Quasitagalis afonsoi sp. nov., by present designation.

Quasitagalis gen. nov. can be separated from other genera of Saicinae of the New World by the combination of the characters presented in the key below; among them, Quasitagalis gen. nov. seems to be closer to Tagalis. However, these two genera can be promptly separated by the following set of characters: ventral portion of the head (gula) with a distal pair of strong setigerous spines, posterior to the eyes, in both genera, while only in Quasitagalis gen. nov., another pair is present below (between) the eyes; scutellum tapering into an erect spine in Tagalis and slightly elevated, subtriangular, without a spine in Quasitagalis gen. nov.; and inner surface of fore tibia with three or four (T. femorata Melo, 2008) very strong setigerous spines implanted close to dorsal surface (Tagalis) or with a simple (male) or double (female) longitudinal median row of numerous shorter spines (Quasitagalis).

Adults. Head: transversal sulcus deep, reaching eyes at hind margin; postocular portion subglobose, faintly depressed at median portion. Eyes globose in dorsal view, suboval in lateral view; strong setigerous spines anteroventrally and posteroventrally from eyes, the former somewhat smaller than the latter and ventrally, on gula, two pairs of similar setigerous spines: one pair below (between) the eyes and other pair, posterior to the eyes, somewhat closer to the neck than to the eyes. Antennal segments slender; segment I longest, clothed with long fine (ciliated) setae in males; segment II and III longer than half or half as long as the first segment; segment IV approximately one third as long as the first segment. Labium: segment II slender, elongate, curved, almost reaching posterior margin of eyes, with a pair of stout spines slightly basal to midpoint; segment III swollen mainly in the first third, where another pair of stout spines are located; segment IV slender, tapering. Thorax: prothorax divided by a transverse deep furrow between fore and hind lobes of pronotum, interrupted at median portion, above which there is a deep small excavation; anterolateral angles prominent as rounded tubercles; fore lobe subquadrate with pairs of lateral somewhat acute dorsal swellings or humps anteriorly and posteriorly, the latter more prominent; a longitudinal shallow and narrow median furrow, slightly larger at midportion, its posterior portion ending at the median deep excavation mentioned above; disc of fore lobe finely rugose; hind lobe trapezoidal, becoming larger to the posterior margin; integument more coarsely rugose on disc, which is slightly depressed; humeral angles rounded. Lateral shallow ridge reaching from tubercles of anterolateral angles towards posterior swellings of fore lobe, prominent at anterior half and shallower posteriorly. Scutellum base broad, apex slightly elevated, subtriangular, spineless. Metanotum with a short erect tubercle followed by a short obliquely erect spine larger at base and blunt at apex. Proepisternal processes projected with a pair of strong setigerous spines, anterodorsal spine moderately curved, posteroventral almost straight. Supracoxal lobes of propleura prominent. Prosternum larger on anterior margin; stridulitrum long, narrow. Mesosternum larger than prosternum and metasternum; meso and metasternum with a longitudinal, thin, shallow median keel. Fore legs stouter and shorter than others; fore coxa elongated, cylindrical, with a long spine on basal third of anterior surface, and three or five spines on inner face; mid and hind coxae ovoid; trochanters triangular, tapering; fore trochanter with four spines on inner side, three of which closer to each other at approximately basal half and the fourth spine variably set more or less apart and more ventrally at the apical half of the segment; fore femur stout, slightly curved in lateral view, armed ventrally with a few short spines and a variable number of few longer ones intermixed, a small subapical ventral protuberance with two or three small spines; a row of short spines on upper margin of inner surface; between the latter and the ventral line of spines, in the females, a row of somewhat more numerous, setigerous spines; fore tibia slightly curved in lateral view, apically expanded, with a single (male) or double (female) longitudinal median row of numerous spines running on approximately 1/2 (male) or 2/3 (female) of the median portion of inner surface. Middle and hind legs long and slender. Tarsi with three segments; first the longest; claws simple; scopula present on ventral portion of apex of third tarsomere of all legs. Forewings with two closed cells; distal cell much larger than basal one. Abdomen elongated, cylindrical in the male and ovoid in the female. In male, pygophore with a medial distal process; parameres symmetrical, short, apex rounded with an apical elongate acute spine acute at its centre.

Brazil, State of Tocantins.

The name of the new genus was composed by the Latin word quasi, meaning almost, nearly, like, and Tagalis, in reference to its apparent proximity to the latter genus. The gender is feminine.

Brazil, Tocantins, Lavandeira, Gruta da Gia [Gia’s Cave], 12°49'42"S, 46°20'43"W, 05–10.i.2009, Robson A. Zampaulo leg.: Holotype (male), 2 Paratypes (females) (MNRJ), 1 Paratype (female), (CTJMSB, 861).

Male. Figures 1–20. Measurements are given in Table 1.

Coloration. General coloration testaceous; approximately distal half of first antennal segment and the other antennal segments darkened; articulations between the segments and extreme apex of antennal segment IV pale; femora somewhat paler, the fore pair even more; fore femora with a small subapical pair of lateral dark spots on inner and outer surfaces just distal or above a small spiny protuberance; middle and hind femora with a subapical faint darkened ring; tarsi pale to whitish; forewings greyish, with the veins slightly darkened; hind wings translucent, veins pale yellowish; abdomen with a reddish tinge on the connexivum and on adjacent portion of tergites; most abdominal segments paler, darkened to the apex, including the ventral visible portion of segment VIII and the genital capsule (Fig. 1).

Vestiture. Body generally covered by sparse, thin, pale, suberect, obliquely erect or adpressed setae. Head with somewhat longer and more numerous setae on anterior portion, clypeus, labrum and anterolateral surfaces of first visible labial segment; on ventral portion of head (gula), several rows of shorter, more numerous, obliquely erect setae as a pubescence; antennal segments I and II covered with adpressed or obliquely semi-erect thin pale setae; on segment I, much longer fine (ciliated) setae, which are approximately three to four times as long as the transverse width of the segment (Fig. 4); segments III and IV covered with more numerous and shorter, adpressed, straight or slightly curved thin setae. Thorax: dorsal portion mostly glabrous, except in the anterior and lateral portions of mesoscutum which are covered by setae; median portion of meso- and metasternum covered with numerous short, erect setae forming a pubescence. Legs: coxae and trochanters covered with thin, decumbent, pale setae; armature of inner face of left fore coxa with three spines, lined on the same direction, at submedian basal, submedian distal and apical positions (Fig. 5); right fore coxa with five spines, four lined at similar positions as the other coxa and other smaller spine more anteriorly located, at the level between the two more basal spines (Fig. 6); fore femora covered with numerous thin, long, obliquely erect setae and more abundant erect setae on ventral surface; among the latter, five setae are even longer, approximately as long as the width of the segment, straight and somewhat larger (Fig. 5); armature with seven spines on upper margin of inner surface (Figs 5, 7); ventrally, at approximately the basal third, two long spines preceded by one spine half as long as the others and very short spines along the approximately distal two thirds of the segment (Fig. 5); a subapical ventral protuberance shallow with three minute spines (Fig. 5); middle and hind femora, tibiae and tarsi generally covered with thin, short and long setae, which are even more numerous towards apical portion of tibiae. Armature of fore tibiae with a single longitudinal median row of 17 (left tibia) (Fig. 5) or 18 (right tibia) spines at inner face, beginning somewhat far from base and ending far from apex of the segment, running by a distance of approximately 1/2 of the length of the segment; the spines are somewhat larger basally, becoming smaller towards distal portion of the row; additionally, three small spines, slightly positioned anteriorly, intercalated with spines which form the row at approximately its distal half (Fig. 5). The distal portion of the fore tibiae generally more densely covered by numerous stout adpressed setae; on inner face, distally to the end of the rows, these are even more numerous and subapically, a small subapical pecten. Forewing mostly glabrous, with thin, long setae on the costal vein, basal portion of the area subjacent to the anal vein, and short sparse setae on the margins of the pterostigma. Hind wing glabrous. Abdomen covered by thin, pale setae, which are generally shorter on the tergites and longer on sternites. Ventral portion of distal third of pygophore covered by very long, thin and numerous setae (Figs 8–10); a tuft of setae anterior to the implantation of parameres (Figs 8–10); posterior surface of medial process of the pygophore with a few short erect setae (Figs 8, 10, 12).

Structure. Venation of both wings similar to that of the female (Figs 34, 37). Segment VIII sclerotised on ventral portion, dorsal portion membranous; spiracles above dorsal margin of sclerotised ventral portion (Figs 8, 9). Male genitalia (Figs 8–20): genital capsule in situ (Figs 8, 9) with the apex of dorsal phallothecal sclerite (adps) prominent, just anterior to medial process of pygophore (mpp) and only the distal third of parameres (pa) visible, within pygophore rim. Pygophore in dorsal view (Fig. 11): somewhat elongated and ovoid in shape; between anterior and posterior genital openings, a dorsal (transverse) narrow bridge; margins of anterior opening subrounded; margins of posterior opening sinuous; in lateral view (Fig. 10): ventral margin rounded; dorsal margin rounded at approximately basal half and almost straight at distal half. Medial process of pygophore (mpp) (Figs 8, 10–12) narrow, somewhat elongated, curved in lateral view, tapering. Paramere (pa) (Figs 8, 9, 10, 13) short, strongly curved at approximately middle third, somewhat wider at distal two thirds; rounded at apex, in which a strong median apical acute spine is implanted in the same direction of the body of the paramere; glabrous at approximately basal two thirds and with sparse elongated somewhat curved thin setae scattered on distal third. Phallus (Figs 14–20): articulatory apparatus with short, stout basal arms (ba) (Figs 14, 15, 19), connected by a narrow basal bridge (bb) (Fig. 14); basal arms with a small, pointed inferior prolongation (Fig. 14, ip). Dorsal phallothecal sclerite (dps) (Figs 14–19) faintly sclerotised, elongated, curved in lateral view (Figs 15, 17); in dorsal view (Fig. 14): larger at basal portion, subtriangular at basal third, progressively narrowing towards middle third and somewhat enlarged at distal third, the latter surpassing the other elements of the phallus (Figs 15, 16); apical margin rounded (Figs 14–16). Ventrally to the dorsal phallothecal sclerite, numerous, elongated, sclerotised, curved, laminar processes (lp) (Figs 14–18, 20). These processes are larger at their bases, narrowing, not uniformly, towards their apices, which vary between presenting from a thin to a wider width as well as their tips which also vary as being rounded to acutely pointed (Fig. 20).

Female. Figures 21–51. Measurements are given in Table 1. Similar to male in general. The recorded differences in size are given in Table 1.

Vestiture. first antennal segment without long ciliated setae (Figs 25, 27); armature of inner face of both coxae with three spines, lined in a same direction, at submedian basal, submedian distal and apical positions (Figs 38, 39, 42); armature of fore femora (Figs 38–40, 43–46) with nine (Figs 39, 44) to ten (Figs 38, 43) spines on upper margin of inner surface; between the latter and the ventral line of spines, an intermediate row with 19 spines, similar in size (Figs 38, 39, 43); ventrally, at approximately the basal two thirds, four to seven long spines intermixed with four or five shorter spines; at approximately distal third, four to eight shorter spines, similar in size or progressively smaller towards the apex of the segment (Figs 38, 39, 43, 45, 46). Armature of fore tibiae (Figs 38, 39, 48, 49) with a double row of numerous spines, the posterior row begins closer to the base of the segment and runs by approximately 2/3 along of the inner surface; the anterior row begins more distally, approximately posteriorly to the third or fifth posterior spine; both rows ending far from the apex; the spines are generally stronger than those recorded in the male; the approximately 11–15 anterior and 16 posterior spines are mostly implanted intercalated in relation to each other and the former are generally smaller than the latter (Figs 38, 39, 48, 49); in one specimen there were five or six small additional spines randomly distributed at the mid portion of the rows and the two or three distal spines of the posterior row are thinner than the other posterior spines.

Structure. Venation of both wings as shown in Figs 34, 37. Female genitalia: posterior view of external genitalia as in Fig. 51.

Brazil, State of Tocantins, Lavandeira municipality, Gruta da Gia [Gia’s Cave], 12°49'42"S, 46°20'43"W.

The new species is named in honour to Professor and Researcher Dr Luiz Afonso Vaz de Figueiredo for his role as an environmentalist, responsible for the training of countless educators; a great supporter of Speleology in Brazil.

Among the two common sexual dimorphic characteristics recorded among Saicinae, such as in species of Tagalis (e.g., Gil-Santana et al. 2010, Gil-Santana 2011), the larger eyes and longer, ciliated setae on the first antennal segment of males, only the latter was undoubtedly recorded in the specimens of Quasitagalis afonsoi examined here (Fig. 4). The eyes of the male holotype showed the same measurements as one female and did not seem much larger (Figs 1–3, 21–23) as it was recorded in males of some species of Tagalis (e.g., Gil-Santana et al. 2010; Gil-Santana 2011; Castro-Huertas and Forero 2014), but not in Tagalis i. inornata in which, the male eyes were shown not to be much larger than those of the females (Gil-Santana 2011). On the other hand, the armature of the fore femora of the females had more spines in general, including an additional row of spines (intermediate between those of upper margin and ventral margin of inner face) (e.g., Figs 5, 38, 39). Yet, in the females, the armature of the fore tibiae was more prominent, formed by a double row of much stronger spines, extending comparatively for a longer distance on the inner surface (Figs 38, 39, 48, 49), while in the male holotype, although there were three small, anterior spines analogous to that of the anterior row of the female, the armature was basically formed by a single row of generally smaller spines, extending for a shorter distance on the inner surface of its tibiae (Fig. 5). Similarly, for Tagalis, McAtee and Malloch (1923) argued that the spines on the fore femora were sexually dimorphic, while the armature of the fore femora showed smaller spines in the males of Tagalis i. inornata than in the females of this species examined by Gil-Santana (2011). However, both the confirmation that the eyes are of similar size in both sexes as well as if the differences in the armature of the fore legs are part of sexual dimorphism or a part of individual variation in Quasitagalis afonsoi would be better evaluated through examining more specimens.

Quasitagalis afonsoi gen. et sp. nov. Brazil, Tocantins, Lavandeira, Gruta da Gia [Gia’s Cave], 05–10.i.2009, 12°49'42"S, 46°20'43"W, Robson A. Zampaulo leg., 2 nymphs (CTJMSB, 861). Tagalis inornata inornata Stål, 1860. Brazil, Rio de Janeiro, Nova Friburgo, 22°17'S, 42°29'W, 1.049 m, 01 female, 09.xii.1997 (CTJMSB, 862).

Two nymphs of Quasitagalis afonsoi were collected with the adults. They were from different and undetermined instars and were not in good condition for formal or complete descriptions. However, the head of the smaller (earlier instar) nymph and fore legs of both of them were well conserved and were used to obtain SEM images (Figs 52–59). Their measurements are given in Table 2.

They are shown to have bi-segmented tarsi (Figs 54, 57) and smaller eyes (Fig. 52), common features found in immature Heteroptera (Schuh and Weirauch 2020; Rédei 2004) and also in the nymphs of Tagalis evavilmae (Gil-Santana et al. 2010). The armature of the head is like those recorded in the adults, i.e., with strong setigerous spines anteroventrally and posteroventrally from eyes, the former somewhat smaller than the latter and ventrally, on the gula, two pairs of similar setigerous spines: one pair below (between) the eyes and other pair, posterior to the eyes, somewhat closer to the neck than to eyes (Fig. 52); labial segments II and III with a pair of stout spines, slightly basal to midpoint and on swollen portion, respectively (Figs 52, 53). The structure and armature of the fore coxa is also like those of the adults in general (a long spine on basal third of anterior surface and three spines on its inner surface) (Figs 54, 55). The armature of the fore trochanter is similar, but with small differences: the apical (fourth) spine is comparatively smaller and there is an additional spine below the third spine in the nymph of earlier instar (Fig. 54). The armature of the fore femora is equivalent to that of the adult female (Figs 54–56, 58): armed ventrally with a few short spines and a variable number of few longer ones intermixed at approximately the basal two thirds and a small subapical ventral protuberance with small spines; a row of spines on the upper margin of the inner surface, which are comparatively longer than those of adults (with ten elements in the earlier instar nymph and nine in the later instar nymph); between the latter and the ventral line of spines, a row of setigerous spines, more variably in size, with approximately 14 elements in both nymphs. The armature of the fore tibia, however, is conspicuously different from that of the adults, with a submedian (somewhat lateral to the median portion of the segment) row of six (earlier instar nymph) (Fig. 54) or eight (later instar nymph) (Figs 55, 56) strong, long spines on the inner surface (besides the more distal spine in both nymphs which is shorter and smaller); at the lateral border of the inner surface, three larger but shorter setigerous spines (Figs 54–56); at the apical portion of the dorsal surface, two or three small curved spines (Figs 54, 56, 57).

The municipality of Lavandeira is located at 12°47'19"S, 46°24'28"W, in the southeast of the State of Tocantins and Northern Brazil (Figs 60, 61, 63) with an altitude of approximately 330 meters and an area of 519,614 km². The climate in the region is tropical with average annual rainfall ranging from 1,400 to 1,600 mm and average temperature ranging from 25 to 27 °C (SEPLAN 2008). The predominant vegetation is the Cerrado (Brazilian savannas), covering 87.8 % of the state’s area, the rest is occupied by forests. It is noteworthy that the Cerrado is considered one of the main biodiversity hotspots (priority areas for biodiversity conservation) worldwide (Myers et al. 2000; Mittermeier et al. 2011; Williams et al. 2011).

The carbonates present in the region are located in the Speleological Province of the Bambuí Group (São Domingos District) and distributed in a north-south direction. At this portion, elevations vary between 400 and 600 m, while elevations below 400 m dominate the northwestern portion of the study area. The Bambuí Group constitutes the largest set of limestone occurrences, favourable to the presence of caves in Brazil (Karmann and Sánchez 1979).

Currently, the State of Tocantins has 939 caves registered in the official government databases (CECAV 2019), with approximately 350 known caves in the southeastern region. However, karst in the region has the potential for thousands of caves. Gruta da Gia [Gia’s Cave] is located in the central portion of the municipality of Lavandeira (Fig. 63) under the 12°49'42"S, 46°20'43"W at 503 m high. Inserted at the top of the limestone massif, this cave is approximately 200 m of long and has only one entrance (Fig. 62). It is a humid cave with clay soil, extensive aphotic zones and low availability of trophic resources. The surrounding landscape is made up of pasture areas, but there are still extensive fragments of native forest associated with karst outcrops and springs areas. In general, the caves in the region have small dimensions, the majority of which are less than 50 m long. They have a predominantly flat floor, and few have underground lakes or rivers.