A synopsis of the Neotropical genus Protoneura (Odonata: Coenagrionidae)
Author
Ellenrieder, Natalia Von
Author
Garrison, Rosser W.
text
Zootaxa
2017
2017-12-01
4361
1
1
76
journal article
31258
10.11646/zootaxa.4361.1.1
b15f5f11-6a00-4158-8ec8-3d0004605c28
1175-5326
1069337
53489D29-C68F-44FD-9EA2-CFCA7B949630
Protoneura cara
Calvert, 1903
Figs. 7
(
♂
habitus), 31 (
♀
habitus), 53 (
♀
mes. plate), 77 (gen. lig.), 103 (
♂
app.), 121 (map)
Protoneura cara
Calvert, 1903
: 140
, 143–144, pl. 5,
Figs. 38
,
45
, pl. 6, fig. 9 (in key, description of
♂
♀
, illustrations of
♂
S10 and body);—
Williamson (1915: 619, 624,
Figs. 5
,
19
; in key, illustration of wings and thoracic pattern)
;—
Johnson (1972: 73; characterization, counties in
Texas
,
USA
)
;—
Gloyd (1981: 133;
El Salvador
)
;—
Paulson (1982: 252;
USA
,
Mexico
,
Guatemala
,
El Salvador
)
;—
Paulson (1984: 51;
El Salvador
)
;—
González-Soriano & Novelo-Gutiérrez (1996: 162;
Mexico
)
;—
Abbott & Stewart (1998: 207;
USA
&
Mexico
)
;—
Abbott (2001: 196;
Texas
)
;—
González-Soriano (2001: 335– 339; notes on biology)
;—
Sibley (2003: 6;
Nicaragua
)
;—
Abbott (2005: 13, 49–51,
Figs. 6B
,
12
; characterization, notes on habits and behavior, illustrations of
♂
S10 and
♀
mesostigmal plates, distribution map in
Texas
)
;—
Urrutia (2005: 33;
Colombia
)
;—
Abbott (2006: 50, 56, 111; flight period and distribution map
Texas
)
;—
Westfall & May (2006: 431, 433, 436–438, Figs. 223D, 224B, 226B; in key to northern representatives of genus, characterization of adults, illustrations of
♂
thorax and S10,
♀
pronotum and mesostigmal plates)
;—
Abbott (2007: 20, 32, 89; flight period and distribution map
Texas
)
;—
Upson
et al.
(2007
: 28
;
Mexico
);—
Behrstock
et al.
(2007
: 32
;
Mexico
);—
Cuevas-Yañez (2007: 86, 87;
Mexico
)
;—
Abbott (2008: 23, 35, 93; flight period and distribution map
Texas
)
;—
Bailowitz
et al.
(2009
: 16
;
Mexico
);—
Paulson (2009h: 189 190;
Texas
, characterization, natural history, flight period, color photos)
;—González-Soriano
et al.
(2011: 184;
Mexico
);—
Bailowitz
et al.
(2012
: 9
;
El Salvador
);—
Garrison
et al.
(2010
: 379
);—
Abbott (2011: 100–101; characterization, notes on habits and behavior, color illustration of adults, picture of pair ovipositing, distribution map in
Texas
)
;—
González-Soriano & Paulson (2011: 303, 307;
Mexico
)
;—
González-Soriano & Novelo-Gutiérrez (2011: 2, Appendix VIII.21.2;
Mexico
)
;—
Escoto-Moreno
et al.
(2014
: 1046
, 1052;
Mexico
);—
Bailowitz
et al.
(2016
: 210
–211; habitat, characterization, field pictures of male, female, and pair, dot map of distribution in
Sonora
).
Primary
types
.
Holotype
♂
.
Mexico
,
Guerrero
,
Iguala
,
ix 1898
,
O.W. Barrett
leg. [
ANSP # 9239
].
Specimens examined.
84 ♂
20 ♀
:
U.S.A.
,
Texas
State, Kimble Co.
:
1 ♂
,
North Llano River
at
Farm
road 1674 (
30°30'58'' N
,
99°48'19'' W
,
506 m
),
14 vii 2001
,
J. Miller
leg. [
FSCA
]
;
Kerr Co.
:
4 ♂
1 ♀
, South Fork
Guadalupe
River near
Hunt
(
30°4'14'' N
,
99°20'15'' W
,
550m
)
18 vi 1985
,
S.W. Dunkle
leg. [
FSCA
]
;
2 ♂
, same data but [RWG]
;
1 ♂
, same data but
J.J. Daigle
leg. [RWG]
;
Real Co.
:
1 ♂
1 ♀
,
Frio River
at highway 337, E of
Leakey
{
29°43' N
,
99°45' W
,
475 m
},
14 vii 2001
,
K.J. Tennessen
leg. [
FSCA
]
;
1 ♂
,
Frio River
,
Garner State Park
{
29°34' N
,
99°43' W
},
19 vi 1979
,
C. Cook
leg. [
FSCA
]
;
Uvalde Co.
:
3 ♂
1 ♀
,
Sabinal River
at
Texas
highway 187,
3.7 mi
S of
Utopia
{
29°36' N
,
99°31' W
,
410 m
},
22 viii 1977
,
D.R. Paulson
&
S. Hills
leg. [
FSCA
]
;
Bexar Co.
:
1 ♂
1 ♀
(in tandem),
Medina River
at
Jungman Road
, W of
San Antonio
(
29°20'45'' N
,
98°47'11'' W
,
216 m
),
16 v 2001
,
S.W. Dunkle
leg. [RWG]
.
MEXICO
,
San
Luis
Potosí
State:
1 ♂
,
El Salto Falls
{
22°21'42'' N
,
99°15'53'' W
,
250 m
},
12 vi 1963
,
R. E. Woodruff
leg. [
FSCA
]
;
1 ♂
,
Coy River
headwaters,
28 km
S of
Ciudad Valles
(
21°43'59'' N
,
98°58'41'' W
,
42 m
),
21 viii 1999
,
R.W. Garrison
&
J.A. Garrison
leg. [
CSCA
]
;
1 ♂
1 ♀
(in tandem),
Municipio de Huehuetlán
,
Micos River
above
Cascada de Micos
,
2 km
N of
Micos
(
22°7'29'' N
,
99°10'46'' W
,
240 m
),
22 viii 1999
,
R.W. Garrison
&
J.A. Garrison
leg. [
CSCA
]
;
Nayarit
State:
1 ♂
1 ♀
,
Jumatán
{
21°30'34'' N
,
104°52'3'' W
,
935 m
},
13 iii 1981
,
E. González
leg. [RWG]
;
4♂
1 ♀
,
Tepic
{
21°28' N
,
104°53' W
,
966 m
},
5–11 xi 1923
,
J.H. Williamson
leg. [
UMMZ
]
;
1 ♂
1 ♀
, same data but [
FSCA
]
;
Michoacan
State:
1 ♂
,
Apatzingan
{
19°5' N
,
102°26' W
,
329 m
},
21 vii 1947
,
T.H. Hubbellleg.
leg. [
UMMZ
]
;
Morelos
State:
2 ♂
,
Lagunas Zempoala National Park
{
19°3' N
,
99°18' W
,
2,800 m
},
10–11 vii 1965
,
O.S. Flint
,
Jr. & M.A. Ortiz B.
leg. [
USNM
]
;
1 ♂
, stream in
Pueblo Cocoyotla
, highway 421 (
18°46'4'' N
,
99°27'26'' W
,
1,070 m
),
22 vii 1992
,
K.J. Tennessen
leg. [
FSCA
]
;
6 ♂
1 ♀
,
Xochitepec
{
18°45'53'' N
,
99°15'39'' W
,
1,100 m
},
12–14 vii 1965
,
O.S. Flint
&
M.A. Ortiz
leg. [
USNM
]
;
12 ♂
, route 95, km 91 near
Xochitepec
,
1 viii 1965
,
O.S. Flint Jr.
leg. [
USNM
]
;
1 ♂, Municipio Tlaltizapán, Las Estacas, spring and stream (18°43'48'' N, 99°6'48'' W,
980 m
),
8 vii 1987
, M.J. Westfall, Jr. leg. [FSCA]
;
1 ♂
,
Amacuzac River
at
Tehuixtla
and stream at
Balneario Las Palmas
(
18°32'51'' N
,
99°16'13'' W
,
866 m
),
20 vii 1992
,
S.W. Dunkle
leg. [
FSCA
]
;
1 ♂
,
1 ♀
, same data but
K.J. Tennessen
leg. [
FSCA
]
;
1 ♂
, same data but
W.F. Mauffray
leg. [
FSCA
]
;
Chiapas
State:
2 ♂
, E of
Cintalapa
,
21 km
at route 190 {
16°41' N
,
93°40' W
,
550 m
},
9 xii 1975
,
C.M. & O.S. Flint Jr.
[
USNM
]
;
4 ♂
3 ♀
, route 190, km 1024, beyond
Cintalapa
{
16°41' N
,
93°40' W
,
550 m
},
8–10 vii 1966
,
O.S. Flint
,
Jr. & M.A. Ortiz B.
leg. [
USNM
]
;
Oaxaca
State:
1 ♂
, route 190, km 698, near
El Coyul
{
16°29' N
,
95°53' W
,
19 m
},
11 vii 1966
,
O.S. Flint
,
Jr. & M.A. Ortiz, B.
leg. [
USNM
]
;
1 ♂
, stream
14 mi
E of
El Camarón
{
16°29' N
,
95°47' W
,
850 m
},
26 viii 1967
,
D.R. Paulson
&
M.L. Paulson
leg. [
FSCA
]
;
Veracruz
State:
2 ♂
,
Verde River
plane, route
140 km
368 {
18°56' N
,
97°2'0'' W
,
1,200 m
},
23 vii 1965
,
O.S. Flint
,
Jr. & M.A. Ortiz B.
leg. [
USNM
]
.
GUATEMALA
,
Alta Verapaz
Dept.:
1 ♀
,
Finca Semuc Champey
,
5 km
SSE of
Lanquín
{
15°32' N
,
89°57' W
},
3 v 1980
,
T.W. Donnelly
leg. [
FSCA
]
;
Zacapa
Dept.:
1 ♂
,
Gualán
{
15°6' N
,
89°21' W
,
141 m
},
14 i 1909
,
E.B. Williamson
leg. [
UMMZ
]
;
El Progreso
Dept.:
1 ♂
,
Estancia de La Virgen
{
14°56' N
,
89°53' W
,
320 m
},
11 viii 1965
,
O.S. Flint
,
Jr. & M.A. Ortiz B.
leg. [RWG]
;
Santa Rosa
Dept.:
2 ♂
,
Agua Caliente
{
14°55' N
,
90°24' W
,
569 m
},
2 vi 1909
,
E.B. Williamson
leg. [
UMMZ
]
;
Chiquimula
Dept.:
1 ♂
, near
Tierra Colorada
{
14°34'37'' N
,
89°20'20'' W
,
800 m
},
19 vii 1962
,
T.W. Donnelly
leg. [
FSCA
]
;
1 ♂
, small stream vicinity
Santa Elena
,
600 m
,
12 vii 1962
,
T.W. Donnelly
leg. [
FSCA
]
;
Escuintla
Dept.:
1 ♂
, stream
5.7 mi
SW of
Escuintla
{
14°22' N
,
90°49' W
,
671m
},
9 vii 1964
,
F.G. Thompson
leg. [
FSCA
]
;
Jutiapa
Dept.:
1 ♂
1 ♀
,
27.2 mi
ESE of
Chiquimulilla
{
14°5' N
,
90°22' W
,
215 m
},
3 ix 1964
,
F. G. Thompson
leg. [
UMMZ
]
;
1 ♂
, same data but [
FSCA
]
.
EL SALVADOR
,
La Libertad
Dept.:
4 ♂
2 ♀
(one pair in tandem),
4.7 mi
S of
Mizata
{
13°30' N
,
89°33' W
,
70 m
},
2 vii 1966
,
O.S. Flint
,
Jr. & M.A. Ortiz B.
leg. [
USNM
]
.
HONDURAS
,
Francisco
Morazán
Dept.:
1 ♀
,
Zamorano
,
Quebrada El Gallo
, small stream (
14°1' N
,
87°2' W
,
650 m
),
7 iii 1990
,
S.W. Dunkle
leg. [
FSCA
]
;
1 ♂
, Zamorano, Escuela Agrícola Panamericana,
30 km
ESE of
Tegucigalpa
, ponds and
Yeguari River
with riffles & pools,
14 xii 1987
,
L.
Córdoba
leg. [
FSCA
]
.
NICARAGUA
:
Chontales
Dept.:
1 ♂
, route 7, km 183,
4.2 mi
W of
Villa Somosa
{
12°3' N
,
84°59' W
,
200 m
},
29 vii 1967
,
O.S. Flint
,
Jr. & M.A. Ortiz B.
leg. [
USNM
]
.
PANAMA
,
Canal Zone
:
10 ♂
1 ♀
,
Sardanilla River
{
9°7' N
,
79°35' W
,
145 m
},
5 xii 1916
,
E.B. Williamson
leg. [
UMMZ
]
.
Characterization.
Male: Epicranium black with orange spots along margin of compound eyes, anterior margin of postfrons and occipital bar; black stripes on thorax; dorsum of abdomen brown to black with pale spots on S1–3, basal portion of S4–7, and distal margin of S9–10; pale colors red to orange and yellow (
Fig. 7
). Pronotum anterior, middle and posterior lobes black with medial and lateral orange spots. Mesepisternum black with middorsal carina orange, a antehumeral stripe along entire sclerite, and a narrow black stripe along distal 1/4 of mesepisternal-mesepimeral suture; mesepimeron black margined in orange; metepisternum orange with black along posterior margin; metepimeron orange to yellow with central dark area to black with only posterior margin orange. Coxa dark brown, remainder of leg mostly pale, yellow to orange, sometimes with two diffuse brownish rings subbasally and subapically on each femur and tibia; tibial spurs shorter than twice intervening spaces. Genital ligula lacking lateral lobes and with a slightly convex distal margin, with laterodistal corners projected ventrally (
Fig. 77
). Cercus about as long as S10 length, about as long as paraproct, longer than wide, with a depression on external dorsolateral surface, dorsal margin and ventral margin smoothly convex in lateral view (
Fig. 103a
), medial surface with a small blunt ventrobasal tooth, a large medial branch ending on a curved tooth separated by a membranous area from reminder of cercus, and a subapical dorso-ventral ridge, visible in dorsal and mediodorsal views (
Figs. 103b, c
). Paraproct about as long as S10 length, at midlength narrowing to half its basal width and ending in a bluntly pointed tip (
Fig. 103
). TL 36–38; Hw 16–17.7.
Female: As male but black more extensive on abdomen; pale lateral area of S9 extending along basal 1/3 of segment to about 2/3 of segment height posteriorly (
Fig. 31
). Middle lobe of pronotum lacking pronounced lateral depressions; posterior lobe directed dorsally and smoothly convex. Mesostigmal plate reniform and convex, with posteromedial corner contiguous through lateral arm of middorsal carina with middorsal carina, which is higher at convergence point than remainder of pterothorax; with a depression on mesepisternum behind plate (
Fig. 53
). TL 32.5–36.5; Hw 18–19.
Diagnosis.
Protoneura cara
can be distinguished from all congeners by the morphology of its male cercus, with a large medial branch ending on a curved tooth separated by a membranous area from remainder of cercus (
Figs. 103b, c
), and by female mesostigmal plate reniform and contiguous with elevated middorsal carina via lateral arms (
Fig. 53
).
Habitat and biology.
Adults found at pools and sluggish backwaters bordered by extensive overhanging bush and undercut root-balls (
Upson
et al.
2007
), at small, slow-flowing streams, sometimes in the open, sometimes with a dense overhead canopy (
Bailowitz
et al.
2009
), small clear streams with sand bottom in woodland or open (
Paulson 2009h
), and along shallow slow clear rivers (
Behrstock
et al.
2007
). According to
González-Soriano (2001)
, adults are commonly seen flying along streams and rivers in
Mexico
, either as solitary males or tandem pairs, and males patrol water surface searching for ovipositing females and switch from one perch to another without defending a particular area.
González-Soriano (2001)
observed only one pair in copula perching on grasses at the edge of a spring-fed pond and reported oviposition as occurring in tandem, with male remaining erect beating its wings, and tandem pairs flying along streams and rivers searching for oviposition sites consisting of submerged grasses, living roots, and floating objects such as small pieces of wood. He also observed males aggregating to form aerial mating swarms, hovering in a stationary fashion
4–5 m
above ground level,
2–3 m
from the stream edge at a forest gap surrounded by small trees with all hovering individuals facing in the same direction and displaying no foraging activity, apparently waiting for receptive females.
Paulson (2009h)
also reported seeing occasional mating swarms, with aggregations of males hovering well above head height in sunny clearings near water, and ensuing tandem formation when females entered swarm. He reported males spending much of their time hovering just above water but also perching on slender stems, and slowly rising into trees when disturbed. According to
Abbott (2011)
, egg laying typically occurs in tandem near the margin of slow moving water in floating and submerged vegetation and debris, with females curving abdomen between wings, and males typically hovering just above water for extended periods of time.
Paulson (2009h)
observed tandem pairs ovipositing in any sort of floating detritus, including tiny wood chips, typically where currents concentrate detritus, but also in grass stems and rootlets of woody plants near shore.
Bailowitz
et al.
(2016)
reported its flight period in
Sonora
,
Mexico
, as spanning from late April through late October, and
Paulson (2009h)
from June to October in
Texas
,
USA
.
Distribution.
Texas
State in
USA
to
Colombia
(
Fig. 121
).