Cuernavaca inexacta ( Walker, 1858 )

Lenicov, Ana María Marino de Remes, Hernádez, María Cristina, Sosa, Alejandro Joaquín & Oleiro, Marina Inés, 2020, The genus Cuernavaca (Hemiptera: Dictyopharidae), associated with Pontederiaceae in South America, with redescription and new records of C. inexacta (Walker), Papéis Avulsos de Zoologia 60, pp. 1-9 : 3-8

publication ID 10.11606/1807-0205/2020.60.11

persistent identifier

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scientific name

Cuernavaca inexacta ( Walker, 1858 )


Cuernavaca inexacta ( Walker, 1858) ( Figs. 1 View Figure 1 A‑G; 2A, B)

Dictyophora inexacta Walker, 1858: 38 , synonymized with T. herbida by Muir, 1931: 474.

Taosa paraherbida Muir, 1931: 474 , synonymized by Fennah, 1947: 9.

Taosa (Cuernavaca) paraherbida Muir, 1931 , according to Metcalf, 1946: 110.

Taosa inexacta ( Walker,1858) restored by Fennah,1947:9.

Cuernavaca inexacta ( Walker, 1858) according to Emeljanov, 2011: 1123; Song et al., 2018: 13.

Emended diagnosis: General color green, slightly tinged reddish brown band overlying submedian frontal carina and base of ocelli. Tegmina and hindwings clear hyaline with green veins; pterostigma opaque yellowish. Body elongate; head with narrow vertex, slightly longer than wide across base, projected for more than half its length in front of eyes, median carina evanescent in apical half, transverse carina anteriorly bounding well-defined trigones,separated from preocular areas by slightly sinuous carina, sharp apices distant from midline; frons long, lateral margins slightly parallel in basal half, barely expanded in apical third; rostrum extending past basal third of hind femora in repose.

Male: Anal segment ventrally covering apex of gonostyles; gonostyles slender, longer than wide, narrowing at base in ventral view, upper margin slightly angular on basal half and gradually narrowing to the apex, with a distinctive long straight process dorso cephalad directed, apically obtuse, and a small ventro caudad curved hook-like process near sub-middle on the outer edge, in lateral view; aedeagus ending in a pair of long sinuous ventro-apical curved spinose processes dorsocaudally directed; phallobase tubular with a pair of slender and apically lanceolated endosomal processes emerging abruptly cephalad in their apical fourth. Segment X distinctly elongate, anal style far beyond apical ventral margin of segment X.

Female: Anal segment relatively short, just above the ovipositor; ovipositor short and strong; gonapophyses VIII (first valvulae) strong, slightly curving upward to apex, anterior connective lamina with about 24 uneven teeth decreasing in size toward base arranged in double row running subparallel in the dorsal margin. Gonoplacs (third valvulae) slightly elongated, the ventral lobe with a narrow densely hairy area along lower margin and a tuft of short stout hairs at apex. Segment X oval to elongate; anal style far beyond apical ventral margin of segment X. Female ectodermal genital ducts ditrysian, surrounding area of copula poris distinguishably colored.


Body ( Fig. 1 View Figure 1 A‑C): Ground color uniformly green, with slightly tinged reddish brown bands overline submedian frontal carina and around ocelli.Tegmina and wings clear hyaline with green veins; pterostigma opaque yellowish.

Head: Vertex narrow in dorsal view, little longer than broad across base (1.1:1), apical margin broadly angulate, projecting in front of eyes by more than half its length, on the same plane in relation to pronotum; lateral carinae slightly ridged in front of eyes, sub-parallel at base and gradually convergent anteriorly; basal margin rather shallowly round excavated between eyes, basal width nearly twice the diameter of eyes, median carina incomplete, only distinct in base; transverse carina dorsally bounding the well-defined subtriangular trigones, separated from preocular areas by a slightly sinuous carina, sharp apices distant from the mid line. Frons relatively short, in the middle a little longer than wide at maximum width (1.3:1), linear, with lateral margins slightly parallel in the basal half, barely expanded in the apical third, carinae distinct, without carina between apex of lateral and intermediate carinae, the median complete and very feeble, the intermediates subparallel, nearly straight, weakly ridged, evanescent next to frontoclypeal suture, laterals strongly ridged; apical margin deeply excavated and inverted V-shaped.Clypeus longer than wide at base (2.1:1), surpassing apex of fore coxae, laterals and median carina distinctly elevated, median extended to apex ( Fig. 1C View Figure 1 ). Labrum free, elongate; rostrum relatively long, surpassing basal third of hind femora, basal segment longer than apical (1.3:1) reaching apices of hind coxae. Compound eyes long oval. Ocelli present.

Thorax: Pronotum wider than head (1.3:1), shorter than vertex at midline (0.5:1); carinae distinctly ridged, median carina percurrent, intermediate carinae distinct in basal third to half, curved closely behind the eyes, with a prominent slightly curved downwards carinae on paranotal lobes not reaching rear edge; anterolateral angles rounded, anterior central margin arcuately convex, posterior margin deeply concave. Mesonotum medially more than six times the length of the pronotum, carinae distinct, extinct at the scutellum, the laterals slightly divergent, feebly incurved basally not reaching median carina; tegmina membranous extending beyond the abdomen, with ratio length to maximum width 3:1; hyperpterous in R,MP and CuA., with a second rank of postnodal closed cells after the nodal cells; pterostigma narrow three-celled. Hind tibiae with 4 lateral black-tipped spines and 8 apical black-tipped teeth; hind tarsomeres I as long as II and III together, tarsomere I and II with seven black-tipped apical teeth, respectively ( Fig. 1D View Figure 1 ).

Male genitalia ( Fig. 1E View Figure 1 ): Pygofer in lateral view distinctly longer ventrally than dorsally (4:1), more than twice ventral width; caudal margins rounded, slightly expanded ventrally; dorsal margin straightly excavated to accommodate segment X; ventral margin narrowly truncate. Gonostyles relatively large, slender; in lateral view with base narrow, strongly expanded in the middle and gradually narrowed towards rounded blunt apex; upper margin slightly angular at basal half, with a long lightly compressed straight hook-like process forward directed, apically obtuse; outer upper edge with a small ventro caudad curved spine near sub-middle. Aedeagus with phalloteca tubular, slightly sclerotized at base and laterally, the remainder membranous with pair of inflatable membranous lobes, ending in slender ventro-apical spinose processes dorso-caudally directed and downward recurved at apex; phallobase tubular with a pair of slender and apically lanceolate endosomal processes extending posteriorly below theca and folding abruptly cephalad in their apical fourth. In lateral view, ventro-apical spinose processes of aedeagus reaching apex of gonostyles and ventrocaudad angle of segment X at rest. Connective slender, elongate and slightly sclerotized, extending obliqually from the base of aedeagus to the base of by: Segment X as long as the length of style in lateral view; in dorsal aspect distinctly elongate, distal portion deeply medially emarginated, with ratio of length to width 2.5:1; in lateral view dorsal and ventral margins straight, the latter slightly longer in lateral view (1.1:1); apex obliquely truncate; anal style (Epr and Ppr) relatively short extended more than the half of its length beyond apical ventral margin.

Female ( Fig. 1B View Figure 1 ): Similar to male in body morphology and coloration.

Genitalia ( Figs. 1F, G View Figure 1 ): Short and strong ovipositor; anal segment relatively short, reaching apex of ovipositor at rest. Gonocoxa VIII (1 st valvifer) triangular, vertically placed, gonapophyses VIII (1 st valvulae) subelliptical in lateral view, dorsal border smoothly and slightly curved, ventral border slightly convex to dorsum, both dorsal and ventral margins narrowing sharply in the apical half, slightly curving upward to apex; anterior connective lamina with subparallel double rows of unequal teeth on dorsal margin, each one consisting of about 10 to 11 sharp teeth, almost same-sized, lightly descending in size toward base. Gonapophyses IX, elongate and narrow. Gonoplacs (3 rd valvulae) with both lobes subequal in length, as long as anal segment, subparallel sided, ventral lobe rounded at apex, with lower margin convex strongly curving upward to apex, with a densely hairy area along lower edge in apical half and a tuft of rigid hairs at tip; dorsal lobe slender truncate at apex, strongly reflected inward. Segment X longish ovate, distal portion deeply medially emarginated, with ratio of length to width 2:1; in lateral view dorsal and ventral margins straight, the latter slightly longer in lateral view (1.1:1); apex slightly truncate; anal style (Epr and Ppr) relatively short extended more than half of its length beyond apical ventral margin. Ectodermal genital ducts ditrysian; copulaporis area darkish colored, corresponding to the inner esclerotized vaginal plates, visible through translucent skin, more evident in cleared specimens.

Measurements: L: male, 11.1 (11-11.3), female, 12.1 (11.6-12.6); B.L: male, 7.75 (7.7-7.8), female, 9.5 (9.6-10); W.L.: male, 9.7 (9.5-10), female, 10.2 (10-10.3).

Taxonomic remarks

Cuernavaca inexacta differs from its congeners in its coloration pattern and head traits. The most distinctive features of the male genitalia are the combination of the angular profile of gonostyle, the longest and slender processes in the aedeagal complex and the endosomal processes. Whereas in female, the distinctive feature is the numerous teeth arranged in two subequal and sub-parallel rows on gonapophyses VIII.

Cuernavaca inexacta can be easily distinguished from the sympatric C. longula , known to occur in the same habitats and feed on the resources in several areas of South America, by the smaller size, the reddish brown coloration on the base of frons and around base of ocelli, the longer vertex, the frons not so strongly expanded near apex, the gonostyles wider at middle in lateral view with straight and forward directed hook-like process, and the more slender, longer and different shaped adeagal and endosomal processes in male, and the female by the distinctly shorter anal segment and ovipositor, the greater number of teeth lying in two rows on the anterior connective lamina of gonapophyses VIII and the darkish coloration on the copulaporis area.

Female genitalia of Cuernavacainexacta is redescribed and redefined based on examination of the Holotype of Dictyophara inexacta Walker ( Figs. 2A, B View Figure 2 ), the paratype female of Taosa paraherbida Muir (specimen designed as residual type by Fennah, 1947: 9) ( Figs. 2C, D View Figure 2 ), and 1 female of Taosa inexacta (Walker) from Guyarmare, Trinidad, Ghauri det. ( Figs. 2E, F View Figure 2 ) (all in BMNH). We observed that the anterior connective lamina of the gonapophyses VIII have two subequal and subparallel rows of teeth dorsally, each with 10 to 11 sharp denticle. These features contrast with those erroneously described and illustrated by Fennah (1945) for T. paraherbida Muir as follow: “First valvulae narrow, sinuate, with a row of eight stout, short, triangular teeth on dorsal margin, with a point at apex”. In the remaining features specimens match the original description in every detail; probably the double row of denticles might not have been observed by the author from a side view (p. 464, pl. 12, fig. 306).

Examination of one male specimen of Taosa paraherbida bearing Muir’s identifcation label from Colombia (BMNH) ( Figs. 2G, H View Figure 2 ) as well as those field collected specimens from Peru and Argentina, allowed us to provide a more complete description. We also noticed very little intraspecific variation in color and morphology among the adults in both sexes.


According to previously published records and voucher specimens from the BMNH, MNHN and MLPA, the three species of Cuernavaca are distributed over Central and South America, reaching north and central Argentina in the southernmost distribution ( Fig. 3 View Figure 3 ).

Cuernavaca inexacta : Colombia ( Walker, 1858); Guyana ( Muir, 1931) and Trinidad ( Fennah, 1945). New records: Panamá (Canal Zone, Gamboa), Venezuela (Etat de Guarico), Peru (Dto Loreto) and Argentina: Formosa, Chaco.

Cuernavaca longula : Argentina Northeast ( Formosa y Chaco) ( Remes Lenicov & Hernández, 2010; Hernández et al., 2011b; Remes Lenicov et al., 2012) Brazil West-Central (Mato Grosso do Sul and Perú ( Remes Lenicov & Hernández, 2010; Remes Lenicov et al., 2012).

Cuernavaca herbida: Trinidad-Tobago ( Walker, 1851) ; Costa Rica (Matina), Ecuador (Tena) and Brazil Southeast

(São Paulo, Dumont, Piracicaba) ( Muir, 1931); Trinidad (St. Agustine) and Venezuela (Ocumare de la Costa) ( Fennah, 1945); Mexico: Morelos ( Cuernavaca ), Nuevo Leon (Monterrey), Chihuahua; Ecuador: Napo Pastaza; Guatemala; British Honduras; Nicaragua; Costa Rica; Panamá; Central America; Colombia; Venezuela; Brazil; Paraguay; Argentina and South America ( Metcalf, 1946).

Material examined

Cuernavaca inexacta (Walker) : Holotype female of Dictyophara inexacta Walker, 1858 , Colombia; Paratype female of Taosa paraherbida Muir, 1931 , Blairmont B.G. , September 1923, F.X.Williams col.; 1 male, Taosa paraherbida Muir , Colombia, Lake Sapatozoa, Chiriguana District , VIII-IX-1924, C. Allen col. (Muir’s handwriting), Muir det.; 1 female of Taosa inexacta (Walker) , Guyarmare, Trinidad, on P. crassipes , Dic. 1964, Ghauri det.; 1 male, Guyarmare, Trinidad, on P. crassipes , 1967; 1 male and 1 female, Canal Zone, Gamboa, Panama, Chagres R. at night, on P. ro - tundifolia, 30-VI-1974, C and L O’Brien and Marshall col.; 3 male, 1 female and 1nymph V,unidentified,Guyarmare, Trinidad, on P. crassipes , Dic. 1967. ( BMNH); 1 male, Venezuela, Etat de Guarico. La Cruz-Ruviera; M. Grisol, 1925; December ; MNHN ( EH) 18726; 1 male, Venezuela, Etat de Guarico. Camagua ; M. Grisol, 1923; MNHN ( EH) 18727; 1 male and 1 female, Colombia; 27-XI-1996; G. Lecourt; MNHN ( EH) 18728-29, ( MNHN). PERU: 6 males and 10 females, Montoya, 04°31′00″S, 73°32′23″W, on P. crassipes , 03-05-1999, Cordo col.; GoogleMaps 3 males and 5 females, Iquitos, Montoya Cocha , 04°31′00″S, 73°32′23″W, on P. crassipes , 03-05-1999, Cordo col.; GoogleMaps 3 males and 3 females, Perú, on P. cordata , 15-11-2001, Cordo col.; 1 female, Tacunare/Nacita, 12°16′22.33″S, 76°53′26.36″W, on P. crassipes , 15-11-2001, Cordo col.; GoogleMaps 1 female and 8 nymphs, Morana, Morana ; 04-11-2001, Cordo col.; 2 males, Iquitos, Carmen Cocha , 03°44′37.22″S, 73°15′05.88″W, on P. crassipes , 02-05-1999, Cordo col.; GoogleMaps 5 males and 20 females, Montoya Cocha, 04°31′00″S, 73°32′23″W, on P. crassipes , 01-05-1999, Cordo col.; GoogleMaps 1 female, Montoya Cocha, 04°31′00″S, 73°32′23″W, on P. rotundifolia ; 03-05-1999, Cordo col.; GoogleMaps 3 males and 3 females, Iquitos, Tipashira Cocha , 04°18′45″S, 73°19′32″W, on P. crassipes , 30-04-1999, Cordo col.; GoogleMaps 5 males, Iquitos, Peruru Cocha , 04°16′51″S, 73°17′03″W, on P. crassipes , 29-04-1999, Cordo col.; GoogleMaps 1 male and 1 female, Yanayacu River, 04°19′29″S, 73°18′11″W, on P. rotundifolia , 30-04-1999, Cordo col.; GoogleMaps 7 males and 4 females, Yarapa Yarapa, 04°28′56′S, 73°23′39″W, on P. crassipes , 13-11-2001, Cordo col.; 1 male, Tacunare/Nacita, on P.crassipes , 14-11-2001, Cordo col., 4 males and 4 females, Norte Caño,on P.crassipes , 14-11-2001, Cordo col.; 1male, Yarapa, Yarapa , 04°28′56″S, 73°23′39″W, on P. crassipes , 14-11-2001; Cordo col. GoogleMaps ARGENTINA: 1 female, Formosa province, Subteniente Perin, 25°46′67″S, 59°88′33″W, on P. cordata , 15-12-1998, Sosa col.; GoogleMaps 1 female, Herradura, 26°30′59″S, 58°16′54″W, on P. crassipes , 30-III-2016, Oleiro , Guala col., GoogleMaps 4 males and 6 females, Palo Santo, 25°33′42.0″S, 59°19′10.6″W, on P. subovata , 15-04-2005, Sacco , col.; GoogleMaps 1 male, La Florencia, 26°29′06.20″S, 58°18′16.65″W, on P. crassipes , 30-03-2016, Oleiro , Guala col.; GoogleMaps 1 female, Irupe, on P. crassipes , 05-05-2016, Oleiro , Jastrzebski col. ( MLPA).

Cuernavaca longula (Remes Lenicov) : Holotype male of Taosa (C.) longula , Argentina, Formosa, Herradura, on P. crassipes , 11-II-2004, Hernandez & Sacco , col.; Paratipo female, same data as Holotype; Paratipo male, Peru, Iquitos, Pururu Cocha , on P. crassipes , 29-IV-1999, Cordo col. ( MLPA).

Cuernavaca herbida (Walker) : Holotype female of Dichoptera herbida Walker , Venezuela; 1 male of Taosa herbida (Walker) , Costa Rica, Matina, 1912, Kirkaldy col., F. Muir, det.; 1 male of Taosa herbida (Walker) , Trinidad, Uvarov, det.; 2 females, Atoyac, Vera Cruz , May H.H.S.; 3 males and 1 female, Teapa , Tabasco, Feb, H.H.S.; 1 female, Durango, Mexico; 1 female, Jalisco, Mexico; 1 female, Belize, Cayo; 1 male, San Gerónimo , Guatemala; 2 females, Matachin , Panamá; 1 male, Trinidad; 1 female, Dumont , San Pablo , Brasil; 11-VII-1923; G.L. R. Hancock ( BMNH). 1 male, El Salvador, La libertad, 04-XI-1958, Duret col., 788/93, MNHN ( EH) 18724; 1 female, Ecuador, Quito, R. Benoist 1930. MNHN ( EH) 18723. ( MNHN).


United Kingdom, London, The Natural History Museum [formerly British Museum (Natural History)]


France, Paris, Museum National d'Histoire Naturelle


Argentina, La Plata, Universidad Nacional de La Plata, Museo de la Plata


Departamento de Geologia, Universidad de Chile


Royal British Columbia Museum - Herbarium


Museum National d'Histoire Naturelle














Cuernavaca inexacta ( Walker, 1858 )

Lenicov, Ana María Marino de Remes, Hernádez, María Cristina, Sosa, Alejandro Joaquín & Oleiro, Marina Inés 2020

Cuernavaca inexacta ( Walker, 1858 )

Song, Z. - S. & Bartlett, C. R. & O'Brien, L. B. & Liang, A. - P. & Bourgoin, T. 2018: 13
Emeljanov, A. F. 2011: 1123

Taosa inexacta ( Walker,1858 )

Fennah, R. G. 1947: 9

Taosa (Cuernavaca) paraherbida Muir, 1931

Metcalf, Z. P. 1946: 110

Taosa paraherbida Muir, 1931: 474

Fennah, R. G. 1947: 9
Muir, F. 1931: 474

Dictyophora inexacta

Muir, F. 1931: 474
Walker, F. 1858: 38