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Rudiloria ‘n. sp. Chagrin’ Means and Marek 2017 Material examined: ♂ holotype (VTEC MTX 0185) from Ohio, Cuyahoga County, Cleveland Metroparks, South Chagrin Reservation, Henry Church Rock Loop Trail, near falls and swimming hole (41.41598°N, - 81.4145°W), 12 June 2010, ex on top dead leaves in maple forest (Coll: P. E. Marek). Material examined listed in Supp Table 2 [online only]. Diagnosis: Adult males of Rudiloria charityae n. sp. are distinct from other apheloriine species based on the following combination of characters: Gonopods. Gonopodal acropodite smoothly oval-shaped (Fig. 12A)—not circular as in Apheloria species. Acropodite (when viewed posteriorly) narrow, onethird width of tibia on leg pair 9. Acropodite gradually tapered to acuminate apex–curved one-dimensionally to apex, not twisted cephalically as in Apheloria species. Acropodite shaft smooth without cingulum (midlength transverse groove) as in Brachoria; without teeth, swellings, joints, or projections as in other Rudiloria species. Prefemur with a nubbin-like prefemoral process, one-tenth length of acropodite (Fig. 12C), not long and scythe-like as in Rudiloria kleinpeteri, Rudiloria trimaculata. Color. Tergites with 3 spots: 1 orange metatergal spot, 2 yellow paranotal spots (Fig. 5Q). Collum with yellow spot anteriorly, gradually fading into orange metatergal spot. Description: Supp Table 3 (online only). Based on Holotype (♂) MTX0185. Measurements (mm): BL = 34.0, CW = 6.1, IW = 4.1, ISW = 1.2, B11W = 7.2, B11H = 4.5. Variation: No significant variation from the holotype was observed. Ecology: Individuals of Rudiloria charityae n. sp. have been collected in deciduous broadleaved forests of maple and beech. Distribution: Rudiloria charityae n. sp. is known from Cuyahoga and Geauga counties in the Erie Drift Plain Region, and Monroe County in the Western Allegheny Plateau Region of eastern Ohio. Etymology: The holotype specimen was collected on the wedding day of PEM during a family hike to his childhood swimming hole in the Chagrin River, Cuyahoga County, Ohio. The species is named for his wife Charity Hall., Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on page 15, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533, {"references":["Means, J. C., and P. E. Marek. 2017. Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae? PeerJ 5: e 3854."]}

Genus Nannaria Chamberlin, 1918 Vernacular Name: ‘TheTwisted-Claw Millipedes’ Type species: Nannaria minor Chamberlin, 1918. Taxa included: 26 species, see Supp Material (online only) (Supp Text 1 [online only], Supp Dataset 2 [online only])., Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on page 16, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533

Nannaria hokie Means, Hennen, Marek, New Species Vernacular Name: ‘The Hokie Twisted-Claw Millipede’, Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on page 17, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533

Cleptoria ‘n. sp. Hagoods’ Means and Marek 2017 Material examined: Type specimens. ♂ holotype (VTEC MPE00291), 1 ♂ paratype (VMNH SPC000389), and 2 ♀ paratypes (VTEC SPC000390, 394). Holotype from South Carolina, Barnwell County, Toby Creek, near Hagoods Mill, SC-64 (33.2119°N, - 81.3201°W, Elev: 46 m), 23 November 2003, (Coll: D. Beamer). Paratypes from South Carolina, Barnwell County, Hagoods Mill, SC-64, 351 rd m W jct w/ Lebanon Rd., nr Toby Ck (33.21206°N, - 81.3198°W, Elev: 46 m), 17 October 2004, beneath leaf litter (Quercus, Acer) (Coll: P. Marek, D. Beamer, M. Beamer). Material examined listed in Supp Table 2 (online only). Diagnosis: Adult males of Sigmoria beameri n. sp. are distinct from other apheloriine species based on the following combination of characters: Gonopods. Gonopodal acropodite broad, curving dorsomedially at an almost 90° angle at anterior bend (Fig. 13A). Basal zone lacking spines or tubercles, prefemoral process absent. Medial flange thick, laminate. Acropodite peak broad, flattened, distally with a wide, laminate lateral flange which is distinctly constricted at its base, setting it off from its ventrally-directed tip, separating it from other species with a birdhead-like acropodal apex (= Cleptoria) Sigmoria abbotti (Hoffman, 1967), Sigmoria bipraesidens (Hoffman, 1967), Sigmoria macra (Chamberlin, 1939), Sigmoria rileyi (Bollman, 1889), Sigmoria robusta Shelley, 1986, Sigmoria shelfordi (Loomis, 1944), and Sigmoria arcuata (Shelley, 1981). Acropodite without a cingulum, as in Brachoria and Appalachioria. Color. Tergites with light-red paranotal spots (Fig. 5R). Dark brown background. Collum with light red anterior and lateral margins. Description: Supp Table 3 (online only). Based on Holotype (♂) MPE00291. Measurements (mm): BL = 47.0, CW = 6.9, IW = 4.5, ISW = 1.5, B11W = 9.4, B11H = 5.8. Variation: No significant variation from the holotype was observed. Ecology: Sigmoria beameri n. sp. was collected from the riparian area around Toby Creek and was found under oak and maple leaves. Distribution: Only known from the type locality, a small wooded area around Toby Creek, located in the Coastal Plain region of South Carolina. Etymology: This species is named for its discoverer, Dr. David Beamer of Nash Community College. The specific name is a genitive noun derived as a patronym., Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on page 16, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533, {"references":["Means, J. C., and P. E. Marek. 2017. Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae? PeerJ 5: e 3854."]}

Appalachioria n. sp. ‘Foster Falls’ Means and Marek 2017 Material examined: Type specimens. ♂ holotype (VTEC SPC000296), 1 ♂ paratype (VMNH SPC000297), and 2 ♀ paratypes (VTEC SPC000294, 295) from Virginia, Wythe County, N slope Fosters Falls Mountain, cove near road (36.89021°N, - 80.8376°W, Elev. 657 m), 29 May 2004, tulip, oak, maple, rhododendron, moist cove (Coll: P. E. Marek). Material examined listed in Supp Table 2 (online only). Diagnosis: Adult males of Appalachioria sierwaldae n. sp. are distinct from other apheloriine species based on the following combination of characters: Gonopods. Gonopodal acropodite strongly curving ventromedially and with a strong distal cingulum, separating it from Apheloria and Rudiloria (Fig. 7). Prefemoral process short and stout. Post-cingulum area expanded, wider than pre-cingulum area. Distal zone strongly curved medially into a long, uncinate, thin tip. Color. Tergites with yellow paranotal spots and yellow metatergal spots (Fig. 5I), but sometimes with orange metatergal spots (Fig. 5H). Black background. Collum with yellow lateral and anterior spots, sometimes with yellow or orange posterior spots, or lacking a posterior spot. Description: Supp Table 3 (online only). Based on Holotype (♂) SPC000296. Measurements (mm): BL = 40.0, CW = 7.0, IW = 4.7, ISW = 1.4, B11W = 9.4, B11H = 5.4. Variation: No significant variation from the holotype was observed. Ecology: Individuals of Appalachioria sierwaldae n. sp. were found in an Appalachian cove forest that included tulip, oak, maple, and rhododendron. Distribution: Only known from the type locality on Fosters Falls Mountain in eastern Wythe County, Virginia. Etymology: This species is named for Dr. Petra Sierwald of the Field Museum of Natural History. The specific name is a genitive noun derived as a matronym., Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on pages 9-11, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533, {"references":["Means, J. C., and P. E. Marek. 2017. Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae? PeerJ 5: e 3854."]}

Genus Idaloria Marek, Means, Hennen, New Genus Vernacular name: ‘The Mount Ida Millipede’ Type species: Cibularia profuga Causey, 1955. Taxa included: monotypic. Diagnosis: Adult males of Idaloria are distinct from other genera of the tribe Rhysodesmini based on the following combination of characters: Gonopods. Acropodite linear, not curved as in the Apheloriini. Acropodite distinctly narrowed at apex, forming angular shoulder. Prefemoral process acicular, longer than one-half the length of the acropodite. Acropodite without a cingulum, as in Brachoria and Appalachioria. Color. Tergites with 2 red paranotal spots. Nucleotide site substitutions. 12S: C (94); 16S: T (146, 451); G (934, 1239); T (1017, 1322); C (1043, 1348); COI: T (265, 2541); 28S: T (1293, 5644). Distribution: Idaloria profuga is known from a population in the Ouachita Mountains of Montgomery County, Arkansas and a second disjunct population in Ouachita Parish, Louisiana. Etymology: This genus is named after where the type was discovered on Mount Ida, Arkansas. The name is a combination of ‘Ida’ and the Latin - orium, ‘place for’. The ending ‘- oria ’ is commonly used for genera in the family., Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on page 15, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533

Genus Sigmoria Chamberlin, 1939. Vernacular name: ‘The Sigmoid Millipedes’ Type species: Sigmoria munda Chamberlin, 1939. Taxa included: 67 species, see Supp Material (online only) (Supp Text 1 [online only], Supp Dataset 2 [online only])., Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on page 15, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533

Brachoria camptera Means, Hennen, Marek, New Species Vernacular Name: ‘The City Lake Mimic Millipede’ Figs. 5L,M and 9. Material examined: Type specimens. ♂ holotype (VTEC MPE01398), 6 ♂ paratypes (VTEC MPE01323, 1325, 1399; VMNH MPE01403, 1410, 1416), and 2 ♀ paratypes (VTEC MPE01405, 1413) from Tennessee, Putnam County, Cookeville, City Lake Natural Area, along trail out of parking lot (36.13143°N, - 85.4439°W, Elev. 303 m), 21 May 2016, ex beech, maple, deciduous woods, tuliptree, oak, umbrella magnolia, hickory, moist litter on hillsides, understory of poison ivy, honeysuckle, mayapple (Coll: J. C. Means, D. A. Hennen). Material examined listed in Suppl Table 2 (online only). Diagnosis: Adult males of Brachoria camptera n. sp. are distinct from other apheloriine species based on the following combination of characters: Gonopods. Gonopodal acropodite D-shaped (Fig. 9A)—not circular as in Apheloria or oval as in Rudiloria species (Fig. 12). Acropodite (when viewed posteriorly) narrow, onethird width of tibia on leg pair 9. Acropodite gradually tapered to acuminate apex, not with a blunt apex as in B. hubrichti. Acropodite distal to anterior bend slightly curved cephalically. Acropodite shaft with cingulum (midlength transverse groove). Prefemur with curved sickle-shaped prefemoral process (Fig. 9B). Color. Tergites black with 2 orange (Fig. 5L), or red (Fig. 5M), paranotal spots, not with purple/purple-gray metatergal stripes as in B. hubrichti and B. initialis. Description: Supp Table 3 (online only). Based on Holotype (♂) MPE01398. Measurements (mm): BL = 39.0, CW = 7.6, IW = 5.0, ISW = 1.5, B11W = 10.0, B11H = 5.2. Variation: Individuals of Brachoria camptera n. sp. are known to exhibit two color morphs, 1) two-spotted red (Fig. 5M) and 2) two-spotted yellow-orange with a faint light blue caudal stripe (Fig. 5L). Ecology: Individuals of Brachoria camptera n. sp. were collected in a mesic deciduous forest with dominant tree species of oak, maple, beech, and tuliptree. Distribution: Known only from the type locality. Etymology: This species is named for the shape of its acropodite, specifically the angular nature of its anterior bend. The specific name is a noun in apposition derived from the Greek kampter, ‘bend, angle’., Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on pages 12-13, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533

Brachoria ‘Clinch’ (Marek and Bond 2006) Appalachioria ‘n. sp. Clinch Mountain’ (Marek and Bond 2007, Means and Marek 2017) Material examined: Type specimens. ♂ holotype (VTEC SPC000282), ♂ paratype (VMNH SPC000280), and ♀ paratypes (VTEC SPC000281, 283) from Virginia, Washington County, Clinch Mountain, CR-612 (36.72317°N, - 82.2985°W, Elev. 635 m), 28 May 2004, 15:00, tulip, birch, oak, sassafras (Coll: P. E. Marek). Material examined listed in Supp Table 2 (online only). Diagnosis: Adult males of Appalachioria bondi n. sp. are distinct from other apheloriine species based on the following combination of characters: Gonopods. Gonopodal acropodite curving ventromedially at apex, with a distal cingulum, separating it from Apheloria and Rudiloria (Fig. 6). Prefemoral process short and stout, basal zone lacking tubercles. Acropodite at anterior bend with three stout, triangular dorsal tubercles, separating it from most other Appalachioria species. Distal zone constricted basally, with a medially curved, blunt, uncinate tip. Color. Appalachioria bondi n. sp. has multiple color morphs, often co-occurring (Fig. 5A–G). Tergites always with white, light-pink, or yellow paranotal spots, sometimes also with concolorous metatergal middorsal spots. Dark-brown to black background. Collum always with concolorous white, light-pink, or yellow anterior and lateral spots, sometimes also with a posterior spot, never with marginal lines connecting the spots. Description: Supp Table 3 (online only). Based on Holotype (♂) SPC000282. Measurements (mm): BL = 42.0, CW = 8.5, IW = 5.2, ISW = 1.4, B11W = 9.9, B11H = 5.8. Variation: The dorsal tubercles on the anterior bend of the acropodite vary in size and number, ranging from 1 to 3 (exceptionally up to 10) small to stout triangular tubercles, sometimes serrated or divided apically. The tubercles can vary from dorsal to medial in position. The tip of the acropodite may be curved medially to ventrally. The prefemoral process ranges from short and thin to long and stout and varies from small to medium in size. Appalachioria bondi n. sp. exhibits six unique color patterns: yellow four-spotted collum (Fig. 5A), red twospotted, collum with one anterior and two lateral red spots (Fig. 5B), yellow two-spotted, collum with one anterior and two lateral yellow spots (Fig. 5E), white four-spotted collum (Fig. 5C), white two-spotted, collum with one anterior and two lateral white spots (Fig. 5F) and red four-spotted collum (Fig. 5D). Ecology: Individuals of Appalachioria bondi n. sp. have been found in mesic deciduous forests of oak, beech, maple, tuliptree, birch, buckeye, and sassafras, with small patches of eastern redcedar nearby. Distribution: Appalachioria bondi n. sp. is only known from a small range of about 8 km 2 on Clinch Mountain in Russell and Washington Counties, Virginia. Etymology: This species is named for Dr. Jason Bond of the University of California, Davis. The specific name is a genitive noun derived as a patronym., Published as part of Means, Jackson C., Hennen, Derek A., Tanabe, Tsutomu & Marek, Paul E., 2021, Phylogenetic Systematics of the Millipede Family Xystodesmidae, pp. 1-26 in Insect Systematics and Diversity 5 (2) on page 9, DOI: 10.1093/isd/ixab003, http://zenodo.org/record/5384533, {"references":["Marek, P. E., and J. E. Bond. 2006. Phylogenetic systematics of the colorful, cyanide-producing millipedes of Appalachia (Polydesmida, Xystodesmidae, Apheloriini) using a total evidence Bayesian approach. Mol. Phylogenet. Evol. 41: 704 - 729.","Marek, P. E., and J. E. Bond. 2007. A reassessment of apheloriine millipede phylogeny: additional taxa, Bayesian inference, and direct optimization (Polydesmida: Xystodesmidae). Zootaxa 1610: 27 - 39.","Means, J. C., and P. E. Marek. 2017. Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae? PeerJ 5: e 3854."]}

Millipedes (class Diplopoda) are ubiquitous in forests worldwide, with about 12,000 described species and an estimated 30,000 undescribed species. The most species-rich order is the Polydesmida, the flat-backed millipedes, which encompasses about 3,500 species. Appalachia has an abundance of Polydesmida, and is a biodiversity hotspot for millipedes in the family Xystodesmidae. These diplopods are chemically defended with hydrogen cyanide and benzaldehyde, and sometimes form mimicry rings based on shared color. The evolutionary dynamics of model and mimic are incompletely known in these rings, so I investigated a mimicry ring in the central Appalachian Mountains to determine if the species Apheloria polychroma functions as a model for the genus Brachoria. I measured the size of the chemical gland to assess toxicity in 15 species, and reconstructed the ancestral dimensions of the gland to determine the direction of volume change over evolutionary time. Using a molecular phylogeny, I traced the miniaturization of chemical glands in the mimic genus Brachoria and found that in areas without Apheloria, Brachoria and related xystodesmids have larger chemical glands. Non-aposematic millipedes of the genus Nannaria have significantly smaller glands, and ostensibly rely on camouflage to avoid predation. This genus is known as the twisted claw millipedes and occur throughout eastern North America, but have their center of diversity in the Appalachian Mountains. About 22 species are described, but many undescribed species are known. To determine the diversity of this group, field collection and examination of museum specimens took place from 2015-2020. Examination of morphology, combined with molecular phylogenetics, revealed two distinct clades in the genus. One is distributed throughout the eastern United States, while the other is found only in the Appalachian Mountains. This Appalachian clade contains six described species, and I describe an additional 18 species, quadrupling the diversity of the group. Additionally, the phylogenetics of the polydesmid genus Pseudopolydesmus is investigated in an integrative taxonomic framework using five genes. I find that the genus is monophyletic, contains 8 species, and are related to one another in a hierarchical way according to a molecular phylogeny. I provide complete distributional records and live photographs of each species. Doctor of Philosophy Millipedes are common animals in forests. There are about 12,000 known species in the world, with an estimated 30,000 undescribed species still awaiting discovery and description. The largest group of millipedes are the flat-backed millipedes, with about 3,500 known species. They are some of the most common millipedes in North America, and many of these species defend themselves with poisons that are harmful to predators, but smell sweet like cherries to humans. Some of these millipedes have bright red or yellow spots against dark colors to warn predators of their toxins, and look similar to other species that live near them. I wanted to know if some of these species are more or less poisonous than others, and measured how large their poison reserves were. I found that one species, called Apheloria polychroma, is more poisonous than similar-looking species called Brachoria, which are less poisonous. If Brachoria doesn't live near Apheloria though, Brachoria is more poisonous. Not all of these millipedes are brightly colored, and a group called Nannaria, or the twisted claw millipedes, are camouflaged with brown colors on the forest floor. These millipedes only live in the eastern United States, especially the Appalachians, and aren't found anywhere else in the world. We know about 20 species of them, but based on specimens stored in scientific collections in museums and through discovery from fieldwork, we knew that more species existed. Each species of twisted-claw millipede only lives in a small area, sometimes only a few miles wide, and could be threatened by habitat loss and other dangers. So, to learn more about them, we need to find them in the forest and describe what they look like and most importantly, giving them a name. I did this by collecting them, illustrating their anatomy, and sequencing their DNA. I found two groups within Nannaria, and focused on the group that only lives in Appalachia. Scientists know 6 of these species, but I found 18 more species and describe them. A related group called Pseudopolydesmus lives throughout North America, and I studied their anatomy and DNA as well, finding 8 species.

Genus Apheloria Chamberlin, 1921 Type species: Fontaria montana Bollman, 1887., Published as part of Marek, Paul E., Means, Jackson C. & Hennen, Derek A., 2018, Apheloria polychroma, a new species of millipede from the Cumberland Mountains (Polydesmida: Xystodesmidae), pp. 409-425 in Zootaxa 4375 (3) on page 415, DOI: 10.11646/zootaxa.4375.3.7, http://zenodo.org/record/1158980, {"references":["Chamberlin, R. V. (1921) On some chilopods and diplopods from Knox Co., Tennessee. Canadian Entomologist, 53, 230 - 233. https: // doi. org / 10.4039 / Ent 53230 - 10","Bollman, C. H. (1887) Description of fourteen new species of North American myriapods. Proceedings of the United States National Museum, 10, 617 - 627. https: // doi. org / 10.5479 / si. 00963801.10 - 670.617"]}

Apheloria polychroma, new species Vernacular name: ���The Colorful Cherry Millipede��� Figs 1���6. Apheloria ���flavissima��� (Hobson 2010), Apheloria ���Stone��� (Marek & Bond 2007; Means & Marek 2017), Apheloria ���Clade A��� (Marek & Bond 2009). Material examined: Type specimens. ♂ holotype (FMNH-INS60792), 1 ♀ paratype (FMNH-INS71228), 3 ♂, 3 ♀ paratypes (VTEC, MMC0309, 310, 313, 305, 306, 308), 1 ♂, 1 ♀ paratypes (VMNH, MMC0314, 312) from Virginia, Lee County, The Cedars, CR-738 (36.65624��N, - 83.20165��W, Elev. 436 m), 28 September 2006, 16:00 (Colls: P. and B. Marek). Non-type material examined listed in Table 1. Diagnosis. Adult males of A. polychroma are distinct from other apheloriine species based on the following combination of characters: Color. Tergites usually with 4 yellow spots (i.e. 1 metatergal, 1 prozonal, 2 paranotal spots, Figs 2A���F, M���P). Collum usually uniformly yellow (1.25Y 7/14). Jet-black background (N 0.5���0.75). Gonopods. Gonopodal acropodite smoothly circular, O-shaped (Figs 3A, B)���without elbow as in A. tigana (Fig. 4C, arrow). Acropodite narrow, half width of tibia on leg pair 9, of uniform width throughout. Acropodite gradually tapered to curved acuminate J-shaped apex, not L-shaped and abruptly twisted as in A. montana, A. tigana, A. virginiensis corrugata (Figs 3C; 4B, D, F). Acropodite shaft without cingulum (midlength transverse groove) or preapical teeth or projections as in Brachoria. Prefemur with a thin sharp prefemoral process, one-ninth length of acropodite (Fig. 3A, arrow), not long and scythe-like as in A. montana, A. tigana, A. virginiensis corrugata (Figs 4B, D, F). Note: The 4-spotted yellow color morph with a uniformly yellow collum unambiguously diagnoses A. polychroma (Fig. 2A, C, E). However, there are at least 6 color morphs of the species with a substantial continuum of hues and patterns among them (Figs 2, 5). There are frequently several color morphs of the species at a single locality, and as color varies considerably, it should be used with caution as a diagnostic character for this species. Description. Based on Holotype (♂) FMNH-INS 60792. Measurements: BL = 48.70, CW = 7.50, IW = 4.10, ISW = 1.00, B11W = 9.50, B11H = 5.40. Head: Antennae extending posteriorly to anterior margin of 3rd tergite, relative antennomere lengths 2>5>3>4>6>1>7. Antennae with 4 sensillum types, sensillum shafts smooth without barbules. Four apical cones (AS) in square cluster on 7th antennomere. Chaetiform sensilla (CS) on antennomeres 5���7. Antennomeres 1���4 smooth with sparse vestiture of CS. Trichoid sensilla (TS) on antennomeres 1���7, coronally encircling apex. Spiniform basiconic sensilla (Bs2) in clusters of 15 on apical dorsal (retrolateral) surface of antennomeres 5, 6; Bs2 shaft 1/10 length of CS. Tergites: Collum with straight cephalic edge, abruptly tapering laterally. Collum with carina present on anterolateral margins, absent medially. Caudolateral corners of paranota rounded cephalically on body rings 1���4; acute, projecting caudally on body rings 5���19. Caudolateral corners of paranota 7���19 with small posteriorprojecting nubbin. Paranota dorsal surface loosely wrinkled, appearing leathery. Ozopores oriented dorsolaterally. Pore formula normal: 5, 7, 9, 10, 12, 13, 15���19. Paranota with anterodorsal region scooped out. Gonapophyses cylindrical apically. Pleural tubercle absent, with slight swelling between paranotal base and spiracle. Sterna without posteriorly projecting spines, with slightly curved caudal margin. Sterna 2���11 sparsely setose (ca. 10 setae), sterna 12���18 lacking setae, ventral surface smooth. Gonopods: In situ configuration���Acropodite base projected posteroventrally, curved anteromedially in smooth O-shaped form (Fig. 3A, B). Terminal arc of acropodite oriented ventrolaterally. O-shaped arc nearly closed, gap 1/8 length of circumference between prefemoral process and apex (Fig. 3A, B). Right, left acropodites crossed midlength, appearing intertwined. Gonocoxae with conical protuberance apically, telopodites arising subapically. Telopodites���Prefemur with thin sharp prefemoral process, one-ninth length of acropodite, with apex tapered to sharp curved point, sickle-shaped, width at base 1/4 its length (Fig. 3A, arrow). Acropodite bent 90�� posteriorly at prefemur (Fig. 3C, arrow), prostatic groove arcs 90�� from cannula to acropodite base (Fig. 3B, arrows). Gonopodal acropodite narrow, half width of tibia on leg pair 9, tapered to curved acuminate apex. Acropodite with ventrobasal surface facing anterolaterally. Acropodite base without spines on dorsal surface. Acropodal basal and apical ventral surfaces not coplanar, apical surface facing ventrolaterally; anterior twist absent (Fig. 3C). Acropodal ventral surface flat, not swollen, smooth. Acropodite without midlength transverse groove, or cingulum. Acropodite shaft of uniform width, tapered to acuminate apex distal to region with setae. Acropodite elliptical in cross-section, tapered acuminate apex thinner, transparent. Acropodite margin rounded, lacking sharp edge; marginal teeth absent. Apical 1/9 of acropodite recurved, J-shaped, projecting cephalically (Fig. 3C). Acropodite region with setae about 1/3 its total length (Fig. 3B). Paratype (♀) FMNH-INS71228���Measurements: BL = 50.30, CW = 8.00, IW = 4.80, ISW = 1.40, B10W = 11.81, B10H = 7.10. Cyphopods: Cyphopod receptacle (at its greatest breadth) equal in width to prefemur length. Receptacle heartshaped, pointed end facing medially. Cyphopodal valves symmetric, anterior valve slightly more convex. Cyphopods with valvular suture facing laterally, without midlength ramp-like swell. Variation. Apheloria polychroma is known from the Cumberland Mountain Thrust Block region at the confluence of southwestern Virginia, northeastern Tennessee, and southeastern Kentucky (Fig. 6). There are at least six color morphs with substantial variation in coloration between them: (1) four-spotted, with yellow to orange paranotal, metatergal, prozonal and collum spots, and legs and collum often completely yellow and sometimes with red legs (Figs 2A���F, M���O); (2) striped, with yellow metatergal, paranotal, anterior collum stripes, and legs (Figs 2G, H); (3) three-spotted, with creamy white paranotal, metatergal and collum spots, and red legs (Figs 2I, J); (4) three-spotted, with yellow paranotal, metatergal and collum spots, and red or yellow legs (Figs 2K, L; 5A, B); (5) striped/four-spotted mix, superimposition of striped and four-spotted yellow morphs (Figs 2M, N); and (6) twospotted, with yellow paranotal spots, and red or yellow legs (Figs 2Q���T). Some three-spotted yellow individuals possess faint metatergal spots, appearing nearly two-spotted (Fig. 2S). The dorsal color of A. polychroma is always yellow and black; however, the pattern varies from two, three, or four maculae, to metatergal stripes. The hue consists of variations of yellow: from light cream-white to brilliantly saturated yellow. Based on the molecular phylogeny of populations of A. polychroma (Fig. 1), some individuals that share a uniform mitochondrial 16S sequence possess three distinct color morphs (morphs 1, 2 and 5 from above). The color morphs lack phylogenetic conservation, whereby individuals sharing the same aposematic appearance are not monophyletic (Fig. 1). Evolutionary shifts between colors, based on reconstructing color morphs on a phylogeny using parsimony, occurred frequently with ca. 29 changes (Fig. 1). Brachoria dentata Keeton, 1959 mimics A. polychroma in color and pattern at six of the seven localities where they were found to co-occur. The other five species of Brachoria cooccur at fewer localities. There is typical sexual size dimorphism between males and females, and negligible variation of measurements within sex. Measurements: ♂ (n = 10) BL = 37.80���49.45 (45.68/3.45). CW = 6.90���8.80 (7.58/0.63). IW = 4.10���5.00 (4.63/0.30). ISW = 1.00���1.40 (1.20/0.13). B10W = 8.80���10.79 (10.11/0.82). B10H = 4.60���6.10 (5.66/0.65). ♀ (n = 7) BL = 41.60���50.30 (46.67/3.07). CW = 6.60���8.30 (7.63/0.59). IW = 4.70���5.40 (5.01/0.29). ISW = 1.20���1.40 (1.31/0.09). B10W = 9.10���11.81 (10.64/0.96). B10H = 6.10���7.10 (6.4/0.37). Ecology. Apheloria polychroma individuals were collected in mesic broadleaf deciduous forests and occasionally in more xeric glades, for example 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 encountered beneath decomposing leaves and walking atop detritus on the forest floor. Individuals of A. polychroma were more often exposed to view than other apheloriines, bolder in behavior, and were more likely to thrash when disturbed. When handled, individuals would emit copious amounts of defense secretions. Due to similarity in color patterns and overlap in distribution, A. polychroma may be confused in the field with six species of Brachoria, which include Brachoria dentata Keeton, 1959; Brachoria cedra Keeton, 1959; Brachoria hoffmani Keeton, 1959; Brachoria insolita Keeton, 1959; Brachoria mendota Keeton, 1959; and Brachoria sheari Marek, 2010. Mimetic resemblance between individuals of A. polychroma and B. mendota at Natural Tunnel State Park (Scott County, Virginia) is one of the most accurate in the mimicry system (Fig. 5). Apheloria polychroma can be distinguished from these taxa in the gonopod morphology, e.g. by the absence of a cingulum, or a midlength transverse groove on the acropodite, and presence of an O-shaped telopodite (Fig. 3A, B). Distribution. Known from southwestern Virginia (Lee, Scott, Norton, and Wise Counties), southeastern Kentucky (Bell and Harlan Counties), and northeastern Tennessee (Campbell, Claiborne, and Hancock Counties, Fig. 6). The distribution of A. polychroma is coincident with the boundaries of the High Knob Landform, and more broadly the Cumberland Mountain Thrust Block (Rich 1934; Browning 2009). Notes. Apheloria polychroma was previously known by the informal names Apheloria ��� flavissima ��� (Hobson, 2010), Apheloria ���Stone��� (Marek & Bond 2007; Means & Marek 2017), and Apheloria ���Clade A��� (Marek & Bond 2009). Etymology. This species is named for its extremely variable coloration. The specific name is a noun in apposition derived from the Greek polu-, ���many���, and khrōma, ���color���., Published as part of Marek, Paul E., Means, Jackson C. & Hennen, Derek A., 2018, Apheloria polychroma, a new species of millipede from the Cumberland Mountains (Polydesmida: Xystodesmidae), pp. 409-425 in Zootaxa 4375 (3) on pages 416-420, DOI: 10.11646/zootaxa.4375.3.7, http://zenodo.org/record/1158980, {"references":["Hobson, C. S. (2010) Powell Mountain Karst Preserve: biological inventory of vegetation communities, vascular plants, and selected animal groups. Natural Heritage Technical Report 10 - 12. Virginia Department of Conservation and Recreation, Division of Natural Heritage, Richmond, Virginia, 30 pp., appendices. [unpublished report submitted to The Cave Conservancy of the Virginias, April 2010]","Marek, P. E. & Bond, J. E. (2007) A reassessment of apheloriine millipede phylogeny: additional taxa, Bayesian inference, and direct optimization (Polydesmida: Xystodesmidae). Zootaxa, 1610, 27 - 39.","Means, J. C. & Marek, P. E. (2017) Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae? PeerJ, 5, e 3854. https: // doi. org / 10.7717 / peerj. 3854","Marek, P. E. & Bond, J. E. (2009) A Mullerian mimicry ring in Appalachian millipedes. Proceedings of the National Academy of Sciences, 106, 9755 - 9760. https: // doi. org / 10.1073 / pnas. 0810408106","Keeton, W. T. (1959) A revision of the millipede genus Brachoria (Polydesmida: Xystodesmidae). Proceedings of the United States National Museum, 109, 1 - 58. https: // doi. org / 10.5479 / si. 00963801.109 - 3411.1","Marek, P. E. (2010) A revision of the Appalachian millipede genus Brachoria Chamberlin, 1939 (Polydesmida: Xystodesmidae: Apheloriini). Zoological Journal of the Linnean Society, 159, 817 - 889. https: // doi. org / 10.1111 / j. 1096 - 3642.2010.00633. x","Rich, J. L. (1934) Mechanics of low-angle overthrust faulting as illustrated by Cumberland Thrust Block, Virginia, Kentucky, and Tennessee. Bulletin of the American Association of Petroleum Geologists, 18, 1584 - 1596.","Browning, W. (2009) The High Knob Landform. Available from: http: // www. highknoblandform. com (accessed 31 December 2017)"]}

Mimuloria rhysodesmoides, new species Fig. 22–23. Type specimens. Male holotype and five fragmented male paratypes (FSCA) collected by H. E. Evans, 12 April 1958, in Cookeville, Putnam Co., Tennessee. One male paratype retained at NCSM. Diagnosis. Acropodite subupright, anterior surface with subapical and subacuminate spur; prefemoral process extending for 1/2 – 2/3 of acropodital length, narrowing strongly apically, angling inward into acropodital arc and directed toward latter’s stem. Holotype. Length ca. 18.4 mm, maximum width ca. 4.4 mm, W/L ratio 23.9%. Somatic features agreeing closely with those of M. castanea with following exceptions. Width across genal apices 1.4 mm. Facial setae as follows: epicranial, interantennal, subantennal, frontal, and genal 0–0, clypeal about 6–6, labral about 8–8, merging with clypeal series and continuing for short distances along genal borders. Antennae reaching back to midlength of 4 th metatergite; relative lengths of antennomeres 6>2=3>4=5>1>7. Caudolateral corners of 2 nd – 4 th paranota rounded; those of 5 th paranota slightly extended and becoming more so on caudal paranota. Sternal spines present on segments 8–19. Relative lengths of postgonopodal podomeres 3>2>4>6>5>1. Coxae through tibiae only sparsely hirsute; prefemoral spines long, overhanging ½ or more of lengths of femora. Tarsal claws spatulate on legs 2–18 (anterior legs of 10 th segment). Gonopods in situ crossing around midlengths, extending anteriad to or beyond midlength of 5 th sternum. Gonopod structure (Fig. 22–23) as follows: Prefemoral process long, broad for most of length, narrowing distad, directed inward toward 2/3 length of acropodite, without trace of secondary process, spur, or lobe. Acropodite leaning mediad for most of length, bending more so distad and overhanging tip of prefemoral process; blade-like and subequal in width for most of length, narrowing at distal bend and more so apically, with subapical spur on outer margin distal to bend. Etymology. The specific name references the apical similarity to the Mexican xystodesmid genus Rhysodesmus Cook, 1895 (Rhysodesmini) in possessing the spur on the outer acropodital surface (Hoffman 1966, 1970). Rhysodesmus also includes two Appalachian species, R. restans Hoffman 1998, from Washington Co., Virginia, and R. agrestis Shelley, 1999, from Knox Co., Tennessee (Hoffman 1998, Shelley 1999). Distribution. Known only from the type locality. Remarks. Though depigmented after 56 years in preservative, the types show an unequivocal banded color pattern. Broad, light bands extend along caudal metatergal margins connecting equally light paranota; the base color appears to be dark olive., Published as part of Hennen, Derek A. & Shelley, Rowland M., 2015, A contribution on the milliped tribe Nannariini (Polydesmida: Xystodesmidae): Revalidation of Mimuloria Chamberlin 1928; identities of Fontaria oblonga C. L. Koch 1847, and Nannaria minor Chamberlin 1918; elucidation of the tribal range; and commentaries on Nannaria Chamberlin 1918, and Oenomaea Hoffman 1964, pp. 1-21 in Insecta Mundi 2015 (418) on pages 15-16, DOI: 10.5281/zenodo.5182058, {"references":["Hoffman, R. L. 1966. The Mexican genera of Xystodesmidae. Transactions of the American Entomological Society 92: 1 - 16.","Hoffman, R. L. 1970. Random studies on Rhysodesmus. I. Notes and redescriptions of miscellaneous species. The Radford Review 24: 143 - 162.","Hoffman, R. L. 1998. An Appalachian species of Rhysodesmus (Polydesmida: Xystodesmidae: Rhysodesmini). Myriapodologica 5: 77 - 83.","Shelley, R. M. 1999. A second east-Nearctic species of Rhysodesmus Cook (Polydesmida: Xystodesmidae). Myriapodologica 6: 19 - 22."]}

Hennen, Derek A., Shelley, Rowland M. (2015): A contribution on the milliped tribe Nannariini (Polydesmida: Xystodesmidae): Revalidation of Mimuloria Chamberlin 1928; identities of Fontaria oblonga C. L. Koch 1847, and Nannaria minor Chamberlin 1918; elucidation of the tribal range; and commentaries on Nannaria Chamberlin 1918, and Oenomaea Hoffman 1964. Insecta Mundi 2015 (418): 1-21, DOI: http://doi.org/10.5281/zenodo.5182058, {"references":["Attems, C. 1899. System der Polydesmiden, II. Theil. Denkschriften der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften 68: 1-185.","Attems, C. 1937. Myriapoda 3 Polydesmoidea I. Fam. Strongylosomidae. Das Tierreich 68: 1-300 + i-xxii.","Attems, C. 1938. Myriapoda 3 Polydesmoidea II. Fam. Leptodesmidae, Platyrhachidae, Oxydesmidae, Gomphodesmidae. Das Tierreich 69: 1-487 + i-xxviii.","Bollman, C. H. 1889a. Description of a new species of insect, Fontaria pulchella, from Strawberry Plains, Jefferson County, Tennessee. Proceedings of the United States National Museum 11: 316.","Bollman, C. H. 1889b. Notes on a collection of Myriapoda from Mossy Creek Tenn., with a description of a new species. Proceedings of the United States National Museum 11: 339-342.","Bollman, C. H. 1893. The Myriapoda of North America. Bulletin of the United States National Museum 46: 1-210.","Causey, N. B. 1942. Six new diplopods of the family Xystodesmidae. Entomological News 53: 165-170.","Causey, N. B. 1950a. On four new polydesmoid millipeds. Entomological News 61: 193-198.","Causey, N. B. 1950b. A new genus and species of diplopod (Family Xystodesmidae). Chicago Academy of Sciences Natural History Miscellanea 71: 1-3.","Causey, N. B. 1950c. Variations in the gonopods of a xystodesmid diplopod. American Midland Naturalist 44: 198-202.","Causey, N. B. 1952. Some records and descriptions of polydesmoid millipeds from the United States. Chicago Academy of Sciences Natural History Miscellanea 106: 1-11.","Causey, N. B. 1955. New records and descriptions of polydesmoid millipeds (order Polydesmida) from the eastern United States. Proceedings of the Biological Society of Washington 68: 21-30.","Chamberlin, R. V. 1918a. New polydesmoid diplopods from Tennessee and Mississippi. Psyche 25: 122-127.","Chamberlin, R. V. 1918b. Myriapods from Nashville, Tennessee. Psyche 25(2): 23-30.","Chamberlin, R. V. 1928. Some chilopods and diplopods from Missouri. Entomological News 39: 153-155.","Chamberlin, R. V. 1949. A new genus and four new species in the diplopod family Xystodesmidae. Proceedings of the Biological Society of Washington 62: 3-6.","Chamberlin, R. V., and R. L. Hoffman. 1958. Checklist of the millipeds of North America. Bulletin of the United States National Museum 212: 1-236.","Hoffman, R. L. 1948. Three new eastern millipeds of the family Xystodesmidae. Journal of the Washington Academy of Sciences 38: 346-350.","Hoffman, R. L. 1949. Nine new xystodesmid millipeds from Virginia and West Virginia, with records of established species. Proceedings of the United States National Museum 99: 371-389.","Hoffman, R. L. 1964. The status of Fontaria pulchella Bollman, with the proposal of a new genus and tribe in the diplopod family Xystodesmidae. Proceedings of the Biological Society of Washington 77: 25-34.","Hoffman, R. L. 1966. The Mexican genera of Xystodesmidae. Transactions of the American Entomological Society 92: 1-16.","Hoffman, R. L. 1970. Random studies on Rhysodesmus. I. Notes and redescriptions of miscellaneous species. The Radford Review 24: 143-162.","Hoffman, R. L. 1980 (1979). Classification of the Diplopoda. Museum d'Histoire Naturelle; Geneve, Switzerland. 237 p.","Hoffman, R. L. 1998. An Appalachian species of Rhysodesmus (Polydesmida: Xystodesmidae: Rhysodesmini). Myriapodologica 5: 77-83.","Hoffman, R. L. 1999. Checklist of the millipeds of North and Middle America. Virginia Museum of Natural History Special Publication Number 8: 1-584.","Holt, S. B. 1976. Harold Cummins (1894-1976). Journal of Medical Genetics 13: 540.","Jeekel, C. A. W. 1971. Nomenclator generum et familiarum Diplopodorum: A List of the genus and family-group names in the Class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Monografieen van de Nederlandse Entomologische Vereniging 5: 1-412 + i-xii.","Kevan, D. K. McE. 1983. A preliminary survey of known and potentially Canadian millipedes (Diplopoda). Canadian Journal of Zoology 61: 2956-2975.","Koch, C. L. 1847. System der Myriapoden. In: Herrich-Schaffer and Panzer. Kritische Revision der Insectenfauna Deutschlands 3: 1-190.","Koch, C. L. 1863. Die Myriapoden. Getrau nac der Natur abgebildet und beschrieben 1: 1-134.","Loomis, H. F., and R. L. Hoffman. 1948. Synonymy of various diplopods. Proceedings of the Biological Society of Washington 61: 51-54.","Marek, P. E., and J. E. Bond. 2006. Phylogenetic systematics of the colorful, cyanide-producing millipedes of Appalachia (Polydesmida, Xystodesmidae, Apheloriini) using a total evidence Bayesian approach. Molecular Phylogenetics and Evolution 41: 704-729.","Marek, P. E., and J. E. Bond. 2007. A reassessment of apheloriine millipede phylogeny: additional taxa, Bayesian inference, and direct optimization (Polydesmida: Xystodesmidae). Zootaxa 1610: 27-39.","Marek, P., T. Tanabe, and P. Sierwald. 2014. A species catalog of the millipede family Xystodesmidae (Diplopoda: Polydesmida). Virginia Museum of Natural History Special Publication Number 17: 1-117 + i-xxii.","McNeill, J. 1887. Descriptions of twelve new species of Myriapoda, chiefly from Indiana. Proceedings of the United States National Museum 10: 328-334.","Shelley, R. M. 1975. The identity of Nannaria conservata Chamberlin, with notes on an abnormal male and descriptions of two new species of Nannaria from North Carolina (Diplopoda: Polydesmida: Xystodesmidae). Proceedings of the Biological Society of Washington 88: 179-188.","Shelley, R. M. 1978. Millipeds of the eastern Piedmont region of North Carolina, USA (Diplopoda). Journal of Natural History 12: 37-79.","Shelley, R. M. 1980. Revision of the milliped genus Pleuroloma (Polydesmida: Xystodesmidae). Canadian Journal of Zoology 58: 129-168.","Shelley, R. M. 1981. Revision of the milliped genus Sigmoria (Polydesmida: Xystodesmidae). Memoirs of the American Entomological Society 33: 1-141.","Shelley, R. M. 1988. The millipeds of eastern Canada (Arthropoda: Diplopoda). Canadian Journal of Zoology 66: 1638-1663.","Shelley, R. M. 1996. Revision of the milliped genus Xystocheir Cook (Polydesmida: Xystodesmidae). Canadian Journal of Zoology 74: 1336-1363.","Shelley, R. M. 1999. A second east-Nearctic species of Rhysodesmus Cook (Polydesmida: Xystodesmidae). Myriapodologica 6: 19-22.","Shelley, R. M. 2000. Annotated checklist of the millipeds of North Carolina (Arthropoda: Diplopoda), with remarks on the genus Sigmoria Chamberlin (Polydesmida: Xystodesmidae). Journal of the Elisha Mitchell Scientific Society 116: 177-205.","Shelley, R. M. 2002. The millipeds of central Canada (Arthropoda: Diplopoda), with reviews of the Canadian fauna and diplopod faunistic studies. Canadian Journal of Zoology 80: 1863-1875.","Shelley, R. M. 2014. A consolidated account of the polymorphic Caribbean milliped, Anadenobolus monilicornis (Porat, 1876) (Spirobolida: Rhinocricidae), with illustrations of the holotype. Insecta Mundi 0378: 1-12.","Shelley, R. M., and C. T. McAllister. 2007a. Composition and distribution of the milliped tribe Pachydesmini west of the Mississippi River (Polydesmida: Xystodesmidae). Western North American Naturalist 66: 45-54.","Shelley, R. M., and C. T. McAllister. 2007b. Distribution of the milliped genus Apheloria Chamberlin, 1921; summaries of peripheral localities and ones of A. virginiensis (Drury, 1770) west of the Mississippi River (Polydesmida: Xystodesmidae). Western North American Naturalist 67: 258-269.","Shelley, R. M., and D. R. Whitehead. 1986. A reconsideration of the milliped genus Sigmoria, with a revision of Deltotaria and an analysis of the genera in the tribe Apheloriini (Polydesmida: Xystodesmidae). Memoirs of the American Entomological Society 35: 1-223.","Shelley, R. M., P. Sierwald, S. B. Kiser, and S. I. Golovatch. 2000. Nomenclator generum et familiarum Diplopodorum II, a list of the genus and family-group names in the class Diplopoda from 1958 through 1999. Pensoft Publishers; Sofia, Bulgaria. 167 p.","Shelley, R. M., C. T. McAllister, and S. B. Smith. 2004. Discovery of the milliped Pleuroloma flavipes (Polydesmida: Xystodesmidae) in Texas, and other records from west of the Mississippi River. Entomological News 114: 2-6.","Shelley, R. M., C. T McAllister, C. M Nagy, M. E. Weckel, R. G. Christie, P. Wilson, and A. Wilson. 2011. Distribution of the American milliped genus Boraria Chamberlin, 1943: Introductions of B. stricta (Brolemann, 1896) in New York and B. infesta (Chamberlin, 1918) in Connecticut; indigenous occurrence of B. profuga (Causey, 1955) in Louisiana (Diplopoda: Polydesmida: Xystodesmidae). Insecta Mundi 0194: 1-8.","Snyder, B. A. 2008. A preliminary checklist of the millipedes (Diplopoda) of the Great Smoky Mountains National Park, USA. Zootaxa 1836: 16-32.","Williams, S. R., and R. A. Hefner. 1928. The millipedes and centipedes of Ohio. The Ohio State University Bulletin 33[Ohio Biological Survey Bulletin 4]: 91-147."]}

Genus Nannaria Chamberlin 1918a Nannaria Chamberlin 1918a: 124. Loomis and Hoffman 1948: 53. Hoffman 1949: 381; 1964: 33; 1980: 159; 1999: 365. Chamberlin and Hoffman 1958: 39–40. Jeekel 1971: 274. Kevan 1983: 2968. Shelley 2000: 196. Marek et al. 2014: 36. Type-species. Nannaria minor Chamberlin 1918a, by original designation. Remarks. Nannaria occurs from westcentral New York and southeastern Virginia to western Ohio and perhaps eastcentral Kentucky / Tennessee and northern Alabama / Georgia, and presently comprises 17 species. However, N. ericacea and N. scutellaria, for example, do not seem congeneric with N. minor and appear to require new genera, and numerous new and undiscovered species and genera undoubtedly await discovery in both the field and preserved holdings. A revision is needed but can only derive as part of a comprehensive tribal review, which is well beyond our scope. We note, however, that the holotypes of the two most important components – N. minor, the type-species, and N. oblonga, the oldest name and thus holding priority over all others – have never been illustrated, and knowledge of their gonopods is essential for generic and tribal revisions to advance. We therefore publish such (Fig. 24–25) to eliminate this void and facilitate future research., Published as part of Hennen, Derek A. & Shelley, Rowland M., 2015, A contribution on the milliped tribe Nannariini (Polydesmida: Xystodesmidae): Revalidation of Mimuloria Chamberlin 1928; identities of Fontaria oblonga C. L. Koch 1847, and Nannaria minor Chamberlin 1918; elucidation of the tribal range; and commentaries on Nannaria Chamberlin 1918, and Oenomaea Hoffman 1964, pp. 1-21 in Insecta Mundi 2015 (418) on page 16, DOI: 10.5281/zenodo.5182058, {"references":["Chamberlin, R. V. 1918 a. New polydesmoid diplopods from Tennessee and Mississippi. Psyche 25: 122 - 127.","Loomis, H. F., and R. L. Hoffman. 1948. Synonymy of various diplopods. Proceedings of the Biological Society of Washington 61: 51 - 54.","Hoffman, R. L. 1949. Nine new xystodesmid millipeds from Virginia and West Virginia, with records of established species. Proceedings of the United States National Museum 99: 371 - 389.","Hoffman, R. L. 1964. The status of Fontaria pulchella Bollman, with the proposal of a new genus and tribe in the diplopod family Xystodesmidae. Proceedings of the Biological Society of Washington 77: 25 - 34.","Hoffman, R. L. 1980 (1979). Classification of the Diplopoda. Museum d'Histoire Naturelle; Geneve, Switzerland. 237 p.","Hoffman, R. L. 1999. Checklist of the millipeds of North and Middle America. Virginia Museum of Natural History Special Publication Number 8: 1 - 584.","Chamberlin, R. V., and R. L. Hoffman. 1958. Checklist of the millipeds of North America. Bulletin of the United States National Museum 212: 1 - 236.","Jeekel, C. A. W. 1971. Nomenclator generum et familiarum Diplopodorum: A List of the genus and family-group names in the Class Diplopoda from the 10 th edition of Linnaeus, 1758, to the end of 1957. Monografieen van de Nederlandse Entomologische Vereniging 5: 1 - 412 + i - xii.","Kevan, D. K. McE. 1983. A preliminary survey of known and potentially Canadian millipedes (Diplopoda). Canadian Journal of Zoology 61: 2956 - 2975.","Marek, P., T. Tanabe, and P. Sierwald. 2014. A species catalog of the millipede family Xystodesmidae (Diplopoda: Polydesmida). Virginia Museum of Natural History Special Publication Number 17: 1 - 117 + i - xxii."]}

Mimuloria davidcauseyi (Causey 1950a), revived combination Fig. 4–8, 16–17. Nannaria davidcauseyi Causey 1950a: 194, fig. 3–4. Hoffman 1999: 366. Marek et al. 2014: 36. Castanaria davidcauseyi: Causey 1950b: 1. Mimuloria davidcauseyi: Causey 1955: 30. Chamberlin and Hoffman 1958: 37. Type specimens. Male holotype (ANSP) and one male, one female, and one juvenile paratypes (FSCA) collected by David Causey, 25 August 1950, at Jasper, Newton Co., Arkansas. Diagnosis. Acropodite sublinear, angling or leaning mediad rather than curving or arching; inner projection of prefemoral process long and blade-like, extending for 1/3 or more of acropodital length. Color in life. Males from Newton Co., Arkansas, were more fully tanned than the M. castanea females, but displayed similar colors and patterns. Paranotal margins and peritremata were clear, translucent, and colorless with irregular, light pinkish-orange spots internally (Fig. 4) and broad, concolorous bands along the anterior margins of the collums. The background color was a moderately dark olive-green that was the same darkness on both the anterior and caudal halves of the metaterga. The head color was an even darker olive green on the epicranium that became slightly but progressively lighter anteriorly. Antennomeres 2–6 were moderately dark and articles 1 and 6 were white and slightly tinted, respectively (Fig. 5). Holotype. Fragmented into four pieces; length 21.8 mm, maximum width 4.0 mm (Causey 1950a), W/L ratio 18.3%. Somatic features agreeing closely with those of M. castanea with following exceptions. Width across genal apices 2.5 mm, segmental widths: collum 3.3 mm, 2 nd – 16 th segments 4.0 mm, 17 ~ 3.5 mm. Facial setae as follows: epicranial 0–0, interantennal 2–2, subantennal 1–1, frontal 1–1, genal 0–0, clypeal 1–1, labral 12–12. Antennae 2.5 mm long, reaching back to midlength of 3 rd tergite; relative lengths of antennomeres 6=3>4=5=2>1>7 (Fig. 5). Collum 1.6 mm long. Terga with faint wrinkling. Anterior corners of 2 nd – 4 th paranota rounded, caudolateral corners slightly extended, becoming more so and triangular on 12 th paranota. Caudal sternal spines strong and robust, anterior ones smaller and broader (Fig. 6). Relative lengths of postgonopodal podomeres 3>6>2>4=5=1. Coxae sparsely hirsute, prefemora with hairs distal to midlengths, becoming progressively denser on distal podomeres (Fig. 7), those on tarsi exceeding lengths of claws; prefemoral spines moderately long (Fig. 6). Tarsal claws broadened, flattened, and somewhat spatulate but not twisted on legs 1–14 (segment 2 through anterior legs on segment 10), apically subacuminate (Fig. 7). Gonopods in situ (Fig. 5, 8) overhanging caudal extremity of 5 th sternum. Gonopod structure (Fig. 8, 16–17) as follows: Prefemoral process long and blade-like, extending for nearly half of acropodital length. Latter sublinear, angling mediad; otherwise as described for M. castanea. Male paratype. The male paratype agrees closely with the holotype. Variation. We examined six males of this species and found no variation. Distribution. Known only from a two county area in northwestern Arkansas. Mimuloria davidcauseyi seems to have “carved out” a small enclave in northwestern Arkansas just inside the peripheries of the generic/tribal ranges and that of M. castanea. Published records. Arkansas: Newton Co. (Causey 1950b), Jasper and 4.8 km (3.0 mi) NW Jasper (Causey 1950a 1955; Chamberlin and Hoffman 1958; Hoffman 1999; Marek et al. 2014). Kentucky: Fayette Co., Lexington (Causey 1955)., Published as part of Hennen, Derek A. & Shelley, Rowland M., 2015, A contribution on the milliped tribe Nannariini (Polydesmida: Xystodesmidae): Revalidation of Mimuloria Chamberlin 1928; identities of Fontaria oblonga C. L. Koch 1847, and Nannaria minor Chamberlin 1918; elucidation of the tribal range; and commentaries on Nannaria Chamberlin 1918, and Oenomaea Hoffman 1964, pp. 1-21 in Insecta Mundi 2015 (418) on pages 10-12, DOI: 10.5281/zenodo.5182058, {"references":["Causey, N. B. 1950 a. On four new polydesmoid millipeds. Entomological News 61: 193 - 198.","Hoffman, R. L. 1999. Checklist of the millipeds of North and Middle America. Virginia Museum of Natural History Special Publication Number 8: 1 - 584.","Marek, P., T. Tanabe, and P. Sierwald. 2014. A species catalog of the millipede family Xystodesmidae (Diplopoda: Polydesmida). Virginia Museum of Natural History Special Publication Number 17: 1 - 117 + i - xxii.","Causey, N. B. 1950 b. A new genus and species of diplopod (Family Xystodesmidae). Chicago Academy of Sciences Natural History Miscellanea 71: 1 - 3.","Causey, N. B. 1955. New records and descriptions of polydesmoid millipeds (order Polydesmida) from the eastern United States. Proceedings of the Biological Society of Washington 68: 21 - 30.","Chamberlin, R. V., and R. L. Hoffman. 1958. Checklist of the millipeds of North America. Bulletin of the United States National Museum 212: 1 - 236."]}

Mimuloria dilatata sigmoidea, new subspecies Fig. 20–21. Type specimen. Male holotype (NCSM) collected by R. M. Shelley and W. B. Jones, 14 October 1978, ca. 7.2 km (11.5 mi) S Decatur, along county road 4274, 0.8 km (0.5 mi) from junction of Tennessee highway 58, Meigs Co., Tennessee. Diagnosis. Acropodital lobe broad, well defined, and distinct; prefemoral process with basal shoulder/ flange on dorsal surface, configuration broadly sigmoidal, curving strongly inward into acropodital arc and then outward, terminating near level of latter’s tip (Fig. 20–21). Etymology. The subspecific name denotes the overall sigmoidal curvature of the prefemoral process, which also passes through numerous vertical planes. Distribution. Known only from the type locality, in the Tennessee River Valley on the east side of the river itself., Published as part of Hennen, Derek A. & Shelley, Rowland M., 2015, A contribution on the milliped tribe Nannariini (Polydesmida: Xystodesmidae): Revalidation of Mimuloria Chamberlin 1928; identities of Fontaria oblonga C. L. Koch 1847, and Nannaria minor Chamberlin 1918; elucidation of the tribal range; and commentaries on Nannaria Chamberlin 1918, and Oenomaea Hoffman 1964, pp. 1-21 in Insecta Mundi 2015 (418) on page 15, DOI: 10.5281/zenodo.5182058

Genus Mimuloria Chamberlin 1928, revalidated Mimuloria Chamberlin 1928: 155. Attems 1940: 490, 530. Causey 1952: 7–8; 1955: 30. Chamberlin and Hoffman 1958: 37. Jeekel 1971: 273. Castanaria Causey 1950b: 1. Jeekel 1971: 254. Type-species. Of Mimuloria, M. missouriensis Chamberlin 1928, by original designation; of Castanaria, C. depalmai Causey 1950b, by original designation. Causey (1952) placed Castanaria in synonymy under Mimuloria, which was accepted by Chamberlin and Hoffman (1958). Diagnosis. Gonopod without solenomere; acropodite arching mediad in situ, crossing opposite member distally or near midlength and terminating over opposing coxa; tip ornamented with either variably rounded lobe or short spiniform projection on anterior surface, or narrow lobes on dorsal and ventral margins and caudally directed, subtriangular lamina extending from inner surface and carrying prostatic groove. Components. We recognize two assemblages that are based primarily on the distal acropodital configuration and named for the dominant species. Distribution. Northeastern Indiana and east central Tennessee to southwestern Missouri and northwestern Arkansas. Remarks. Representatives of the dilatata group possess anterior lobes/ornamentations at the same terminal position as the dorsal/ventral ones of the castanea group. We combine these conditions and define Mimuloria as having apically lobed or ornamented acropodites. The pregonopodal tarsal claws in Mimuloria are the second type that Hoffman (1964) characterized in Note 1 at the conclusion of his work. They are broad and somewhat flattened/ spatulate but not twisted; they also narrow apically and are subacuminate rather than truncate (Fig. 7). Gonapophyses are broad basally and relatively long for such relatively small-bodied millipeds (Fig. 5). The Castanea Group. Acropodites of the castanea group angle or arch variably mediad in situ, thereby crossing midlines of the bodies and the opposite members, terminating at the level of the latter’s coxa. The apical acropodital lobes are short, narrow, and inconspicuous in ventral view; one must reorient the structure or focus down on the terminus to see them. Instead of terminating at this position, a short, subtriangular, caudally-directed lamina arises from the inner acropodital surface that carries the prostatic groove to its opening (Fig. 9, 11–13, 15–17). Prefemoral processes vary from short to nubbin-like, and variable projections may or may not arise from its corners. That from the inner corner (relative to the acropodital arc) is longer and more prominent, being blade-like and extending for 1/3 of the acropodital length in M. davidcauseyi and short, narrow, and acuminate in M. castanea. Components. M. castanea (McNeill 1887), M. missouriensis Chamberlin 1928, and M. davidcauseyi (Causey 1950a). Distribution. The castanea group is the exclusive inhabitant of the western 1/3 of the tribal distribution, extending from northeastern Indiana and western Tennessee to northwestern Arkansas and southwestern Missouri., Published as part of Hennen, Derek A. & Shelley, Rowland M., 2015, A contribution on the milliped tribe Nannariini (Polydesmida: Xystodesmidae): Revalidation of Mimuloria Chamberlin 1928; identities of Fontaria oblonga C. L. Koch 1847, and Nannaria minor Chamberlin 1918; elucidation of the tribal range; and commentaries on Nannaria Chamberlin 1918, and Oenomaea Hoffman 1964, pp. 1-21 in Insecta Mundi 2015 (418) on pages 5-6, DOI: 10.5281/zenodo.5182058, {"references":["Chamberlin, R. V. 1928. Some chilopods and diplopods from Missouri. Entomological News 39: 153 - 155.","Causey, N. B. 1952. Some records and descriptions of polydesmoid millipeds from the United States. Chicago Academy of Sciences Natural History Miscellanea 106: 1 - 11.","Causey, N. B. 1955. New records and descriptions of polydesmoid millipeds (order Polydesmida) from the eastern United States. Proceedings of the Biological Society of Washington 68: 21 - 30.","Chamberlin, R. V., and R. L. Hoffman. 1958. Checklist of the millipeds of North America. Bulletin of the United States National Museum 212: 1 - 236.","Jeekel, C. A. W. 1971. Nomenclator generum et familiarum Diplopodorum: A List of the genus and family-group names in the Class Diplopoda from the 10 th edition of Linnaeus, 1758, to the end of 1957. Monografieen van de Nederlandse Entomologische Vereniging 5: 1 - 412 + i - xii.","Causey, N. B. 1950 b. A new genus and species of diplopod (Family Xystodesmidae). Chicago Academy of Sciences Natural History Miscellanea 71: 1 - 3.","Hoffman, R. L. 1964. The status of Fontaria pulchella Bollman, with the proposal of a new genus and tribe in the diplopod family Xystodesmidae. Proceedings of the Biological Society of Washington 77: 25 - 34.","McNeill, J. 1887. Descriptions of twelve new species of Myriapoda, chiefly from Indiana. Proceedings of the United States National Museum 10: 328 - 334.","Causey, N. B. 1950 a. On four new polydesmoid millipeds. Entomological News 61: 193 - 198."]}