Brachoria flammipes, Marek, 2010

Marek, Paul E., 2010, A revision of the Appalachian millipede genus Brachoria Chamberlin, 1939 (Polydesmida: Xystodesmidae: Apheloriini), Zoological Journal of the Linnean Society 159 (4), pp. 817-889 : 846-849

BRACHORIA FLAMMIPES SP. NOV.

‘THE FIRE- FOOTED MIMIC MILLIPEDE’

( FIG. 16A–D, FIG . 40F–G)

Diagnosis: Adult males of B. flammipes are distinct from other Brachoria species based on the combination of: Colour. Striped ( Fig. 40F), yellow metatergal stripes, collum stripe, legs. Striped ( Fig. 40G), red metatergal stripes, collum stripe, legs. Exoskeleton. Caudolateral corners, paranota 1–8 rounded, lateral margins 1–4 angled cephalically. Gonopods. Gonopodal acropodite narrow, ~seven-tenths width of tibia on leg pair 9, distinctly narrowed halfway until apex. Prefemur without prefemoral process, or with tiny thorn-like process. Acropodite bent anteromedially at 50°, at cingulum, one-quarter of its length from prefemur ( Fig. 16C ). Distinctly narrowed distal portion, meathook-like, curved dorsally ( Fig. 16B ). Acropodal apex projected posterolaterally, sinuate, acuminate tip slightly curved dorsally ( Fig. 16B , arrow). Note: B. flammipes and B. dentata appear similar as a result of close evolutionary relationship. However, the two can be distinguished by the form of the gonopods and phylogenetic location. Specifically, B. flammipes has an unadorned meathook-like acropodite distal beyond the cingulum, whereas B. dentata has a single (or double in some Kentucky populations), stout sawtooth projection ventromedially at the base of the narrowed distal portion ( Fig. 11B , arrow). In addition, the acropodite of B. flammipes is bent anteromedially at 50°, at the cingulum, one-quarter of its length from the prefemur (Fig. 39V); whereas in B. dentata the acropodite is bent anteromedially at a smaller angle (45°), one-eighth of its length from the prefemur ( Fig. 11C ). Although colour should be met with suspicion as a diagnostic character in Brachoria , generally B. dentata are either two-spotted or three-spotted and B. flammipes striped.

Description: Holotype (♂) FMNH-INS43068 – measurements: BL = 45.87. CW = 7.77. IW = 5.30. ISW = 1.50. B10W = 10.40. B10H = 6.55. AL = 2.20. Head: antennae extending backwards to middle of third tergite, relative antennomere lengths 2> 3> 5> 4> 6> 1> 7. Tergites: collum with curved cephalic edge, tapering laterally. Collum without carina present on anterior margin. Paranota 1–8 with caudolateral corners rounded, lateral margins 1–4 angled cephalically. Paranotal dorsal surface tightly wrinkled, appearing leathery. Ozopores orientated dorsolaterally. Paranota with anterodorsal corner scooped out. Gonapophyses goblet-shaped apically. Pleural tubercle absent. Sterna without posteriorprojecting spines, with flat caudal margin. Sterna 2–8 sparsely setose (Ể ten setae), sterna 9–18 without setae, ventral surface smooth. Gonopods: in situ configuration– acropodite projected ventrally, bent anteriorly at 50°, at cingulum, one-quarter of its length from prefemur ( Fig. 16C ). Distinctly narrowed distal portion, meathook-like, curved dorsally ( Fig. 16B ). Acropodal apex projected posterolaterally, sinuate, acuminate tip slightly curved dorsally. Terminal arc of acropodite orientated anterolaterally. Left, right acropodites crossed midlength, appearing intertwined. Telopodites – prefemur with tiny thornlike process, ~1/32 length of acropodite, equal to its width at its base. Gonopodal acropodite narrow, ~seven-tenths width of tibia on leg pair 9, distinctly narrowed halfway until apex. Acropodite with ventrobasal surface facing medially, flat without flange or swelling. Acropodite base without spines on dorsal surface. Acropodal basal and apical ventral surfaces not coplanar, apical surface rounded, facing laterally; anterior twist well defined, acropodite immediately distal beyond cingulum curved cephalically, apical arc recurved posteriorly. Acropodal ventral surface not swollen, smooth. Acropodite with midlength transverse groove, cingulum. Acropodite distinctly narrowed halfway until apex, about 2.8 ¥ narrower in distal half than basal half at its widest section. Acropodite immediately distal to cingulum equal width to acropodite immediately basal to cingulum. Postcingular region of acropodite without ventromedial or lateral teeth. Acropodal apex projected posterolaterally. Acropodite with postcingular region ~one-quarter of its total length. Anterolateral edge of acropodal postcingular region with small flange ( Fig. 16A , arrow). Acropodite with an undivided apex. Acropodal apex narrow, ~one-third width of postcingular region, sinuate acuminate tip slightly curved dorsally ( Fig. 16B ). Paratype (♀) FMNH-INS43070 – measurements: BL = 43.55. CW = 7.70. IW = 5.87. ISW = 1.65. B10W = 10.25. B10H = 7.20. Cyphopods: cyphopods with receptacle (at its greatest breadth) narrower than prefemur length. Receptacle rectangular, one side slightly wider, widest edge facing anterodorsally. Cyphopodal valves nearly symmetric. Cyphopods with valvular suture wavy, facing laterally.

Variation: Brachoria flammipes is known only from the type locality and seven specimens (4♂ and 3♀ from type series collected in 2005). There is some variation in coloration amongst these specimens with three morphs, all striped (red or yellow): (1) striped ( Fig. 40F), yellow (10YR 7/12) paranota and anterior collum stripe, orange (7.5YR 6/12) metatergal stripes, legs (1.25YR 4/12); (2) striped, yellow (1.25Y 8/14) paranota, anterior collum stripe, yellow (1.25Y 7/14) metatergal stripes, legs; (3) striped ( Fig. 40G), red metatergal stripes, anterior collum spot, legs. There is some variation in length amongst these specimens, specifically I observed two length classes in males: between 45.87–50.05 and 40.71–41.67. Measurements: ♂ (N = 4) BL = 40.71–50.05 (44.58/4.28). CW = 7.29–8.20 (7.70/0.39). IW = 4.98–5.80 (5.27/ 0.38). ISW = 1.40–1.75 (1.52/0.16). B10W = 9.30–10.50 (9.90/0.64). B10H = 5.55–7.10 (6.23/0.73). ♀ (N = 3) BL = 43.55–55.09 (48.60/5.90). CW = 7.70–7.90 (7.82/ 0.10). IW = 5.80–5.89 (5.85/0.05). ISW = 1.65–1.73 (1.69/0.04). B10W = 10.25–11.19 (10.65/0.49). B10H = 7.20–7.70 (7.38/0.28).

Ecology: Brachoria flammipes specimens were collected during the evening (19:00) in a moist forest comprising maple, hemlock, dogwood, sycamore, birch, and tulip poplar trees. Individuals were found on a north-facing slope beneath layers of decomposing leaf material on the side of a trail above (south) Cawood Branch. Other xystodesmids encountered co-occurring with B. flammipes were B. splendida , A. v. corrugata and an undescribed species of Nannaria . Note: individuals of B. flammipes may be confused in the field with B. splendida and A. v. corrugata because of similarity in colour patterns, but can be distinguished from B. splendida by the diagnostic characters (specifically, B. flammipes has a narrow acropodite distally and B. splendida a bulky S-shaped acropodite distally), and from A. v. corrugata by the presence of the following characters (viewed with a 20¥ loupe): acropodite with midlength transverse groove, cingulum; prefemur without prefemoral process, or with tiny thorn-like process; acropodite with distinctly narrowed distal portion, meathooklike, curved dorsally; acropodite not circular like in A. v. corrugata . Nannaria is easy to distinguish; it has an olive drab colour and a body length ~threequarters that of B. flammipes .

Distribution: Known only from the type locality. The closest known Brachoria species , B. splendida , co-occurs with B. flammipes at Cawood Recreation Site. Brachoria gracilipes occurs about 17 km east in the town of Pine Mountain, Harlan County, Kentucky.

Etymology: The specific epithet was derived from my first impression of some specimens’ bicoloured legs while collecting them at Cawood. Some individuals’ legs are brilliant orange at the base and bright yellow at the tip, imparting a flame-like appearance. (The specific name is a noun derived from the Latin words: flamma = flame, and pes = foot.)

Material examined: Type specimens. ♂ holotype ( FMNH-INS43068 *: EU127870 ) , 1 ♂, 1 ♀ paratypes ( FMNH-INS43069 , 70 ) , 1 ♂, 1 ♀ paratypes ( USNM,

SPC001086, 1021) from Kentucky, Leslie County, Daniel Boone National Forest, Cawood Recreation Site , about 5.4 rd km north of junction KY-221 and US-421 (36.93645°N, - 83.37294°W, 404 m), 27.vii.2006, 19:00 (colls: P. and B. Marek). Nontype specimens. Kentucky: 1♂ (A8705), 1♀ (A8704), Leslie County , 26.4 km south of Hyden , Cawood Recreation Area , Daniel Boone National Forest (36.9356°N, - 83.3724°W), 11.vi.2001 (coll: R. Shelley). 1♂ (SPC001086), same as type locality, 9.v.2007, 11:00 (colls: P. Marek, D. and M. Beamer, C. Hall) .

BRACHORIA GLENDALEA ( CHAMBERLIN, 1918) ‘THE GLENDALE HILLS MIMIC MILLIPEDE’

( FIG. 17A–D, FIG . 40H)

Fontaria glendalea Chamberlin, 1918: 123 . Brachoria glendalea – Chamberlin & Hoffman, 1958: 212; Keeton, 1959: 28.

Diagnosis: Adult males of B. glendalea are distinct from other Brachoria species based on the combination of: Colour. Two-spotted ( Fig. 40H), yellow paranotal spots, anterior collum stripe, legs. Tergites frequently brown, not black ( Fig. 40H). Exoskeleton. Caudolateral corners, paranota 1–4 rounded, lateral margins angled cephalically. Gonopods. Gonopodal acropodite very narrow, one-third width of tibia on leg pair 9, swollen distally beyond cingulum twofold. Acropodite projecting straight from prefemur, not curved medially before cingulum ( Fig. 17A ). Acropodite beyond cingulum circularly coiled anteriorly ( Fig. 17C , arrow). Acropodite apex tapered distally, with flanged acuminate tip ( Fig. 17B , arrow). Note: most species of the genus have brilliantly variable coloration throughout their distribution. However, B. glendalea has the most uniform colour pattern throughout its range of any species in the genus ( Fig. 40H).

Variation: Brachoria glendalea is known from Davidson County south to Maury, Bedford, and Coffee counties. There is negligible variation in coloration and measurements amongst these specimens. Measurements: ♂ (N = 5) BL = 38.30–44.73 (40.51/2.51). CW = 7.55–8.45 (8.03/0.37). IW = 4.79–5.62 (5.19/ 0.34). ISW = 1.45–1.65 (1.53/0.08). B10W = 9.94–11.28 (10.66/0.66). B10H = 5.65–6.70 (6.10/0.43). ♀ (N = 4) BL = 40.39–44.30 (42.72/1.67). CW = 7.35–8.40 (7.90/ 0.44). IW = 5.48–5.89 (5.65/0.17). ISW = 1.75–1.90 (1.81/0.06). B10W = 10.30–11.60 (10.80/0.56). B10H = 6.75–7.60 (7.08/0.39).

Description: Male described by Chamberlin (1918); redescribed by Keeton (1959).

Ecology: Brachoria glendalea specimens were collected during the day (9:30 and 13:00) in two localities: at the type locality, in a disturbed area in Nashville, and at Radnor Lake State Nature Preserve. I found two millipedes within the city limits of Nashville, one of which, a large female, I did not collect. The Nashville locality is a shaded and disturbed gully with Hedera helix L. blanketing a large part of the soil. No other xystodesmids were encountered sympatrically here. Brachoria glendalea from Radnor Lake were found in a maple, birch, and tulip poplar forest in a moist gully. One other xystodesmid encountered co-occurring at the Radnor Lake locality is Falloria mimetica ( Chamberlin, 1918) . Note: individuals of B. glendalea may be confused in the field with F. mimetica , but can be distinguished by the presence of the following characters (viewed with a 20¥ loupe): two-spotted, yellow paranotal spots, anterior collum stripe, legs; F. mimetica has red paranota and purple metatergal stripes; acropodite beyond cingulum circularly coiled anteriorly, F. mimetica has a bulky acropodite, not circularly coiled.

Distribution: Known from the Interior Plateau of Tennessee in Davidson County south through Marshall County and into Maury, Bedford, and Coffee counties; and west to Hickman County. The closest known Brachoria species , B. initialis occurs south and east of B. glendalea . Brachoria hubrichti occurs east in the Cumberland Plateau and Sequatchie Valley.

Material examined: Type specimens. ♂ holotype ( MCZ) , ♂, ♀ paratypes ( USNM) from Davidson County, Tennessee, Glendale Hills, 21.iv.1917 (coll: H. Cummins) (vidi paratypus). Nontype specimens. Tennessee : 1♂ ( PMLN0165 ), Bedford County, 14.4 km south of Shelbyville (35.3907°N, - 86.4945°W), 8.x.1960 (coll: L. Hubricht) . 1♂, 1♀ (A2670), Coffee County, Old Stone Fort (35.4859°N, - 86.1015°W), 11.v.1979 (coll: R. Shelley) . 1♂ (SPC000481*: EU127850 ), Davidson County, Nashville, gully draining into West Fork , near junction of Scenic Dr and Glendale Ln. (36.09468°N, - 86.79347°W, 165 m), 22.v.2005, 9:30 (coll: P. Marek) . 1♂ (SPC000484), 1♀ (SPC000489), Davidson County, Radnor Lake State Nature Preserve, South Cove Trail , in gully near loop trailhead (36.05945°N, - 86.80724°W, 263 m), 22.v.2005, 13:00 (coll: P. Marek) . 1♂ ( PMLN0060 ), Hickman County, Duck River bluff, north of Centerville (35.7891°N, - 87.4673°W), 24.ix.1955 (coll: L. Hubricht) . 3♂, 2♀ (A2655), Marshall County, Henry Horton State Park campground (35.5855°N, - 86.6923°W), 5.v.1979 (coll: R. Shelley) .

BRACHORIA GRACILIPES ( CHAMBERLIN, 1947)

Figures

Figure 11. Brachoria dentata Keeton, 1959. A–D, holotype specimen gonopods: A, left male gonopod, anterior view; B, left male acropodal apex, magnified; C, left male gonopod, medial; D, same, posterior (acropodite length = 2.00 mm).

Figure 16. Brachoria flammipes sp. nov. A–D, holotype specimen gonopods: A, left male gonopod, anterior view; B, left male acropodal apex, magnified; C, left male gonopod, medial; D, same, posterior (acropodite length = 2.20 mm).

Figure 17. Brachoria glendalea (Chamberlin, 1918). A–D, paratype specimen gonopods: A, left male gonopod, anterior view; B, left male acropodal apex, magnified; C, left male gonopod, medial; D, same, posterior (acropodite length = 1.80 mm).