Brachoria ligula, KEETON, 1959

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 : 868-872

BRACHORIA LIGULA KEETON, 1959

‘THE WELCH TONGUE MIMIC MILLIPEDE’

( FIG. 30A–D, FIG . 41D)

Brachoria ligula Keeton, 1959: 40 .

Diagnosis: Adult males of B. ligula are distinct from other Brachoria species based on the combination of: Colour. Striped ( Fig. 41D), lemon-yellow metatergal stripes, anterior collum spot, 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, uniform width. Acropodite bent medially at 60°, one-quarter its length from prefemur. Acropodite distal to 60° angle curved cephalically 180° in C-shape form ( Fig. 30C ). Arc immediately distal beyond cingulum orientated anterolaterally. Acropodal apex, pointed posteriorly, long, rectangular with rounded distal edges ( Fig. 30D ). Posterior apical corner with thin, tongue-shaped carina ( Fig. 30C arrow, hence the name ligula ). Note: B. ligula can be distinguished from B. laminata by the presence of the tongue-shaped carina on the posterior apical corner ( Fig. 30C , arrow); whereas B. laminata has thin, finshaped carina appressed to the laminate apex ( Fig. 29D , bottom arrow).

Variation: Brachoria ligula is known only from near the type locality and ten specimens (2♂ and 1♀, the type series, collected in 1950, and 3♂ and 4♀ collected in 2004–5). There is some variation in coloration amongst individuals; specifically, a single specimen had a reddish brown tinge on the anterolateral paranotal corners, very similar to A. v. corrugata in the area. There is negligible variation in measurements amongst these specimens. Measurements: ♂ (N = 5) BL = 41.55–43.84 (42.56/0.83). CW = 7.55–8.00 (7.69/0.18). IW = 4.90–5.30 (5.09/ 0.17). ISW = 1.40–1.60 (1.51/0.08). B10W = 10.00– 10.60 (10.20/0.25). B10H = 5.65–6.20 (5.94/0.23). ♀ (N = 3) BL = 42.51–46.01 (44.45/1.78). CW = 7.50–7.70 (7.57/0.12). IW = 5.06–5.50 (5.32/0.23). ISW = 1.50– 1.70 (1.60/0.10). B10W = 10.04–10.60 (10.35/0.28). B10H = 6.70–7.00 (6.84/0.15).

Description: Male described by Keeton (1959).

Ecology: Brachoria ligula specimens were collected during the day (13:00) in a forested cove comprising birch, tulip poplar, and maple. Individuals were found beneath layers of decomposing leaf material in close proximity to a large yellowjacket nest. At the other site (near the roadside overlook), individuals were found in a forest comprising rhododendron, maple, hemlock, tulip poplar, and sassafras. Other xystodesmids encountered sympatrically at the first locality are: Sigmoria latior latior ( Brölemann, 1900) (two-spotted orange), A. v. corrugata (striped yellow) and Pleuroloma flavipes (three-spotted yellow). Note: individuals of B. ligula may be confused in the field with A. v. corrugata as a result of similarity in colour pattern, but can be distinguished by the presence of the following characters (viewed with a 20¥ loupe): acropodite with midlength transverse groove, cingulum; acropodite not circular, without an acuminate apex as in A. v. corrugata .

Distribution: Known only from the type locality (sites about 1.2 air km apart). The closest known Brachoria species , B. laminata , B. hoffmani , and B. campcreekensis , occur about 30 air km west, south, and east in Virginia and West Virginia, respectively.

Material examined: Type specimens. ♂ holotype, 1 ♂, ♀ paratypes ( USNM) from McDowell County, West Virginia, 2.08 km east of Premier, 28.vi.1950 (coll: L. Hubricht) (vidi). Nontype specimens. West Virginia: 1♂ (SPC000324*: DQ 490688 ), McDowell County , Welch , US-52, 0.75 rd km north-east of roadside overlook (37.43356°N, - 81.57610°W, 491 m), 16.viii.2004, 13:33 (coll: P. Marek). 1♀ (SPC000322) McDowell County , Welch, US-52, 125 rd m north-east from water tower, roadside pull-out (37.42377°N, - 81.58258°W, 447 m), 16.viii.2004, 12:00 (coll: P. Marek). 2♂ (SPC000756, 762), 3♀ (SPC000757, 759, 760), ibid., 22.vii.2005, 13:00 (colls: P. Marek and C. Spruill) .

BRACHORIA MENDOTA KEETON, 1959 ‘THE MENDOTA MIMIC MILLIPEDE’

( FIG. 31A–D, FIG . 41E–I)

Brachoria mendota Keeton, 1959: 42 . Appalachioria mendota – Marek & Bond, 2006. Brachoria mendota – Marek & Bond, 2007.

Diagnosis: Adult males of B. mendota are distinct from other Brachoria species based on the combination of: Colour. Two-spotted ( Fig. 41E), red paranotal spots, anterior collum spot, legs. Three-spotted ( Fig. 41I), yellow metatergal spots, paranotal spots, anterior and posterior collum spots (usually merged together in an hourglass shape), legs. (Note: these are the most frequent colour morphs. Brachoria mendota has five colour morphs with considerable variation amongst them. Colour can vary wildly and be completely misleading for identification purposes.) Exoskeleton. Paranota with lateral margins 1–4 angled cephalically. Gonopods. Gonopodal acropodite straplike, four-fifths width of tibia on leg pair 9, tapered to an acuminate apex distally beyond cingulum ( Fig. 31C ). Prefemur with long, thin icepick-like prefemoral process, one-third length of acropodite ( Fig. 31A, C, D ). Acropodite projecting straight from prefemur, not curved medially ( Fig. 31A ). Acropodite bent medially 45° and anteriorly 45°, half its length from prefemur. Acropodal apex bent 90°, with twisted falcate apex ( Fig. 31B ).

Variation: Brachoria mendota is widespread throughout the Cumberland Mountains and Valley and Ridges of Tennessee and Virginia. There is considerable variation in coloration and measurements amongst these specimens. There are five colour morphs with substantial variation amongst them: three-spotted yellow ( Fig. 41I), and red ( Fig. 41G); two-spotted yellow ( Fig. 41F) and red ( Fig. 41E); and four-spotted yellow ( Fig. 41H, i.e. with paranotal, metatergal, prozonal spots). There are regional differences in coloration and measurements amongst three areas: the Valley and Ridges, Cumberland Mountain massif, and Jefferson County, Tennessee. In the Valley and Ridges, individuals tend to be smaller and two-spotted red; whereas, in the Cumberland Mountain area, individuals tend to be larger and threespotted or four-spotted yellow. An apparently disjunct population was recently discovered in Jefferson County, Tennessee with two-spotted red or yellow coloration and wider metatergites than average for northern individuals in Virginia (average width of body ring 10:11.83 for Jefferson County specimens versus 9.99 for predominately Virginia specimens). Measurements: ♂ (N = 8) BL = 37.25–42.90 (40.05/ 2.25). CW = 6.68–8.91 (7.55/0.66). IW = 4.70– 6.00 (5.23/0.40). ISW = 1.30–1.79 (1.56/0.14). B10W = 9.05–12.30 (9.93/1.07). B10H = 5.49–7.50 (6.11/0.63). ♀ (N = 7) BL = 39.37–43.40 (41.22/1.49). CW = 7.24–9.25 (7.88/0.68). IW = 5.47–5.81 (5.61/ 0.15). ISW = 1.50–1.72 (1.68/0.08). B10W = 9.91–11.35 (10.58/0.53). B10H = 6.70–7.20 (6.97/0.16).

Description: Male described by Keeton (1959).

Ecology: Brachoria mendota is common in the Appalachian mixed mesophytic deciduous forests of Virginia and Tennessee. Near the type locality, individuals were collected from a maple and oak forest with a understory of Impatiens and Urtica on a steep slope from beneath decaying leaves. Many other xystodesmids co-occur with B. mendota . Note: individuals of B. mendota may easily be confused in the field with A. v. corrugata , Rudiloria kleinpeteri ( Hoffman, 1949) , B. cedra , and B. dentata because of similarity in colour patterns, but can be distinguished from the other Brachoria by the diagnostic characters and from A. v. corrugata and R. kleinpeteri by the presence of the following characters (viewed with a 20¥ loupe): acropodite with midlength transverse groove, cingulum; prefemur with long, thin icepicklike prefemoral process, one-third length of acropodite; acropodite projecting straight from prefemur, not curved medially ( Fig. 31C ); acropodite not circular like in A. v. corrugata . On Clinch Mountain, north-east of Gate City, B. mendota and R. kleinpeteri overlap in distribution and have similar coloration. However, A. v. corrugata does not appear similar in this area. However, in the highlands around Cumberland, Wallen, and Powell mountains, R. kleinpeteri is not present and A. v. corrugata appears extremely similar to B. mendota .

Distribution: Known from south-western Virginia and north-eastern Tennessee on Clinch Mountain, north-west through Wallen and Powell mountains, to Pennington Gap in Lee County Virginia. There is one, apparently disjunct population 80 air km west in Jefferson County, Tennessee. Brachoria mendota was previously only known from Clinch Mountain from Fugate Gap in Russell County to Gate City in Scott County in Virginia; however, I discovered a population in Jefferson County, Tennessee, 80 km southwest of the nearest known population at Flat Gap in Scott County, Virginia. Later, populations were found at Pennington Gap, Virginia south-east to Snake Hollow on Newman’s Ridge in Tennessee. Brachoria mendota co-occurs with B. cedra and B. dentata forming a component of a Müllerian mimicry ring with them.

Material examined: Type specimens. ♂ holotype ( USNM) from Russell County, Virginia, Fire Tower, Mendota , 30.vii.1941 (colls: Dr and Mrs Brooks) (vidi). Nontype specimens. Tennessee : 1♀ ( MMC0449 ), Hancock County, Newman’s Ridge, Snake Hollow Rd (36.53873°N, - 83.31737°W, 500 m), 4.x.2006, 15:00 (coll: P. Marek) . 1♂ (SPC000450), 1♀ (SPC000448), Jefferson County, Jefferson City, N. Ridge Dr, near Mossy Creek (36.53873°N, - 83.31737°W, 350 m), 19.v.2005, 15:00 (coll: P. Marek). Virginia : 1♀ ( MMC0286 ), Lee County, Poor Valley, Pennington Gap , VA-621, about 1.6 km east US-421, on north-facing slope (36.77681°N, - 83.01574°W, 460 m), 28.ix.2006, 9:30 (colls: P. and B. Marek) . 1♂ ( PMLN0033 ), Lee County, about 3.2 km north of Stickleyville , VA-738 (36.7355°N, - 82.9021°W), 23.v.1994 (coll: C. Hobson) . 1♀ ( MMC0270 ), Lee County, Lovelady Gap, CR 619 (36.763661°N, - 82.88194°W, 815 m), 27.ix.2006, 12:15 (colls: P. and B. Marek) . 2♂ ( PMLN0031 ), Russell County, junction VA-606 and Copper Creek , VA-678 (36.8240°N, - 82.2381°W), 23.iv.1994 (coll: M. Donohue) . 1♂ (SPC000525*: EU127853 ), Russell County, Clinch Mountain, Fugate’s Gap , about 6.3 rd km up the mountain from VA-802 and Mendota , CR-612, just over ridge of mountain, north slope (36.72868°N, - 82.30478°W, 822 m), 10.vi.2005, 11:00 (coll: P. Marek) . 1♂ ( PMLN0020 ), Scott County, 7.2 km west of Fort Blackmore , 2.88 km north-west of junction routes 649 and 656 (36.7589°N, - 82.6770°W), 19.viii.1985 (coll: J. Mitchell) . 1♂ ( MMC0202 ) , 1♀ ( MMC0166 ), Scott Co., Quillin Spring , CR- 619, 160 m north of junction with CR-664 (36.69112°N, - 82.57847°W, 427 m), 19.viii.2006, 10:30 (colls: P. Marek and C. Spruill) . 1♂ ( MMC0256 ) , 1♀ ( MMC0245 ), Scott County, Flat Rock, Power’s Gap , VA-628, 0.2 rd km north of junction with VA-629 (36.60566°N, - 82.82960°W, 540 m) 26.ix.2006, 9:00 (colls: P. and B. Marek) . 1♀ ( MMC0080 ), Washington County, Clinch Mountain, CR-612 below Fugate’s Gap , 3.84 rd km south of junction with CR-613 (36.72894°N, - 82.30668°W, 813 m), 10.viii.2006, 15:30 (colls: P. Marek and C. Spruill) .

BRACHORIA OCHRA ( CHAMBERLIN, 1918) ‘THE MISSISSIPPI MIMIC MILLIPEDE’

( FIG. 32A–D, FIG . 41J–K)

Fontaria ochra Chamberlin, 1918: 123 . Brachoria sequens Chamberlin, 1939 ; synonymized by Keeton, 1959. Anfractogon tenebrans Hoffman, 1948 ; synonymized by Keeton, 1965. Brachoria ochra – Chamberlin & Hoffman, 1958. I agree with the synonymy of B. sequens and B. tenebrans with B. ochra . I have seen the types (USNM) and agree with Keeton that they do not exhibit significant enough differences from B. ochra to warrant nomenclatural status.

Diagnosis: Adult males of B. ochra are distinct from other Brachoria species based on the combination of: Colour. Combined striped and two-spotted ( Fig. 41J), purple-grey metatergal stripes with white caudal border, pinkish-red paranotal spots, legs. Collum purple, with pinkish-red paranota, and black central spot. Two-spotted ( Fig. 41K), red paranotal spots, anterior collum spot, legs. Exoskeleton. Caudolateral corners, paranota 5–19 with posterior-projecting nubbin, not rounded as in most Brachoria species. Lateral paranotal edges with curved aileron-like flange, projecting posterior past medial caudal edge. Gonopods. Gonopodal acropodite narrow, half width of tibia on leg pair 9, of uniform width throughout. Acropodite curved laterally at base, then medially in an oval-shaped form; not curved anteriorly, as in other species. Arc immediately distal beyond cingulum not elbowed as in B. initialis ( Fig. 26A , arrow), continuous oval-shaped form ( Fig. 32D ). Acropodal apex ( Fig. 32B ) distal to postcingular arc projecting dorsally, with curved sickle-shaped apex (not paring knife-shaped as in B. initialis , Fig. 26C ). Apex with raised shelf-like carina at base, imparts an elbowed appearance. ( Fig. 32B , arrow) Some specimens with an accessory sawtooth-shaped denticle on the acropodal apex (the ‘ B. tenebrans ’ morph), or with strongly pronounced carina (the ‘ B. anfractogon ’ morph).

Variation: Brachoria ochra has the second largest distribution of any species in the genus ( B. initialis has the largest) from northern Alabama to northeastern Mississippi. There is some variation in coloration amongst individuals, with two morphs present: two-spotted ( Fig. 41K) red and combined ( Fig. 41J) striped-spotted purple-grey and red (at Bankhead National Forest both morphs co-occur). There is some variation in length and width amongst these specimens, to a greater extent in males than in females. Measurements: ♂ (N = 9) BL = 33.68–46.99 (40.54/5.16). CW = 6.20–8.78 (7.53/1.01). IW = 4.30– 5.56 (4.87/0.43). ISW = 1.25–1.70 (1.47/0.19). B10W = 8.79–12.10 (10.42/1.19). B10H = 5.05–7.20 (6.02/0.70). ♀ (N = 3) BL = 37.64–40.47 (38.60/1.62). CW = 6.90–7.45 (7.08/0.32). IW = 4.99–5.20 (5.06/ 0.12). ISW = 1.44–1.60 (1.50/0.09). B10W = 10.25– 10.30 (10.28/0.03). B10H = 6.30–6.75 (6.53/0.23).

Description: Male described by Chamberlin (1918).

Ecology: Brachoria ochra specimens were collected during the day (17:30, Tombigbee locality) in a mixed evergreen and deciduous forest about 50 m south of Choctaw Lake. Millipedes were found beneath a layer of decaying oak leaves and pine duff. Brachoria ochra individuals were also collected from the banks of Bordon Creek in Bankhead National Forest. Here, individuals were not found under leaves but on top of them, actively walking about. Other xystodesmids encountered sympatrically at these localities were Dicellarius bimaculatus fictus ( Chamberlin, 1943) (Tombigbee National Forest) and C. g. latassa (Bankhead National Forest), the second of which appears similar in coloration to B. ochra . The species D. b. fictus, a uniform pale white colour, does not appear similar to B. ochra . At Bankhead National Forest, C. g. latassa has two colour morphs: striped yellow and striped red. The striped red colour morph appears similar to B. ochra (to the combined striped-spotted purple-grey and red morph). Note: individuals of B. ochra may be confused in the field with C. g. latassa because of similarity in colour patterns and broad overlap in distribution, but can be distinguished by the presence of the following characters (viewed with a 20¥ loupe): dorsal cuticle smooth, not rugose like C. g. latassa; acropodite oval-shaped, not straight as in C. g. latassa; acropodite projecting straight from prefemur, whereas in C. g. latassa (and other individuals in the tribe Rhysodesmini ) acropodite is bent at 90° from prefemur. Individuals of B. ochra may also be confused in the field with B. initialis because of similarity in colour patterns and overlap in distribution (co-occurring in northern Alabama, and northeastern Mississippi), but can be distinguished by the presence of the following characters (viewed with a 20¥ loupe): arc immediately distal beyond cingulum not elbowed as in B. initialis ( Fig. 26A , arrow), continuous oval-shaped form ( Fig. 32D ); acropodal apex ( Fig. 32B ) curved sickle-shaped (not paring knifeshaped as in B. initialis Fig. 26C ); apex with raised shelf-like carina at base ( Fig. 32B , arrow), imparts an elbowed appearance, not curved 90° as in B. initialis ( Fig. 26A ). Some specimens with an accessory sawtooth-shaped denticle on acropodal apex, or a strongly pronounced carina.

Distribution: The second largest distribution of any Brachoria species. Known from northern Alabama, south to Tuscaloosa, west into north-eastern and central Mississippi to Holmes County. Not extensively distributed throughout the coastal plain, as in B. initialis .

Material examined: Type specimens. ♂ holotype ( RVCC) from Oktibbeha County, Mississippi, ‘ Mississippi Agricultural College’ (Mississippi State University), Fall 1916 (coll: J. Bailey). The type specimen is missing from the collection. Nontype specimens. Alabama : 1♂ ( PMLN0166 ), Cullman County, wooded hillside, Wilburn (33.9518°N, - 87.0342°W), 20.x.1962 (coll: L. Hubricht) . 2♂ (SPC000077: DQ490655 , SPC000078), Lawrence County, Bankhead National Forest, Black Warrior WMA, Sipsey River picnic grounds area, Bordon Creek Trail , about 8 rd km west of junction Cranal and Bunyan Hill rds (34.30959°N, - 87.39433°W, 185 m), 12.vi.2003, 10:00 (colls: P. Marek and B. Hendrixson) . 1♂ ( PMLN0001 ), Lawrence County, King’s Cove , hemlock-beech woods (34.9698°N, - 85.7453°W), 29.vi.1950 (coll: G. Ball) . 1♀ (A3142), Morgan County, 12.8 km south of Decatur, on AL-41 at Flint Creek (34.5007°N, - 86.9673°W), 24.v.1980 (coll: R. Shelley) . 1♂ (SPC000937), Tuscaloosa County, Tuscaloosa, Rice Mine Rd. , 0.3 km east of junction with North Ridge Rd. , Black Warrior River bank (33.2341°N, - 87.5188°W, 52.7 m), 5.iii.2006 (coll: D. Beamer). Mississippi : 1♂, 2♀ (A4096), Pontotoc County, 1.6 km south-east of Ecru , mesic deciduous hillside, mixed woods (34.3357°N, - 89.0139°W), 19.v.1983 (coll: R. Shelley) . 1♂ (SPC000090), 1♀ (SPC000091), Choctaw County, Tombigbee National Forest, 4 air km east of Ackerman , Choctaw Lake , Chata Trail (33.27277°N, - 89.14778°W, 192 m), 16.vi.2003, 17:30 (colls: P. Marek and B. Hendrixson) . 1♂ ( PMLN0002 ), Holmes County, Holmes County State Park (33.0290°N, - 89.9162°W), 15.vii.1986 (coll: P. Lago) . 1♂ ( PMLN0003 ), Marshall County, mixed woods 3.2 km east of Slayden (34.9472°N, - 89.3960°W), 27.ii.1961 (coll: L. Hubricht) .

Figures

Figure 26. Brachoria initialis Chamberlin, 1939. A–D. Coopers, AL, specimen (PMLN0058) gonopods: A., left male gonopod, anterior view; B, left male acropodal apex, magnified; C, left male gonopod, medial; D, same, posterior (acropodite length = 1.82 mm).

Figure 29. Brachoria laminata 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 = 2.13 mm).

Figure 30. Brachoria ligula Keeton, 1959. A–D, holotype specimen gonopods: A, left male gonopod, anterior view; B, left male gonopod, medial; C, left male acropodal apex, magnified; D, same, posterior (acropodite length = 1.95 mm).

Figure 31. Brachoria mendota 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 = 1.97 mm).

Figure 32. Brachoria ochra (Chamberlin, 1918). A–D, Chata Trail, MS, specimen (SPC000090) 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).