Nannaria fowleri Chamberlin, 1947 Figs 15, 16 Vernacular name: “Fowler’s Twisted-Claw Millipede"

Nannaria fowleri Chamberlin, 1947: 29, fig. 14. Chamberlin and Hoffman 1958: 40. Shelley 1988: 1656. Hoffman 1999: 366. Shelley 2002: 1872. Marek et al. 2014: 37. Means et al. 2021: S70.

Nannaria cayugae Chamberlin, 1949: 4, fig. 3. Chamberlin and Hoffman 1958: 40. Shelley 1988: 1656.

Material examined.

United States - Maryland • 1 ♂; Allegany County, E of Flinstone, 1600 ft. summit of Town Hill Mtn.; 39.6903°N, - 78.4067°W; 19 May 1958; L. Hubricht leg.; VMNH NAN0352 • 1 ♂; Allegany County, Cumberland, Knobby Mountain; 39.6527°N, - 78.7626°W; 29 March; K. Holbrook; NCSM NAN0463; SCAU - New York • 1 ♂; Cattaraugus County, Allegany St. Pk., Anderson Trail; 42.0023°N, - 78.8357°W; 10 May 1958; Muchmore leg.; VMNH NAN0347 • 1 ♂; McKean County, 8 Oct. 1959; VMNH NAN0348 • 1 ♂; Monroe County, Mendon Ponds Park; 42.9988°N, - 77.5641°W; 17 Apr. 1958; Muchmore leg.; VMNH NAN0344; SCAU - Pennsylvania • 1 ♂; Centre County, Toftrees State Game Lands; 40.9680°N, - 77.9154°W; 1993; T. McCoy leg.; NCSM NAN0451 • 3 ♂♂; Clinton County, Rauchtown, Ravensburg State Park, bank of Rauchtown Run, north of restrooms beside playfield; 41.1034°N, - 77.2435°W; elev. 368 m; 18 Sep. 2017; hand collected; J. Means & D. Hennen leg.; VTEC MPE03071 , 3094, 3906 • 10 ♀♀; same collection data as preceding; VTEC MPE03072, MPE03095 -102, 3907 • 1 ♂; Franklin County, Caledonia State Park, Thaddeus Stevens Trail; 39.9079°N, - 77.4777°W; elev. 287 m; 15 Sep. 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE03017 • 1 ♂; Franklin County, low woods, near creek 1.8 miles S of Caledonia; 39.7359°N, - 77.4967°W; 8 June 1956; L. Hubricht leg.; VMNH NAN0345 • 1 ♂; Franklin County, road to Cowants Gap State Park; 39.9957°N, - 77.9154°W; 11 Apr.1959; R. Mighton leg.; VMNH NAN0349 • 2 ♂♂; Potter County, Patterson Park, Susquehanna State Forest, 10 mi SSE of Coudersport, roadside logs; 41.6406°N, - 77.9471°W; 20 Aug. 1971; W. Shear leg.; VMNH NAN0050 • 1 ♂; Potter County, Austin Dam ruins; 41.6944°N, - 78.1197°W; 12 June; W. Shear leg.; VMNH NAN0342 • 1 ♂; Potter County, Denton Hill State Park, 14 mi. E of Coudersport, beech-birch-maple litter; 41.7685°N, - 77.8334°W; 22 Aug. 1971; W. Shear leg.; VMNH NAN0037 • 1 ♂; Potter County, Rock Ridge Road, 2 mi from Lyman Run State Park, wet birch logs near spring; 41.6609°N, - 77.8864°W; 23 Aug. 1971; W. Shear leg.; VMNH NAN0033; SCAU - Virginia • 5 ♂♂ ♀♀; Bath County, GWNF Long Spring Run above Little Back Creek; 38.2213°N, - 79.8387°W; 24 July 1992; J. Pagels, D. Kobuszewski leg.; VMNH NAN0651 • 1 ♂; same collection data as preceding; 11 Sep. 1992; D. Kobuszewski leg.; VMNH NAN0654 • 2 ♂♂; same collection data as preceding; 17 July 1992; J. Pagels leg.; VMNH NAN0659 • 1 ♂; Bath County, “F” GWNF H’Town; 37.9993°N, - 79.8315°W; 1 July 1992; J. Pagels leg.; VMNH NAN0653 • 4 ♂♂; Highland County, Locust Spring Rec. Area, 8 mi. NW of Hightown; 38.5152°N, - 79.6899°W; elev. 1158 m; 23 May 1973; R. Hoffman leg.; VMNH NAN0343 • 1 ♂; same collection data as preceding; 28 Apr. 1972; R. Hoffman, Knight leg.; VMNH NAN0346 • 1 ♂; Highland County, GWNF west bank, Laurel Fork (between Bear Hollow & Newman Rd.); 38.2862°N, - 79.7468°W; 17 May - 15 June 1993; VMNH NAN0658; SCAU - West Virginia • 2 ♀♀; Pendleton County, old road headed up to Spruce Knob; 38.6947°N, - 79.5146°W; elev. 1010 m; 24 May 2015; hand collected; M. Kasson leg.; VTEC MPE00433 , 446 • 1 ♂; Pendleton County, Spruce Knob; 38.7308°N, - 79.4913°W; elev. 1112 m; 2016; C. Stauder leg.; VTEC MPE01816 . For detailed collection data see Suppl. material 7 .

Diagnosis.

Adult males of N. fowleri are distinct from other Nannaria and the sympatric N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gradually curving medially before apex, not straight as in N. shenandoa . Distal zone short, bent medially forming 130° angle with acropodite (Fig. 15). Telopodite with basal zone> 1/4 length of prefemoral process. Tip and distal zone simple, rectangular, blunt, <1/8 length of acropodite, not large, curving with flange as in N. shenandoa . Prefemoral process arising from base of prefemoral spine, paralleling medial curve of acropodite (Fig. 15A), not directed laterally and crossing acropodite when viewed medially as in N. shenandoa or arising from top of prefemoral spine as in N. tasskelsoae sp. nov. Prefemur with stout, tooth-like prefemoral spine (Fig. 15C, red triangle). Color. Tergites with orange paranotal spots (Fig. 16). Dark brown background. Dorsum of collum smooth with orange margin. Chamberlin (1947) described the holotype as having yellow paranota, but his description was from a preserved specimen which had likely lost its color due to its preservation in alcohol.

Measurements.

♂ (VTEC, MPE03017): BL = 25.1, CW = 3.7, IW = 2.3, ISW = 0.8, B11W = 4.6, B11H = 2.9; ♀ (VTEC, MPE03098): BL = 28.8, CW = 3.7, IW = 2.3, ISW = 0.8, B11W = 4.4, B11H = 3.1.

Variation.

A male taken from Cumberland, Maryland (NCSM NAN0463), has a slight acropodite medial swelling, which has not been observed in other individuals of N. fowleri .

Distribution.

Nannaria fowleri has a larger distribution than most Nannaria minor species group Nannaria, perhaps due to post-glacial northward expansion from the eastern Appalachians (Virginia, West Virginia) north into Maryland, Pennsylvania and New York (Virginia: Bath and Highland counties; West Virginia: Pendleton and Pocahontas counties; Maryland: Allegany County; Pennsylvania: Centre, Clinton, Franklin, McKean, and Potter counties; New York: Cattaraugus and Monroe counties; Suppl. material 7; Fig. 127). Distribution area: 43,054 km2; status: WRE.

Ecology.

Individuals of N. fowleri have been collected from mesic hardwood forests composed of maple, hemlock, and scattered pine, often buried 1-2 cm under dark, sandy soil.

Etymology.

Chamberlin (1947) gave no etymology for the name fowleri but it is reasonable to assume that the name refers to one of the collectors of the holotype, H. W. Fowler.

Type locality.

United States, Maryland, Garrett County, Jennings.

Notes.

In the original description, Chamberlin (1947: 29) examined and designated the holotype male (ANSP Type no. 9951) collected in July 1907 by W. Stone and H. W. Fowler.