3E40F84FAF6AFB98FF4CFF11FDFB.taxon	discussion	Note: The most recent revision of Pleuroloma was accomplished by Shelley, 1980, Canadian Journal of Zoology, 58.	en	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): 315-350, DOI: 10.11646/zootaxa.5701.3.4, URL: https://doi.org/10.11646/zootaxa.5701.3.4
