Nannaria castanea (McNeill, 1887) Figs 32, 33 Vernacular name: "The Bloomington Twisted-Claw Millipede"

Polydesmus castaneus McNeill, 1887: 329, fig. 8

Fontaria castanea: Bollman 1893: 123.

Mimuloria castanea: Chamberlin 1928: 155. Causey 1952: 8, fig. 6c; 1955: 30. Chamberlin and Hoffman 1958: 37. Hennen and Shelley 2015: 6, figs 3, 9-14.

Nannaria castanea: Chamberlin 1949: 4. Hoffman 1999: 365-366. Marek et al. 2014: 36. Means et al. 2021:17, S68.

Castanaria castanea: Causey 1950a: 1.

Castanaria depalmai: Causey 1950a: 1-3, fig. 1.

Mimuloria depalmai: Causey 1952: 9. Chamberlin and Hoffman 1958: 37-38.

Nannaria depalmai: Hoffman 1999: 366. Marek et al. 2014: 36.

Material examined.

Lectotype (here designated): United States - Indiana • ♂; Monroe County, Bloomington; NMNH Type #38.

Paralectotype: United States - Indiana • 1 ♀; same collection data as holotype; NMNH Type #38 .

Other material.

United States - Arkansas • 11 ♀♀; Carroll County, Eureka Springs, Lake Leatherwood, Hyde Trail south of lake; 36.4307°N, - 93.7576°W; elev. 317 m; 17 May 2017; hand collected; J. Means, D. Hennen, V. Wong leg.; VTEC MPE02765 - 70, MPE02797 - 99, MPE03766, MPE03767 • 1 ♂; Fulton County, Hwy. 62; 24 Apr. 1952; N. Causey leg.; VMNH NAN0196; SCAU - Mississippi • 1 ♂; Choctaw County, 4.5 mi. S. of Ackerman, upland woods; 33.2809°N, - 89.1690°W; 2 Dec. 1961; L. Hubricht leg.; VMNH NAN0238 • 3 ♀♀; Tishomingo County, Burnsville, Divide Wildelife Management Area, forest beside pulloff on road; 34.8051°N, - 88.3063°W; elev. 159 m; 14 May 2017; hand collected; J. Means, D. Hennen, V. Wong leg.; VTEC MPE02789 -91 • 1 ♂; Tishomingo County, wooded hillside 1.6 miles W of Burnsville; 34.8388°N, - 88.3298°W; 27 Feb. 1961; L. Hubricht leg.; VMNH NAN0241; SCAU - Missouri • 1 ♂; Crawford County, 5 miles W of Berryman; 37.9188°N, - 91.1871°W; 2 Apr. 1955; R. Crabill leg.; VMNH NAN0242. For detailed collection data see Suppl. material 7 .

Diagnosis.

Adult males of Nannaria castanea are distinct from other Nannaria based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially, not curving ventromedially before apex as in N. hokie . Distal zone with medial flange at 135° to solenomere, and lateral flange at 90° angle to solenomere (Fig. 32B, red triangle), not encircling tip as in N. hokie . Acropodite tip curving caudally. Acropodite robust, simple; not thin, with small medial flange near apex as in N. hokie . Telopodite basal zone height ca. 1/2 length of acropodite, not ca. 1/3 length as in N. hokie . Prefemoral process small, thin, curving medially, arising from top of projected, stout prefemoral spine (Fig. 32A, red arrow), not arising from thin, projected prefemoral spine as in N. hokie . Color. Tergites with orange paranotal spots (Fig. 33) and occasionally faint orange stripes (Fig. 33). Dark to light brown background. Dorsum of collum smooth with orange margin.

Measurements.

Lectotype ♂ (NMNH, Type #38): CW = 3.5, IW = 2.2, ISW = 0.8. BL, B11W and B11H were unmeasurable due to the poor quality of the specimen.

Variation.

Castanaria depalmai was synonymized with N. castanea by Hennen and Shelley (2015) without justification and the relationship between C. depalmai and N. castanea has not been tested using molecular evidence. Morphological differences between the two are extremely slight, however, so it is likely that the synonymy of C. depalmai and N. castanea will be supported by molecular phylogenetics. Interestingly, there are six specimens of N. castanea collected from two localities in Mississippi by Leslie Hubricht in 1961 that Hennen and Shelley (2015) were apparently unaware of and did not mention in their treatment of Mimuloria . The northern Mississippi specimen is remarkably similar in form to N. missouriensis, however specimens collected by the authors are genetically more similar to N. castanea from Arkansas than N. missouriensis, and we therefore identify the two Mississippi populations as N. castanea, awaiting more detailed investigations.

Distribution.

Known from central Indiana west to Missouri and south into Arkansas and Mississippi (Indiana: Monroe County; Missouri: Jefferson, Crawford, Dent, Wayne, Wright counties; Arkansas: Carroll, Searcy, Stone, Fulton counties; Mississippi: Tishomingo, Choctaw counties; Suppl. material 7; Fig. 129). Distribution area: 113,102 km2; status: WRE.

Ecology.

Individuals of N. castanea were collected from both mesic broadleaved and xeric semi-evergreen forests, often beneath 1-2 cm of soil.

Etymology.

McNeill (1887) gave no etymology for his choice of castaneus but he mentioned the dark chestnut coloration of the species, suggesting that his choice of name may have been derived from the Latin castaneus for 'of the color of chestnuts.'

Type locality.

United States, Indiana, Monroe County, Bloomington.

Notes.

In the original description, McNeill (1887: 329) stated that he examined three specimens but did not designate a holotype, and no lectotype has been designated by subsequent authors. Therefore, we consider two type specimens we examined as syntypes and here we designate a lectotype male. There is no information on who collected the type material and when. The label of " NMNH Type #38" was presumeably added by an unknown individual after the specimens were deposited at the NMNH.