A revision of the Appalachian millipede genus Brachoria Chamberlin, 1939 (Polydesmida: Xystodesmidae: Apheloriini)
Author
Marek, Paul E.
text
Zoological Journal of the Linnean Society
2010
2010-07-26
159
4
817
889
http://dx.doi.org/10.1111/j.1096-3642.2010.00633.x
journal article
10.1111/j.1096-3642.2010.00633.x
0024-4082
5755069
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
).
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).
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).
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).
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
)
.