Apheloria corrugata ( Wood, 1864 ) 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 : 319-322

Apheloria corrugata ( Wood, 1864) , new status

Vernacular name: “The pink and yellow cherry millipede”

Figs 2–6 , 10

Polydesmus (Fontaria) corrugatus Wood, 1864: 6 .

Fontaria butleriana Bollman, 1888: 407 , new synonymy.

Leptocircus inexpectatus Attems, 1931: 67 . Synonymized by Hoffman, 1999: 307.

Apheloria adela Chamberlin, 1939: 10 . Synonymized by Hoffman, 1957: 186.

Apheloria iowa Chamberlin, 1939: 10 , new synonymy.

Apheloria reducta Chamberlin, 1939: 11 , new synonymy.

Apheloria pinicola Chamberlin, 1947: 26 . Synonymized by Hoffman, 1999: 305.

Apheloria asburna Chamberlin, 1949a: 101 . Synonymized by Hoffman, 1999: 307.

Detailed taxonomic history in Appendix 1.

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

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 , 5A , 6 ). ♂ Gonopods. Gonopodal acropodite smoothly circular, O-shaped ( Fig. 3 , 4B )—without elbow as in A. virginiensis ( Fig. 7A , 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. 4B )—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. 4B , pfp)—not short, scythe-like as in A. polychroma ( Fig. 8A , pfp). With distinct bend tubercle at prefemur-acropodite junction ( Fig. 4B , bt), not with acute angle at junction as in A. polychroma , A. uwharrie ( Figs 8B , 9B ).

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.

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 , 5A , 6 ); (2) striped, with yellow metatergal—paranotal stripes, anterior collum stripe, and legs ( Fig. 10A ); (3) two-spotted, with yellow paranotal spots, and yellow legs ( Fig. 10B ); (4) three-spotted, with yellow paranotal, metatergal and collum spots, and yellow legs ( Fig. 10C ); (5) striped/three-spotted superimposition of striped (morph 1) and three-spotted yellow morphs (morph 4) ( Fig. 10D ); (6) striped, with pink metatergal—paranotal stripes, anterior collum stripe, and legs ( Fig. 10E ); and (7) striped/three-spotted superimposition of striped (morph 6) and three-spotted yellow morphs (morph 4) ( Fig. 10F ). Some that appear like morphs 2 or 4 have small brick red crescents next (anterior) to their yellow paranotal spots ( Figs 10A, 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). B10W = 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). B10W = 9.06–10.64 (9.79/0.46).

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, 1949b , Pleuroloma Rafinesque, 1820 , and Rudiloria Causey, 1955 : Appalachioria bondi Marek, Means, Hennen, 2021 ; Appalachioria calcaria ( Keeton, 1959) ( Fig. 5B ); 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 , 4A ).

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).

Figures

FIGURE 2. Apheloria corrugata (Wood, 1864), pink-spotted yellow stripe morph (specimen # MPE00770 from Price Mountain, Montgomery Co., Virginia).

FIGURE 3. Apheloria corrugata (Wood, 1864), ventral views showing in situ arrangement of gonopods. A: Ventral view of anterior rings illuminated with ultraviolet light (specimen # MPE01930); B: Ventral view illuminated with white light; C: Magnification of the gonopods from A showing their situ arrangement.

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

FIGURE 5. Müllerian mimicry between Apheloria corrugata (Wood, 1864) (A) and Appalachioria calcaria (Keeton, 1959) (B) in Blacksburg, Virginia.

FIGURE 6. The typical habitats of Apheloria corrugata (Wood, 1864) individuals are mesic broadleaf deciduous forests.

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 10. Apheloria corrugata (Wood, 1864), color morphs. A: Striped, with yellow metatergal—paranotal stripes, anterior collum stripe, and legs; B: Two-spotted, with yellow paranotal spots, and yellow legs; C: Three-spotted, with yellow paranotal, metatergal and collum spots, and yellow legs; D: Striped/three-spotted superimposition of striped (morph 1) and three-spotted yellow morphs (morph 4); E: Striped, with pink metatergal—paranotal stripes, anterior collum stripe, and legs; F: Striped/threespotted superimposition of striped (morph 6) and three-spotted yellow morphs (morph 4).

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.