Cephisus siccifolia (Walker, 1851)

Aphrophora siccifolia Walker, 1851: 698 .

Cephisus siccifolius [sic] Stål, 1866: 384.

Sphodroscarta siccifolia Stål, 1869: 18 .

Cephisus siccifolia Hamilton, 2012: 51–52, figs 1B, 2B, 13A–C, 21.

Material examined

Cephisus siccifolia: 15♂, 15♀ “ Brasil, PR, Centro Politécnico \ UFPR, Árvores prox. ro RU, 09.X.2023, A.C. Domahovski, \ A. Paladini, L. Alasmar, leg.”; 1♂ “ Brasil, Rondônia. Flona \ Jamari, 3/ 5.ix.2012 (light) \ 9°08'75"S 63°00'70"W \ 110m / R. R. Cavichioli”; 1♂ “ Brasil, Acre, Bujari, FES do \ antimary, 9°20'01"S - \ 68°19'17"W, Arm. Luminosa \ A.A Agudelo, F.F Xavier, D. M. Mendes, J. A. Rafael leg”; 1♂ “ Ipixuna - AM, Rio \ Gregório; Com. Lago \ Grande, Seringal \ Recreio \ 7°10'6"S 70°49'6"W ” “ 145m - light trap \ 18-23V.2011 \ Cavichioli, Gonçalves & \ Takiya”; 1♂ “ Encruzilhada - Bahia \ Brasil, 580m, XI/74 \ M. Alvarenga leg.”; 1♂ “ Brasil, MT, Chapada dos \ Guimarães, Aldeia Velha \ 15°26'14"S 55°45'30"W \ 698m 1-31.X.2016 Luz \ P. R. O. A. Correa ”; 1♂ “ Sinop Mato Grosso \ Brasil X-1975 \ M. Alvarenga leg.”; 1♂ “ Brasil, MS, Dourados, \ Reserva Florestal Faz. \ Coqueiro 22°12'34"S \ 54°54'46"W 19.II.2009 A. \ Paladini, D. R. Parizotto, P. C. \ Grossi leg. Luz”; 1♂ “ Brasil, Santa Catarina, \ Chapecó, Monte Belo \ 01-30.iv.2013 \ (Malaise) M. Savaris, & \ S. Lampert leg”; 1♂ “ Brasil, Paraná, Tibagi, \ P. E. do Guartelá, 1000m \ 24°33'41"S 50°15'26"W \ 21-24.XI.2016 Sweep \ A. C. Domahovski leg.”.

Adult redescription

Coloration (Fig. 2). Head dark brown with paler whitish area near ocelli; frons light brown, blackish apically; clypeus, lorum, genal lobe and last rostral segment blackish. Pronotum dark brown with whitish areas on the anterior and posterior thirds, above the pronotal rugosity. Scutellum dark brown, whitish apically. Tegmina blackish brown with paler areas forming three crescent-shaped oblique bands on the basal and median thirds, in some specimens the bands are less apparent. Thorax and abdomen light brown, legs light brown with femora and tibia blackish. Some specimens of the same brood show a light-colored head, frons, pronotum, thorax and abdomen.

Structure (Fig. 2). Head as wide as anterior margin of pronotum, humeral angle of pronotum surpassing the length of eyes, tylus triangular with fine pubescence and without median carina, vertex rectangular with ocelli equidistant to each other (approximately three diameters of one ocellus) and equidistant to the eye, compound eyes disposed obliquely; frons inflated, strongly projected backwards, median carina strongly marked in the median and posterior third, lateral grooves strongly marked, rostrum surpassing the metathrocanter; pronotum hexagonal, anterior margin convex, anterolateral margin straight, posterolateral margin with a slight concavity medially, posterior margin with a strong median concavity, anterior third of pronotum with a median depression forming a slightly marked carina, median and posterior portion of pronotum elevated, punctuated with strongly marked transverse rugae; scutellum with a depressed area at middle with transverse striae slightly marked. Metafemur with a conspicuous apical spine, metatibia with two lateral spines, the basal one smaller than the apical, apex of tibia with two rows of strong spines apically, basitarsus with one row of 7–8 apical spines.

Male genitalia (Fig. 3). Pygofer with a rounded, inconspicuous process between the anal tube and subgenital plates, anal tube surpassing subgenital plates (Fig. 3A), subgenital plates with a truncate apex (Fig. 3B), tip of the dorsal margin turned inward forming an apical, minute triangular process. Style (Fig. 3C, 3D) with dorsal margin convex, apical process turned upward with a minute apical teeth slightly sclerotized and turned inward, another small inner process turned inward. Aedeagus (Fig. 3E–G) robust with a curved shaft armed with a pair of lateroapical acute processes; each lateral apical process curved toward ventral side of shaft, and laterally flattened, and a pair of ventroapical processes directed toward shaft base.

Female genitalia (Fig. 4). First valvifer (Fig. 4A) taller than long, covered with long setae; first valvula of ovipositor (Fig. 4A) long with rounded apex, basal process undeveloped directed downward, dorsal portion of blade, in lateral view, with striate sculptures (Fig. 4B); second valvula (Fig. 4B) long with rounded apex dorsal margin undulate without denticulation, blade in lateral view with striate, slightly marked sculpture (Fig. 4D); second valvifer (Fig. 4E) longer than wide, setae present along ventral portion and forming a group on basal third and; third valvula (Fig. 4E) short and wide, covered apical and ventrally by long setae.

Remarks. According to Hamilton (2012), this species is widely distributed in lowland regions of South America, from Uruguay to Panama. This species is very similar to C. diminutus and C. magnificus both in terms of external morphology and male genitalia, and both species are distributed in Brazil. It also presents morphological similarity with C. variolosus, which is known to occur in North and Central America.

Immature description

First instar (Figs 5, 10A, 12A, 13A). Body length 1.38 (1.05–1.7); head width 0.51 (0.475 –0.675); head length 0.42 (0.32–0.52); pronotum width 0.47 (0.4–0.5); pronotum length 0.11 (0.075 –0.125); frons width 0.353 (0.3–0.35); frons length 0.24 (0.2–0.47). Measurements in mm.

General color yellowish, dorsum (Fig. 5A) with a conspicuous longitudinal white line from head apex to mesonotum, abdomen reddish, mostly in ventral portion. Head broadly rounded, slightly wider than long, without ocelli. Antennae (Fig. 12A) with indistinct supra-antennal ledge, flagellum with 7 flagellomeres, first and second fused; apex rounded; scape and pedicel combined length surpassing flagellum length. Frons, in ventral view (Fig. 5B), with inconspicuous muscular impressions; in lateral view (Fig. 5C), strongly inflated. Rostrum with 3 segments, surpassing base of coxa III. Pronotum and mesonotum rectangular, much wider than long, metanotum restricted to lateral margins. Pronotum length half length of mesonotum plus metanotum. Wing pads indistinct. Hind tibia (Fig. 13A) without any visible setae, apical crown of spines slightly visible; two visible tarsomeres. Abdomen, in ventral view (Fig. 5B), with a slight median groove, distal portion undifferentiated.

Second instar (Figs 6, 10B, 12B, 13B). Body length 4.32 (3.6–4.85); head width 1.11 (0.9–1.25); head length 0.91 (0.75–1.25); pronotum width 1.18 (1.4–1.05); pronotum length 0.42 (0.3–0.45); frons width 0.85 (0.7–0.95); frons length 0.53 (0.4–0.7). Measurements in mm.

General color yellowish to brownish, dorsum (Fig. 6A) with a conspicuous longitudinal white line from head apex to abdomen apex, broadened in mesonotum, and some yellow to pale brown spots laterally of median line. Abdomen pale, slightly reddish ventrally in median portion. Head similar to that of the anterior instar. Antennae (Fig. 12B) with supra-antennal ledge starting to differentiate, flagellum similar to that of first instar, with length surpassing scape and pedicel combined length; apex subacute. Frons, in ventral view (Fig. 6B), with conspicuous muscular impressions; in lateral view (Fig. 6C), similar to that of the anterior instar, but more convex. Rostrum with 3 segments, reaching base of coxa III. Pronotum and mesonotum rectangular, wider than long, metanotum broadened, but restricted to lateral margins. Pronotum length almost as long as mesonotum plus metanotum. Wing pads indistinct. Hind tibia (Fig. 13B) with some visible setae, apical crown of spines visible similar to that of the anterior instar; two visible tarsomeres, first with few apical setae. Abdomen, in ventral view (Fig. 6B), with a conspicuous median groove, abdominal tergites extended downwards as flaps; distal portion undifferentiated.

Third instar (Figs 7, 10C, 12C, 13C). Body length 5.83 (4.5–7.2); head width 1.87 (1.7–2.1); head length 1.37 (1.2–1.65); pronotum width 2.04 (1.8–2.45); pronotum length 0.75 (0.6–0.85); frons width 1.36 (1.2–1.55); frons length 0.87 (0.7–1.2). Measurements in mm.

General color similar to that of preceding instar, with head and thorax darker with paler markings dorsally, longitudinal white line dissipating in thorax, conspicuous in abdomen. Head similar to that of the anterior instar, with ocelli spots slightly distinct, transocular width about 1,3x of vertex width. Antennae (Figs 5, 12C) with supra-antennal ledge and flagellum similar to that of anterior instar, with a more distinct sulcus between first and second flagellomeres, length of flagellum two times the length of scape and pedicel combined. Frons, in lateral view (Fig. 7C), similar to that of the anterior instar. Rostrum with 3 segments, surpassing base of coxa II. Pronotum similar to that of anterior instar, lateral margins of mesonotum and metanotum starting to differentiate in wing pads; length two-thirds as long as mesonotum plus metanotum. Hind tibia (Fig. 13C) with more visible setae. Abdomen with morphology similar to that of anterior instar, in ventral view (Fig. 7B) with the abdominal tergite flaps quadrangular and darkened, distal portion of abdomen undifferentiated.

Fourth instar (Figs 8, 10D, 12D, 13D). Body length 12.21 (9.56–16.34); head width 3.22 (2.81–3.75); head length 1.80 (1.5–3.12); pronotum width 4.06 (3.37–4.68); pronotum length 1.65 (1.25–2.00); frons width 1.89 (1.62–2.12); frons length 0.91 (0.62–1.37). Measurements in mm.

General color (Fig. 8) more yellowish to that of anterior instar, with head and thorax more marked with brown maculae, without a longitudinal white line in body. Head similar to that of the anterior instar, with ocelli distinct, transocular width about 1,75x of vertex width. Antennae (Fig. 12D) with distinct supra-antennal ledge, flagellum with 7 well delimited first and second flagellomeres completely separated; flagellum thin, more than 4x longer than scape and pedicel combined. Frons and rostrum similar to that of the anterior instar. Pronotum posterior margin sinuate, mesonotum median portion produced posterad, forming the scutellum. Wing pads well developed, much longer than pronotum, in dorsal view, reaching half of third abdominal segment. Hind tibia (Fig. 13D) with more visible setae and in greater quantity, lateroapical spine inconspicuous, starting to differentiate; tarsomeres with plantar spines inconspicuous, starting to differentiate. Abdomen similar to that of anterior instar, with abdominal tergite flaps rounded and distal portion differentiated in male or female genital plates.

Fifth instar (Figs 9, 10E, 11, 12E, 13E). Body length 13.21 (10.94–14.98); head width 3.34 (2.81–3.56); head length 1.80 (1.56–2.06); pronotum width 4.06 (3.19–4.69); pronotum length 1.55 (1.40–1.75); frons width 1.99 (1.62–2.12); frons length 0.92 (0.75–1.06). Measurements in mm.

General color and head morphology similar to that of anterior instar. Antennae (Fig. 12E) with distinct supra-antennal ledge, flagellum with 8 flagellomeres. Frons and rostrum similar to that of the anterior instar. Pronotum posterior half transversely striated, with posterior margin sinuate, mesonotum median portion with well-delimited scutellum, transversely strigated. Wing pads well developed, with veins slightly distinct. Hind tibia (Fig. 13E) with lateral spines distinctly visible, apex with three rows of spines, plantar surface of tarsomeres with inverted triangular row of setae. Abdomen similar to that of anterior instar, distal portion differentiated in male or female parts. Male plates form a single broad and short structure, divided only in apical portion, forming a V-shaped notch (Fig. 11A). Female plates are a pair of two elongated triangular structures, formed by sternites VIII and IX (Fig. 11B).

Instar discrimination. Size variation for each instar and morphometric parameter is given in Table 1. Considering head length, pronotal length and pronotal width, there is a steep growth among first and second instar, with mean growth ratio of 3.04, and a lesser growth rate among second, third and fourth instar, and size stabilizes among fourth and fifth instar (Table 1).

Four discrete size classes could be individuated by head and pronotal width (Fig. 14). First, second and third instars can be discriminated by these morphometric parameters, but fourth and fifth instars overlap (Fig. 14; Tables S1 and S2). The other morphometric parameters, i.e. length of head, frons, pronotum and thorax, and width of frons, do not allow discriminating size classes (Fig. S1, Tables S1 and S2).

On the other hand, it is possible to identify nymphal instars through qualitative evaluation of morphological changes. Muscular impressions can be observed from the second instar onwards (Figs 5–10). Ocellar spots, the differentiation of wing pads, and the quadrangular projections of urosternites, are visible from the third instar, and can be used to distinguish it from the second instar (Figs 6, 7, 10). The differentiation between the third and fourth instars can be made by the former presenting distinctly more developed wing pads, surpassing the tergun II, by the triangular shape of the posterior margin of the pronotum, indicating the starting of scutellar development (Figs 7, 8, 10). Fourth and fifth instars can be reliably separated by seven antennal flagellomeres in the fourth instar, whereas the fifth presents eight (Fig. 12D, E). Other characteristics that can aid in discriminating fourth and fifth instars are the presence of, on the fifth instar, transverse grooves on the posterior margin of the pronotum (Fig. 9A), the distinct presence of the scutellum with rugosities, the beginning of vein formation (Fig. 9C) in both the forewing and hindwing, development of lateral spines on the tibiae and the presence of spines on tarsomere’s plantar surfaces (Fig. 13E).

SD, standard deviation; IQR, interquartile range.

Comparative morphology of male genitalia

To highlight some of the intraspecific variations found in C. siccifolia, in addition to the specimens from spittle masses collected in Curitiba, Paraná (Figs 1–4, 15), we studied nine specimens from different Brazilian locations and geographic regions (Fig. 20): North (states of Rondônia, Acre, and Amazonas), Northeast (Bahia), Midwest (Mato Grosso and Mato Grosso do Sul), and South (Santa Catarina and Paraná) (Figs 16A–I, 17). The range of length of males was between 11.2 mm (Fig. 16F, G, from Mato Grosso) to 13.5 mm (Fig. 16C, from Amazonas) while the size range for males from Curitiba (Paraná) was between 11.7 to 13.2 mm.

We also found intraspecific variation of the male genitalia, specifically the aedeagus and style (Figs 3, 17). On the aedeagus, the lateroapical processes can be wide (Fig. 17S, W) or narrow (Fig. 17M, Z); strongly (Fig. 17M) or moderately curved (Fig. 17D), or almost straight (Fig. 17S); more strongly curved anteriorly in lateral view (Fig. 17B) or less curved (Fig 17K); the ventroapical processes can be shorter (Fig. 17K, T), moderately long (Fig. 17E, Q) or longer (Fig. 17A). The style apex varies mainly in the shape of the apical processes, with the dorsal process being shorter and wider (Fig. 17C), or more elongated (Fig. 17 Ab), or more triangular with the lateral margins almost straight (Fig. 17U, Y), or more curved anteriorly (Fig. 17O, Ab); the small process of posterior margin varies in size, being smaller (Fig. 17O, R, Ab) or larger (Fig. 17L, U).

We selected nine other specimens, from different geographic regions (from Mexico to southern Brazil), that may be separated into three morphological groups (putative groups 2, 3 and 4) showing similarities with some of the currently recognized species of Cephisus (Figs 16J–R, 18). The first group (Fig. 16J–L, 18A–I) represents the smallest male specimens studied (9.1 to 10.7 mm), all from the State of Bahia (Northeastern Brazil), which we determined as Cephisus aff. variolosus or brevipennis, due to the shape of the style and the short and strongly tapered lateroapical processes of aedeagus. However, these three specimens have the lateroapical processes shorter than C. variolosus or C. brevipennis, and smaller body size. As the range of intraspecific variation is unknown within these species it is doubtful to classify them as belonging to a single species or even as a new species.

The second group, comprising specimens from Rio de Janeiro (Brazil) (Figs 16M, O, 18J–L, P–R), Ecuador (Figs 16N, 18M–N), and Mexico (Figs 16P, 18S–U), is more similar to C. laticeps mainly due to the apical portion of style narrower than of its congeneric species, and the thin and curved dorsally lateroapical processes of the aedeagus. However, the specimen from Ecuador could also be interpreted as morphologically similar to C. variolosus despite being outside the distribution range of this species proposed by Hamilton (2012).

The third group is represented by one specimen from Argentina (Fig 16Q, 18W–Y) and one from Minas Gerais state, Brazil (Fig. 16R, 18Z –Ab). These specimens are more similar to C. jacobii (style not illustrated by Hamilton 2012) by the lateroapical and ventroapical processes of the aedeagus with blunt or truncated apices and bearing many small denticles or serrations. However, they also share similarities with C. diminutus in having a similar stylus with posterior process prominent, produced posterad. Moreover, they differ from C. jacobii due to the fewer serrations present on the ventrolateral processes and the current distribution range documented for this species (Peru, British Guiana); and from C. diminutus, which lacks serrations on aedeagal processes.

Additional specimens examined

Cephisus aff. variolosus / brevipennis: 1♂ “Encruzilhada - Bahia \ Brasil X-1975 \ M. Alvarenga leg.”; 2♂ “ Brasil, Bahia, 10 km a \ NE de Encruzilhada, \ 15.483° S 40.824° W, \ 830m, 15.xii.2012 \ G. Melo & P. Grossi \ Arm luminosa”.

Cephisus aff laticeps: 2♂ “ Brasil, RJ, Nova Friburgo, \ San Souci \ 20.XII.2008 - 15.I.2009 \ P.C. Grossi leg. Luz”; 1♂ “ Ecuador 85-01-6 \ Pichincha \ Dulluriquin \ Legit: Valencia”; 1♂ “ México, Jalisco, 14 Km \ SW de Hostotipaquilo \ 4-10.viii. 2013, 990m \ 21.012° N 104.179° W \ Malaise Melo & Rosa”.

Cephisus aff. j acobii: 1♂ “ Argentina, Jujuy, P.N. \ Calilegua 600-1110m, \ 23º41’38”S 64º52’4”W \ 16-15.I.2008 K. Ramos”; 1♂ “ Brasil, MG, 12 km a N de \ Águas Vermelhas, Faz. \ Faceiro, 15.640° S \ 41.477° W, 845m, \ 12.xii.2012, G. Melo & P. \ Grossi. Arm luminosa”.

Cephisus spp.: 1♀ “ Brasil, PR, Antonina \ Res. Rio Cachoeira \ 25.316°S 48.696°W \ 50m, 20-25.xi.2014 \ Luminosa suspensa \ Entomologia UFPR ”; 1♀ “Rosário Oeste \ MT, Brasil, XI.63 \ M. Alvarenga leg”; 1♀ “Jacaré - P. N. Xingu \ M. Grosso– Brasil \ 25-30/11/1965 \ M. Alvarenga leg”; 1♀ “ Brasil, RJ, Nova Friburgo, \ San Souci \ 20.XII.2008 - 15.I.2009 \ P.C. Grossi leg. Luz”; 1♀ “Ipixuna - AM, Rio \ Gregório; Com. Lago \ Grande, Seringal \ Recreio \ 7°10’6”S 70°49’6”W ” “ 145m - light trap \ 18-23 V.2011 \ Cavichioli, Gonçalves & \ Takiya”.