Soyedina Ricker, 1952 (Plecoptera: Nemouridae) in the eastern Nearctic: review of species concepts, proposed morphology-based species groups, and description of a new species from North Carolina
Author
Grubbs, Scott A.
Author
Baumann, Richard W.
text
Zootaxa
2019
2019-08-22
4658
2
223
250
journal article
25938
10.11646/zootaxa.4658.2.2
349f1d0e-6dfc-443a-998d-0322ea62b866
1175-5326
3375788
A4D55B3B-9BE9-4918-89C6-8049F5895CF2
Soyedina carolinensis
(
Claassen, 1923
)
Carolina Forestfly
http://lsid.speciesfile.org/urn:lsid:
Plecoptera
.speciesfile.org:TaxonName:606 5
(
Figs. 25–34
,
80
)
Nemoura carolinensis
Claassen 1923:287
.
Holotype
♂
(MCZC),
Morganton
, (
Burke Co.
),
North Carolina
,
USA
Nemoura carolinensis
Needham & Claassen, 1925:215
Nemoura
(
Soyedina
)
washingtoni carolinensis
Ricker, 1952:53
Soyedina carolinensis
Illies, 1966:249
Soyedina carolinensis
Stark, 2017:187
Distribution.
USA
: DE,
MD
,
NC
, PA,
TN
, VA,
WV
(
DeWalt
et al
. 2019
).
Material examined.
USA
,
Maryland
,
Frederick Co.
, spring seep
Raven Rock Run
,
17 March 1990
,
R.M. Duffield
, 1 pharate male,
10 larvae
(
CHNC
)
;
Fishing Creek
,
15 March 1991
,
R.M. Duffield
,
1 male
,
1 female
(
CHNC
)
;
tributary to
Steep Creek
,
39.56006
,
-77.48271
,
9 March 1997
,
S.A. Grubbs
,
1 male
,
2 females
(
WKUC
)
.
North Carolina
,
Buncombe Co.
, stream,
Tanbark
(
Ridge
)
Tunnel
,
Blue Ridge Parkway
,
5 March 1991
,
R.W. Baumann
&
S.M. Clark
,
2 males
(
BYU
)
;
spring into
Ox Creek
,
Blue Ridge Parkway
,
35.67078
,
-82.47066
,
18 April 2009
,
S.A. Grubbs
,
1 male
(
WKUC
)
;
Graham Co.
, spring seeps, FS 81,
Nantahala National Forest
,
35.34784
,
-83.99531
,
1 March 2008
,
A.L. Sheldon
,
1 male
(
WKUC
)
;
seep into
Sand Creek
, FS 81F,
Nantahala National Forest
,
35.36208
,
-83.99576
,
2 March 2008
,
A.L. Sheldon
,
1 male
(
WKUC
)
;
Haywood Co.
,
Haywood Co.
,
GRSM
ATBI
Plot
,
Purchase Knob
, (
malaise trap
),
35.59004
,
-83.06024
,
7–27 March 2002
,
P.F. Super
&
J. Lowe
,
3 males
,
9 females
(
BYU
)
;
Big
East Fork
,
Pigeon River
,
14 March 2006
,
J. Robinson
,
1 male
,
2 females
(
BYU
)
;
tributary to
Laurel Creek
,
Pisgah National Forest
,
35.75263
,
-83.09060
,
1 March 2008
,
S.A. Grubbs
,
7 males
,
3 females
(
WKUC
)
;
tributary to
Laurel Creek
,
Pisgah National Forest
,
35.75255
,
-83.08755
,
1 March 2008
,
S.A. Grubbs
,
4 males
,
4 females
(
WKUC
)
;
tributary to
Flat Branch
,
Pisgah National Forest
,
35.74612
,
-83.07928
,
1 March 2008
,
S.A. Grubbs
,
1 male
,
1 female
(
WKUC
)
;
Cold Springs Creek
, FR 148,
Pisgah National Forest
,
35.77850
,
-82.95590
,
2 March 2008
,
1 male
(
WKUC
)
;
same but
18 April 2009
, S.A.
Grubbs
,
1 male
,
1 female
(
WKUC
)
;
tributary to
Hemphill Creek
,
35.57690
,
-83.07230
,
2 March 2008
,
S.A. Grubbs
,
14 males
,
6 females
(
WKUC
)
;
Jackson Co.
,
3 mi.
SW Cashiers
,
Whiteside Mountain
,
9 May 1997
,
C.R. Nelson
,
1 male
(
BYU
)
;
Macon Co.
,
Rattlesnake Springs
,
Nantahala National Forest
,
35.17060
,
-83.57741
,
18 March 1982
,
J. Weaver
,
R. Holzenthal
,
2 males
(
BPSC
)
;
same but
1–18 May 1983
, J.
Weaver
& R.
Holzenthal
,
31 males
,
93 females
(
BYU
)
;
same but
23 March 2002
, S.A.
Grubbs
,
6 males
,
8 females
(
WKUC
)
;
tributary to
Jarrett Creek
,
Nantahala National Forest
,
35.15903
,
-83.62873
,
23 March 2002
,
S.A. Grubbs
,
1 male
,
5 females
(
WKUC
)
;
same but
8 March 2004
, S.A.
Grubbs
,
1 male
,
1 female
(
WKUC
)
;
Robin Branch
,
Nantahala National Forest
,
35.16832
,
-83.58465
,
B.P. Stark
,
19 May 1983
,
1 male
(
BYU
)
;
same but K.W.
Stewart
,
1 March 1986
,
1 male
,
1 female
(
BYU
)
;
same but
28 February 2008
, A.L.
Sheldon
,
1 male
,
1 female
(
WKUC
)
;
same but
15 March 2009
, B.P.
Stark
,
3 males
, 3 pharate males,
1 female
(
BPSC
)
;
upper Ball Creek
,
Coweeta Hydrologic Laboratory
,
12 March 1983
,
A.D. Huryn
,
1 male
(
ADHC
)
;
same but
7–20 April 1984
, A.D.
Huryn
,
1 male
(
BYU 1824
)
;
Kit Springs Branch
, FS 711,
Nantahala National Forest
,
35.20191
,
-83.58354
,
29 February 2008
,
A.L. Sheldon
,
1 male
(
WKUC
)
;
Sassafras Branch
, FS 711,
Nantahala National Forest
,
35.20230
,
-83.59406
,
29 February 2008
,
A.L. Sheldon
,
1 male
(
WKUC
)
;
seep into
Split Whiteoak Branch
, FS 711,
35.20458
,
- 83.58046
,
29 February 2008
,
A.L. Sheldon
,
1 male
(
WKUC
)
;
tributary to
Split Whiteoak Branch
, FS 711,
Nantahala National Forest
,
35.20640
,
-83.59059
,
29 February 2008
,
A.L. Sheldon
,
3 males
,
2 females
(
WKUC
)
;
Blue Valley
Overlook,
Hwy
106,
12 March 1995
,
B.P. Stark
,
R. Simmons
& R.
Frederick
,
3 males
(
BYU
)
;
same but
14 March 2006
, B.P.
Stark
,
4 males
(
BPSC
)
;
Swain Co.
, tributary to
Connelly Creek
,
Nantahala National Forest
,
35.35226
,
-83.35175
,
23 March 2002
,
S.A. Grubbs
,
1 male
,
5 females
(
WKUC
)
;
Yancey Co.
,
Big Crabtree Creek
, FR 5511,
Pisgah National Forest
,
35.79264
,
-82.17388
,
24 March 2002
,
S.A. Grubbs
,
2 males
,
7 females
(
WKUC
)
.
Pennsylvania
,
Chester Co.
,
East Branch
,
White Clay Creek
,
0.6 mi.
W London Grove
,
24 February 1980
,
D.H. Funk
,
2 males
,
1 female
(
BYU
)
;
same but
March 6 1980
, D.H.
Funk
,
7 males
,
2 females
(
BYU
)
;
tributary,
White Clay Creek
,
Stroud Laboratory
,
26 April 1964
,
J.K. Gelhaus
&
D P. Koenig
,
11 males
,
22 females
(
BYU
)
;
[
Philadelphia Co.
],
Philadelphia
,
24 April 1910
,
G.M. Greene
,
1 male
(removed from pin) (
USNM
)
.
Tennessee
,
Carter Co.
,
Twin Springs
,
Cherokee National Forest
,
36.12128
,
-82.08654
,
10 April 2002
,
S.A. Grubbs
,
1 male
,
1 female
(
WKUC
)
;
Sevier Co.
,
Middle Prong Little Pigeon River
,
Great Smoky Mountains National Park
,
22–23 April 1978
,
C.H. Nelson
,
2 males
,
3 females
(
CHNC
)
;
Ramsay Prong Little Pigeon River
,
Great Smoky Mountains National Park
,
22–23 April 1978
,
C.H. Nelson
,
2 males
,
2 females
(
CHNC
)
;
Gatlinburg
,
Great Smoky Mountains National Park
,
28 April 1978
,
F.F. Sherberger
,
1 male
,
1 female
(
BYU
)
;
Cades Cove
,
3.5 mi.
E Cabin Mill
,
Great Smoky Mountains National Park
,
25 March 1980
,
B.P. Stark
,
K.W. Stewart
& J.
Feminella
,
4 males
,
4 females
(
BYU
)
;
Rocky Spur Branch
,
Great Smoky Mountains National Park
,
35.68027
,
-83.46450
,
25 March 1999
,
B.C. Kondratieff
,
3 males
,
3 females
(
WKUC
)
;
Road Prong
,
Great Smoky Mountains National Park
,
35.60975
,
-83.44918
,
15 April 2014
,
S.A. Grubbs
,
1 male
,
1 female
(
WKUC
)
.
Virginia
,
Bedford Co.
,
Hemp Mill Branch
,
Sheep Creek
,
37.40222
,
-79.65056
,
25 February 1980
,
C.F. Burgoon
&
C.E. Dunn
,
2 males
(
BYU
)
;
Floyd Co.
,
Tory Creek
,
Hwy
58, west of
Meadows of Dan
,
36.72798
,
-80.47042
,
11 March 1991
,
R.W. Baumann
&
R.F. Kirchner
,
6 males
(
BYU
,
WKUC
)
;
Grayson Co.
, spring on top of
Straight Mountain
,
Rt.
603,
22 April 1979
,
B.C. Kondratieff
,
1 male
,
1 female
(
BYU 1941
)
;
unnamed spring,
Rt.
603, near
Lewis Fork Bridge
,
18 February 1990
,
R.F. Kirchner
&
K.W. Stewart
,
8 males
,
4 females
(
BYU
)
;
seep into
Lewis Fork
,
Jefferson National Forest
,
36.66883
,
-81.53128
,
13 April 2008
,
A.L. Sheldon
,
5 males
,
2 females
(
WKUC
)
;
same but
36.67208
,
-81.52467
,
13 April 2008
, A.L.
Sheldon
,
2 males
,
2 females
(
WKUC
)
;
same but
36.66790
,
-81.53016
,
13 April 2008
, A.L.
Sheldon
,
2 males
(
WKUC
)
;
Madison Co.
,
Rapidan River
,
Criglersville
,
27 March 1938
,
B.D. Burks
,
2 males
, (
INHS
Plecoptera
7755)
;
Rapidan River
,
Rapidan Camp, R.M
.
Duffield
,
30 April 1988
,
1 male
(
CHNC
)
;
spring into
Hogcamp Brook
,
Dark Hollow Falls
Trail,
Shenandoah National Park
,
38.51867
,
-78.42349
,
15 May 1995
,
S.A. Grubbs
,
1 male
,
4 females
(
WKUC
)
;
Washington
Co.
, spring, FR 90,
Mount Rogers National Recreation Area
,
36.65629
,
-81.70547
,
22 March 2002
,
S.A. Grubbs
,
2 males
,
2 females
(
WKUC
)
.
West Virginia
,
Randolph Co.
,
Marsh Fork of Buckhannon River
,
Route
47 near
Pickens
,
38.66300
,
-80.23180
,
24 March 1992
,
S.M. Clark
,
13 males
,
23 females
(
BYU
)
.
Remarks.
The adult male was described and illustrated in
Claassen (1923)
and subsequently in
Needham & Claassen (1925
, with the same image of the male in lateral view; their plate 38, fig. 7).
Ricker (1952)
demoted
S. carolinensis
as a subspecies of
S. washingtoni
(both as subgenera of
Nemoura
), noting that he found the two taxa as nearly indistinguishable. Full species status, however, was re-elevated by
Illies (1966)
. The combination of the asymmetric dorsal (
Figs. 25–27, 30
) and ventral sclerites (
Figs. 25–26
), plus the inner epiproct member that is flanged and directed laterally with the flange lip approximately same width apically (
Figs. 25–27
), aligns this species closely with
S. merritti
and
S. washingtoni
. We view
S. carolinensis
as a valid, albeit variable, species that is distinct from
S. washingtoni
. Although the outer paraproct lobe of
S. carolinensis
is variable in form (see below), for
S. washingtoni
it is consistently quadrate or subquadrate across its range (
Figs. 49–50
).
The outer paraproct lobe of
S. carolinensis
varies in form. For example, early-emergent populations (i.e. late winter–early spring) from east
Tennessee
and western
North Carolina
have an outer lobe that has a narrow profile in apical ½ with anterior and posterior margins ca. parallel and without a basoposterior expansion (
Fig. 31
). This is a very common phenotype. A slight variation in this theme is present on populations from central
West Virginia
(
Fig. 32
). In contrast, populations from higher-elevation streams and seeps in southwestern
Virginia
and east
Tennessee
have an outer paraproct lobe that is more sickle-shaped and with a distinct anteriorly-directed spur compared to other phenotypes (
Fig. 33
). Populations further northward along the Blue Ridge and upper Piedmont provinces into central
Maryland
and southeastern
Pennsylvania
have an outer lobe that is different along the anterior and posterior margins, and has a more bulbous basoposterior portion (
Fig. 34
). Because of this, we contend that morphology alone cannot effectively address the question regarding of if there are evolutionarily independent units of
S. carolinensis
that represent undescribed species. Although the larva is undescribed,
Stark (2017)
noted one cercal feature that is distinct from that of
S. vallicularia
.
Soyedina carolinensis
is known from the southern Appalachian Mountain region northeastward to the Piedmont Uplands of southeastern
Pennsylvania
and northern
Delaware
(
Fig. 80
).