Nephepeltia flavifrons ( Karsch, 1889 )

Ellenrieder, Natalia Von, 2014, A synopsis of the Neotropical genus Nephepeltia (Odonata: Libellulidae), including description of a new species, synonymies, and a key to males, Zootaxa 3796 (1), pp. 121-146 : 125-128

Nephepeltia flavifrons ( Karsch, 1889)

Figures 2 c–d; 3c–d; 4; 7; 9c; 10d–f; 11c; 13c–f; 14a–d; 15

Nannothemis prodita Hagen, 1861: 320 (nomen nudum);— Hagen (1877: 94; synonymy with N. inermis );— Calvert (1906: 230; synonymy with N. flavifrons based on examination of Hagen’s ♀ specimen from Pernambuco).

Nannothemis inermis Hagen, 1867: 90 (nomen nudum);— Hagen (1877: 94; synonymy with N. prodita );— Calvert (1906: 230; synonymy with N. flavifrons based on examination of Hagen’s ♂ specimen).

Neothemis flavifrons Karsch, 1889: 256 –257 (description based on one ♂ from Lambaré, Central Dep., Paraguay {25°20'59''S, 57°38'37''W, 47 m}, R. Rohde leg., deposited in ZMHB).

Nephepeltia flavifrons — Calvert (1906: 230–231; transference of species to genus, diagnosis from N. phryne , description of ♀);— Calvert (1909: 243; notes on ♂ and ♀ from Brazil, Cuiabá, Cachoeira, and Rio Grande do Sul);— Ris (1911: 393, 395–396, figs. 241, 243; in key, notes on additional specimens, illustrations of ♂ wings and genital fossa);— Santos (1950: 21–26, figs. 3–4, 13, 16, 35–36, 40, 42, 46; redescription based on ♂ and ♀ from Brazil; illustrations of ♂ anterior lamina, genital fossa, S10, and vesica, and of ♀ pronotum and S9–10);— Rácenis (1953: 135, records from Venezuela);— Geijskes (1971: 670; record from French Guiana);— Costa et al. (2000: 16, records from São Paulo State, Brazil);— Förster (2001: 116; in key for Central America).

Nephepeltia chalconota Ris, 1919: 1131 (description based on 5 ♂ and 1 ♀ from Puerto Barrios, Guatemala);— Santos (1950: 13–15, transcription of Ris’ description);— Boomsma & Dunkle (1996: 27; record from Belize); González-Soriano & Novelo-Gutiérrez (1996: 167; mention from Oaxaca and Veracruz States, Mexico);— González-Soriano (1997: 255; records from Veracruz State, Mexico);— Förster (2001: 117; in key for Central America);—González-Soriano & Novelo- Gutiérrez (2011: 7 in Appendix VIII.21.2; records from Veracruz State, Mexico). N ew synonymy.

Specimens examined. Total: 218 ♂, 61 ♀: PARAGUAY, Guaira Dep.: 1 ♂, Villarica {25°45'S, 56°26'W, 125 m}, 17 i 1947, F.H. Schade leg. [MLP]. ARGENTINA, Corrientes Prov.: 1 ♀, Bañado Pariopá and stream, on prov. road 94, aprox. 12 km N of Santo Tomé (28°26'53''S, 56°00'34''W), 24 ii 2003, P. Pessacq & J. Muzón leg. [MLP]; 1 ♂, Ituzaingó, Natural Reserve Rincón de Santa María, artificial ponds (27°28'23''S, 56°34'38''W), 30 ix 2003, J. Muzón & P. Pessacq leg. [MLP]; 1 ♂, Laguna Iberá, Colonia Pellegrini (28°32'16''S, 57°11'12''W, 57 m), 2003, J. Muzón & P. Pessacq leg. [MLP]; 1 ♂, Estancia El Dorado (28°44'34''S, 58°7'36''W), 20–21 ii 2003, P. Pessacq & J. Muzón leg. [MLP]. BOLIVIA, Santa Cruz Dep.: 2 ♂, Guarayos Prov., Ascención de Guarayos, 13 km south on highway 9, Progreso Hacienda laguna and wetlands (15°59'24''S, 63°07'10''W, 234 m), 17–18 viii 2003, B. Mauffray leg. [FSCA]; 1 ♀, same data but 5 km N on highway 9, small hillside seepage stream (16°29'S, 60°57'W, 427 m), 27 viii 2003 [FSCA]; 2 ♂, Velasco Prov., pond NW of El Carmen de la Ruiza, 14 km NW on highway to Concepción, marsh and pond (16°02'10''S, 61°05'56''W 345 m), 11 xi 1999, B. Mauffray leg. [FSCA]; Warnes Prov.: 1 ♀, 10 km E Warnes (17°40'S, 63°00'W, 329 m), 19–23 viii 1983, R. Wilkerson leg. [FSCA]. BRAZIL, Rio Grande do Sul State: 2 ♂, Imbé municipality, Braço Morto, channel near restaurant (29°58'05"S, 50°07'12"W, 1 m), 19 ii 2004, A.P. Pinto & J.G. Da Silva leg. [DZRJ-847–848, not examined]; 1 ♂, Guaíba muncipality, Sítio Prof. Matzembacher (30°10'48"S, 51°23'27"W, 32 m), 20 xi 2004, A.P. Pinto leg. [DZRJ-853, not examined]; 1 ♂, Pelotas {31°46'S, 52°20'W, 3 m}, 4 iii 1959, C. Biezanko leg. [FSCA]; Santa Catarina State: 1 ♂, Papanduva municipality, 02–07 ii 2005, N. Ferreira, Jr. leg. [DZRJ-1981, not examined]; Minas Gerais State: 1 ♂, Vespasiano (19°40'S, 43°55'W, 870 m, ii 1954), A.B.M. Machado leg. [FSCA]; São Paulo State: 1 ♂, Sitio Primavera, Rio Claro (22°24'33''S, 47°36'44''W, 569 m), 13 ii 2002, F.A.A. Lencioni leg. [RWG]; 1 ♂, 1 ♀, Jacareí Municipality, Fazenda Santana do Rio Abaixo (23°14'52''S, 46°59'58''W, 624 m), 23 xii 2000, F. & F.A.A. Lencioni leg. [RWG]; Bahia State: 2 ♂, Barreiras Municipality (16°04'S, 39°20'W, 72 m), 4 i 1908, J.D. Haseman leg. [CMNH]; 1 ♂, Lauro de Freitas municipality, K13, Encontro das Aguas (12°51'47''S, 38°18'50''W, 23 m), 22 iv 2006, R. Penalva leg. [RWG]; 1 ♂, same but 18 ii 2005 [RWG]; Rondônia State: 1 ♂, 1 km N Cacaulândia on B-65, linea B-65, ponds near Fernandes trail (10°19'25''S, 62°54'00''W, 164 m), 16 xi 1991, M. J. Westfall, Jr. leg. [FSCA]; 3 ♂, Governador Jorge Teixeira Municipality, Fazenda Rancho Grande, Linea C-20, 7 km E of B-65 (10°31'48''S, 62°48'00''W, 165 m), 16 xi 1991, M.J. Westfall, Jr. [FSCA]; 3 ♂, same data but 8–20 iv 1997, A.C. Rehn & C. Alexander leg. [ACR]; 1 ♂, same data but farm of Harald Schmitz and environs within about 15 km radius, 16–24 iii 1989, S.W. Dunkle leg. [FSCA]. SURINAME, Sipaliwini Dis.: 1 ♂, Kwamalasamutu, ditch at village (02°21'17''N, 56°47'11''W, 211 m), 8 ix 2010, NvE leg. [CSCA]. PERU, Loreto Dep.: 1 ♂, Tamshiyacu-Tahuayo Reserve, Tabano lake, lake with water lettuce (04°20'19''S, 73°15'38''W), 4 iii 2010, TF [TF]; 4 ♂, same data but [USNM]; 1 ♂, Explorama Lodge, 50 mi NE Iquitos on Amazon River at junction with Yanamono River, Manatí river and pasture pond at Lodge (03°21'59''S, 72°47'56''W, 103 m), 16 viii 1989, S.W. Dunkle leg. [RWG]; 4 ♂, Explornapo Camp at 100 mi NE Iquitos, at Lorenso, a blackwater lake (03°16'33''S, 72°56'18''W, 97 m), 14–15 vii 1990, S.W. Dunkle & A. Menke leg. [RWG]; 8 ♂, same data but [FSCA]; 1 ♂, same data but [CSCA]; 4 ♂, Aguas Negras (00°31'22''S, 75°15'24''W, 145 m}, 13–16 iii 1994, J. Louton leg. [RWG]. ECUADOR, Napo Prov.: 6 ♂, Lago Anangu, near Providencia (00°31'32''S, 76°26'22''W, 300 m), 9 v 1975, T.C. Emmel leg. [FSCA]. VENEZUELA, Falcón State: 3 ♂, Tucacas {10°47'52''N, 68°19'03''W}, 23 iii 1920, J.H., E.B. Williamson & W.H. Ditzler leg. [UMMZ]; Zulia State: 1 ♂, 1 ♀, Encontrados {09°03'51''N, 72°13'55''W, 13 m}, 26 iv 1920, J.H. & E.B. Williamson & W.H. Ditzler leg. [UMMZ]; 4 ♂, same data but [FSCA]; Guárico State: 1 ♂, Caño Caracol, Fundo Masaguaral, 45 km south of Calabozo (08°34'12''N, 67°34'48''W, 69 m), 18 i 1983, O.S. Flint Jr. leg. [RWG]; 1 ♂, same but 1 iii 1986, R.B. Miller & L.A. Stange [FSCA]; Apure State: 2 ♀, Hato San Leonardo, 125 km SW of San Fernando on Río Capanaparo (07°01'30''S, 67°34'00''W, 160 m), 18–24 ii 1997, G.A. Kareofelas leg. [ACR]. COLOMBIA, Magdalena Dep.: 25 ♂, 14 ♀, Río Frío {10°55'N, 74°10'W, 43 m}, 4–8 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 2 ♀, same data but [FSCA]; 1 ♂, 1 ♀, same data but 30 iii–11 iv 1925, F.W. Walker leg. [UMMZ]; 1 ♂, Sevilla, Guatemala Farm {10°46'N, 74°09'W, 33 m}, 3 xii 1925, F.W. Walker leg. [UMMZ]; 75 ♂, 10 ♀, Fundación {10°31'17''N, 74°11'12''W, 45 m}, 10–13 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 2 ♂, same data but [FSCA]; 1 ♂, Puerto Colombia, Arroyo Grande {09°43'N, 74°26'W, 93 m}, 10 xii 1916, J.H.& E.B. Williamson leg. [UMMZ]; 5 ♂, 4 ♀, El Banco {09°02'50''N, 73°58'41''W, 46 m}, 23–24 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 3 ♂, same data but [RWG]; Bolivar Dep.: 2 ♀, Zambrano {09°44'58''N, 74°49'05''W, 5 m}, 20 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 2 ♂, 2 ♀, near Magangue {09°14'29''N, 74°45'12''W, 21 m}, 21 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 1 ♂, Río Nuevo, between Magangue and El Banco, on lower Magdalena river Valley {08°29'N, 74°26'W, 46 m}, 22 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 1 ♀, Boyaca, Sogamoso, 134 leagues from Barranquilla {05°43'14''N, 75°55'47''W, 2,455 m}, 29 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; Córdoba Dep.: 1 ♂, Monteria {08°45'27''N, 75°53'24''W, 24 m}, 3 ix 1980, S.W. Dunkle leg. [FSCA]; 1 ♀, same data but 1 ix 1980, may have been ovipositing in flight over a grassy bank [RWG]; Cesar Dep.: 3 ♂, 10 ♀, Gamarra {08°19'45''N, 73°44'40''W, 32 m}, 27 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; Antioquia Dep.: 1 ♀, Sautata, M.A.Carriker {07°49'41''N, 77°03'55''W, 3 m}, 30 i 1918, J.H. & E.B. Williamson leg. [RWG]; 1 ♂, Casabe {07°00'28''N, 73°54'51''W, 70 m}, vii 1961, native collectors for M. Bos [FSCA]; 2 ♂, Boca Murindo {06°59'26''N, 76°45'24''W, 21 m}, 10 ii 1918, M.A. Carriker leg. [UMMZ]; 4 ♂, Puerto Berrio {06°30'N, 74°24'W, 69 m}, 8 ii 1917, J.H. & E.B. Williamson leg. [UMMZ]; 1 ♂, 1 ♀, Cristalina {06°29'N, 74°50'W, 320 m}, 17–19 ii 1917, J.H. & E.B. Williamson leg. [UMMZ]. COSTA RICA, Heredia Prov.: 1 ♀, Puerto Viejo, La Selva Biological Station (10°28'N, 84°01'W, 60 m), 16 viii 1996, C.l. & S.L. Staines leg. [FSCA]; Limón Prov.: 1 ♂, swampy pond 1.2 km W of Río Parismina on road to Guapiles, small stream partly dammed by road (10°13'N, 83°47'W, 268 m), 5 iii 1987, S.W. Dunkle leg. [FSCA]. GUATEMALA, Suchitepequez Dep.: 2 ♂, 19 mi W Siquinala, pond {14°12'31''N, 91°14'02''W, 180 m}, 24 vi 1966, D.R. Paulson leg. [FSCA]; Escuintla Dep.: 1 ♂, Finca San Bernardo, Cushin I (14°18'18''N, 90°47'06''W, 275 m), 3 vi 1986, M. Dix leg [FSCA]; Izabal Dep.: 1 ♂ syntype of N. chalconota Ris, 1919 , Puerto Barrios {15°43'N, 88°36'W, 2 m}, 26 v 1909, E.B. Williamson leg. [FSCA]; 2 ♂ syntypes of N. chalconota Ris, 1919 , same data but [UMMZ]. BELIZE, Toledo Dis.: 2 ♂, 2 ♀, Río Temash {15°58'35''N, 88°56'31''W, 11 m}, v 1934, J.J. White leg. [UMMZ]; 1 ♂, 1 ♀, Punta Gorda (16°06'10''N, 88°50'00''W, 18 m}, vi 1933, J.J. White leg. [FSCA]; 1 ♀, same data but [UMMZ]. MEXICO, Tabasco State: 1 ♀, Villahermosa {17°58'N, 92°55'W, 8 m}, 17 iii 1964, R. Duncan Cuyler leg. [FSCA]; Veracruz State: 4 ♂, Arroyo near Playa Escondida, about 30 km NE of Catemaco (18°40'46''N, 95°09'16''W, 35 m), 28 viii 1988, R.W. Garrison leg. [RWG]; 1 ♂, same data but temporal pond, 17–24 vii 1981, R. Lopez leg. [RWG]; 4 ♂, Arroyo on road to Jicacal Beach and pond near Rancho La Esperanza, 28 viii 1988, S.W. Dunkle leg. [FSCA]; 2 ♂, same data but [RWG]; 1 ♂, same data but M.J. Westfall leg. [FSCA].

Justification of synonymy of Nephepeltia chalconota Ris, 1919 . Nephepeltia flavifrons was briefly described by Karsch (1889) based on one male from Paraguay, without accompanying illustrations. The holotype is currently deposited in ZMHB, and Mélanie Turiault kindly photographed it at my request. The photographs agree with numerous additional specimens deposited in several collections (see specimens examined), and with its original description, except that they show that there are two cells under the subtriangle in the left Fw and not one in both wings ( Fig. 4 ). Ris (1919) described N. chalconota based on a series of five males and one female from Guatemala, also without accompanying illustrations. Because his specimens presented two cells under the Fw subtriangle, he distinguished his species only from N. phryne , the only other species of Nephepeltia described at that time thought to share that character state. Direct examination of two of the male syntypes of N. chalconota deposited at the UMMZ and one deposited at FSCA ( Figs. 2 d; 7i; 9c; 10d; 13f), and of photographs of an additional syntype at UMMZ kindly sent to me by Mark O’Brien, revealed that they do not differ in any morphological character from the holotype of N. flavifrons , except for the relative height of the thoracic ventral tubercle (lower in the holotype of N. flavifrons ).

In his key to Central American dragonflies, Förster (2001) included both N. chalconota and N. flavifrons and separated them by the presence of two cells between Fw subtriangle and posterior edge of wing and male ventral thoracic tubercle at least as high as wide, versus one cell between Fw subtriangle and posterior edge of wing and male ventral thoracic tubercle wider than high, respectively (based on S.W. Dunkle in litt.). Most of the additional specimens I examined from South America have only one cell under the subtriangle in both wings (96% of 251 specimens), while about two thirds of the specimens from Costa Rica to Mexico have two cells in both wings (69% of 27 specimens). However, I found specimens collected at the same locality and dates which have one and two cells under the Fw subtriangle, in some cases with both states on right and left wings of the same individual as is the case of the holotype of N. flavifrons , evidence that this character is variable, as it also is in N. berlai . For example, of seven males I examined from near Rancho La Esperanza in Mexico, six have two cells in both wings and one has two cells in one wing and one in the other; of four specimens from Río Temash in Belize, one male and one female have one cell in both wings, and the other male and female have two cells in both wings; of 85 specimens from Fundación in Colombia, three have one wing with two cells; of 29 specimens from Río Frío in Colombia, two have one wing with two cells; of nine specimens from Río Frío in Colombia, two have one wing with two cells; of five males from Aguas Negras in Peru, three have both wings with one cell, one has one wing with two cells, and one has two cells in one wing and one in the other.

Even though the thoracic ventral tubercle from most specimens I examined from Mexico, Guatemala, and Belize is relatively more prominent than that of most specimens from Costa Rica and South America, in which it is usually relatively lower and more rounded, there is no clear-cut difference between two groups, with the height of the tubercle varying along the range regardless of locality and number of cells under the subtriangle. Some specimens from the same locality and date both in North and South America show both lower rounded tubercles and higher more prominent tubercles, irrespective of the number of cells between subtriangle and hind margin of wing ( Fig. 7 ), thus displaying a similar degree of variability in the length and shape of the tubercle to that observed in N. phryne ( Fig. 8 ). Since the specimens do not differ in any other character and all share the same hind tibia armature and morphology of genital lobe, vesica spermalis, and caudal appendages, which are diagnostic for other species in this genus, I consider differences in frequency of the number of cells under the subtriangle and relative height of the ventral thoracic tubercle observed between South and North American populations to loosely reflect geographical differences of variable characters within the same species that do not justify recognizing two separate species, rendering N. chalconota Ris, 1919 a junior subjective synonym of N. flavifrons ( Karsch, 1889) .

Diagnosis. Males of Nephepeltia flavifrons can be distinguished from all congeners by the spurs of hind tibiae short and more closely set at basal 0.50–0.75 ( Figs. 2 c–d; spurs of about uniform placement and length or closely set and peg-like at medial 0.33 in other species, Figs. 2 a–b, e, g), and by the distal segment of vesica spermalis with short rectangular latero-ectal processes and vestigial medio-ectal process ( Fig. 11 c; latero-ectal processes triangular, oval, or spine-shaped and medio-ectal process vestigial or clearly longer than wide in other species, Figs. 11 a–b, d–f). They share venter of thorax with a tubercle about as long as wide or shorter with N. aequisetis , N. berlai , and N. flavipennis ( Figs. 6 a–c; 7; smoothly convex or with a spine about as long as twice its width in N. leonardina and N. phryne respectively, Figs. 6 d; 8). Females can be recognized from those of N. leonardina by the radial planate consisting of three cells and the base of Fw discoidal planate with one cell (as in Fig. 3 c–d; four or five and two respectively in N. leonardina , Fig. 3 f). The number of cells between Fw subtriangle and hind margin of wing is variable, and females are indistinguishable morphologically from those of N. aequisetis , N. berlai , and N. phryne (females of N. flavipennis unknown).

Distribution. Nephepeltia flavifrons is found from S Mexico south to NE Argentina and S Brazil ( Fig. 15 ), representing the most widespread species of the genus together with N. phryne . Adults ( Figs. 14 a–d) are commonly seen flying at vegetated ponds, pools, marshes, and ditches.

Figures

FIGURE 2. Male hind tibiae, lateral view of inner surface.

FIGURE 3. Male wings. d. f.: base of discoidal field; r. p.: radial planate; sub.: cells between subtriangle and hind margin of wing.

FIGURE 4. Nephepeltia flavifrons, holotype male, dorsal view with details of Fw subtriangles.

FIGURE 6. Male thoracic venter, lateral view.

FIGURE 7. Male thoracic venter, lateral view, variability in N. flavifrons. [n – n]: number of cells under Fw subtriangle in left and right wings.

FIGURE 8. Male thoracic venter, lateral view, variability in N. phryne.

FIGURE 11. Male vesica spermalis, lateral (above) and ventral (below) view. l. e.: latero-ectal sclerotized distal process; m. e.: medio-ectal sclerotized distal process.

FIGURE 14. Habitus pictures of N. flavifrons and N. phryne showing variability in extension of pruinescence and pale markings on thorax and abdomen in males. (a) Male of N. flavifrons, Brazil, Rio de Janeiro, Reserva Ecologica de Guapi Açu, 24 x 2013; (b) Male and female of N. flavifrons in copula, same data but 14 x 2013; (c) Male of N. flavifrons, Costa Rica, Limón Prov., Tortuguero, 1 viii 2010; (d) Male of N. flavifrons, Brazil, Rio de Janeiro, Vecchi wetlands, 21 x 2013; (e) Male of N. phryne, Brazil, Rio de Janeiro, Reserva Ecologica Guapi Açu, 14 x 2013; (f) Male of N. phryne, Panama, Canal Area, Old Gamboa Road, Summit ponds, 26 viii 2012. (a – b, c – e), by Dennis Paulson; (c, f) by Netta Smith.

FIGURE 15. Distribution of Nephepeltia species.