Research Article |
Corresponding author: Hélcio R. Gil-Santana ( helciogil@uol.com.br ) Academic editor: Laurence Livermore
© 2020 Hélcio R. Gil-Santana, Jader Oliveira, Robson de A. Zampaulo.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gil-Santana HR, Oliveira J, Zampaulo RA (2020) Quasitagalis afonsoi, a new genus and a new species of Saicinae (Hemiptera, Reduviidae) inhabiting a cave in Brazil, with an updated key to the genera of Saicinae of the New World. ZooKeys 966: 9-39. https://doi.org/10.3897/zookeys.966.52930
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Quasitagalis afonsoi gen. et sp. nov. of Saicinae (Hemiptera, Reduviidae) is described based on a male and three female specimens collected in a cave in the State of Tocantins, Brazil. Additionally, some characteristics from two nymphs of different instars of the same species are also recorded. An updated key to the New World genera of Saicinae is provided.
Heteroptera, male genitalia, Neotropics, Paratagalis, Tagalis
There are ten genera of Saicinae in the Neotropical region, five of which are currently monotypic (
Little is known of the biology and natural history of Saicinae (
Among Neotropical Saicinae, sexual dimorphism consisting of larger eyes and longer, ciliated setae on the first antennal segment of males was observed in Paratagalis spinosus Monte, 1943 (
On the other hand,
A scopula, previously documented for some taxa of Emesinae (
Our knowledge of immature stages of Saicinae is very limited (
Quasitagalis afonsoi gen. et sp. nov. of Saicinae (Hemiptera, Reduviidae) is described based on a male and three female specimens collected in a cave in the State of Tocantins, Brazil. Additionally, some characteristics from two nymphs of different instars of the same species are recorded too. An updated key to the New World genera of Saicinae is provided.
All fieldwork, including the collection of the specimens inside a cave, was undertaken by the third author (RAZ), who also provided Figs
Photographs of the male holotype and a female paratype of Quasitagalis afonsoi gen. et sp. nov. (Figs
Scanning electron microscopy images (Figs
All remaining figures were produced by the first author (HRG-S). The fixed adults, microscopic preparations, and genitalia were photographed using a digital camera (Sony DSC-W830). Drawings were made using a camera lucida. For clarity, the general vestiture (setation) in several ink drawings was completely (Figs
Observations were made using a stereoscope microscope (Zeiss Stemi) and a compound microscope (Leica CME). Measurements were made using a micrometer eyepiece. General morphological terminology mainly follows
The holotype and two female paratypes will be deposited in the Entomological Collection of the “Museu Nacional da Universidade Federal do Rio de Janeiro”, Rio de Janeiro, Brazil (
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) (
Male. Figures
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.
Measurements (mm) of adult types of Quasitagalis afonsoi gen. et sp. nov.
Male holotype | Female paratypes (N = 3) | |||
---|---|---|---|---|
Maximum | Minimum | Mean | ||
Body length to tip of forewing | 6.00 | 6.40 | 6.00 | 6.20 |
Body length to tip of abdomen | 5.50 | 6.20 | 5.70 | 5.90 |
Head length (excluding neck) | 0.70 | 0.70 | 0.70 | 0.70 |
Anteocular portion length | 0.20 | 0.30 | 0.20 | 0.23 |
Postocular portion length | 0.20 | 0.30 | 0.20 | 0.25 |
Head width across eyes | 0.60 | 0.60 | 0.60 | 0.60 |
Interocular distance (synthlipsis) | 0.30 | 0.30 | 0.30 | 0.30 |
Transverse width of eye | 0.20 | 0.20 | 0.10 | 0.17 |
Length of eye | 0.25 | 0.25 | 0.20 | 0.23 |
Antennal segment I length | 3.10 | 3.30 | 3.10 | 3.20 |
Antennal segment II length | 1.80 | 2.00 | 1.90 | 1.93 |
Antennal segment III length | 1.80 | 1.80 | 1.50 | 1.67 |
Antennal segment IV length | 1.00 | 1.00 | 0.90 | 0.97 |
Labial segment II length | 0.50 | 0.50 | 0.50 | 0.50 |
Labial segment III length | 0.30 | 0.30 | 0.30 | 0.30 |
Labial segment IV length | 0.20 | 0.25 | 0.20 | 0.21 |
Fore lobe of pronotum length | 0.60 | 0.60 | 0.60 | 0.60 |
Fore lobe of pronotum max. width | 0.65 | 0.70 | 0.65 | 0.68 |
Hind lobe of pronotum length | 0.50 | 0.50 | 0.50 | 0.50 |
Fore lobe of pronotum max. width | 0.90 | 1.00 | 0.90 | 0.97 |
Forewing length | 4.10 | 4.70 | 4.00 | 4.37 |
Fore coxa length | 0.80 | 0.80 | 0.70 | 0.76 |
Fore femur length | 2.10 | 2.20 | 1.90 | 2.06 |
Fore tibia length | 1.90 | 2.00 | 1.70 | 1.86 |
Fore tarsus length | 0.30 | 0.30 | 0.30 | 0.30 |
Mid femur length | 2.90 | 3.10 | 2.50 | 2.86 |
Mid tibia length | 3.80 | 3.80 | 3.40 | 3.66 |
Mid tarsus length | 0.25 | 0.25 | 0.25 | 0.25 |
Hind femur length | 4.10 | 4.50 | 3.80 | 4.23 |
Hind tibia length | 6.00 | 6.30 | 5.50 | 5.93 |
Hind tarsus length | 0.25 | 0.30 | 0.20 | 0.23 |
Abdomen length | 3.10 | 3.50 | 3.00 | 3.20 |
Abdomen maximum width | 0.90 | 1.40 | 0.70 | 1.06 |
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.
Quasitagalis afonsoi gen. et sp. nov., male holotype 4 first antennal segment (general vestiture omitted, except the ciliated setae) (A apex, B base) 5 left fore leg, lateral view, inner face (setae omitted, except the longer setae, pointed by arrows) 6 right fore coxa and trochanter, lateral view, inner face 7 left fore femur, dorsal view. Scale bars: 0.5 mm (4, 5, 7); 0.2 mm (6).
Structure.
Venation of both wings similar to that of the female (Figs
Quasitagalis afonsoi gen. et sp. nov., male holotype 8, 9 apical margin of abdominal segment VII, abdominal segment VIII and genital capsule 8 lateral view 9 dorsal view 10–13 male genitalia 10 pygophore and left paramere, lateral view 11 pygophore without parameres, dorsal view 12 medial process of pygophore, lateral view 13 right paramere. Abbreviations: adps apex of dorsal phallothecal sclerite, mpp medial process of pygophore, pa paramere, VIII abdominal segment VIII. Scale bars: 0.2 mm (8–11); 0.05 mm (12, 13).
Quasitagalis afonsoi gen. et sp. nov., male holotype, phallus 14 dorsal view 15–17 lateral view 16 distal portion 17 median portion. Abbreviations: adps apex of dorsal phallothecal sclerite, ba basal arm, bb basal bridge, dps dorsal phallothecal sclerite, ip inferior prolongation of the basal arm, lp laminar process (es). Scale bars: 0.1 mm (14, 15); 0.02 mm (16, 17).
Quasitagalis afonsoi gen. et sp. nov., male holotype, phallus. 18, 19 Basal portion, lateral view 20 shape of apical portion of some laminar processes of endosoma. Abbreviations: ba basal arm, bdps basal portion of dorsal phallothecal sclerite, lp laminar process. Scale bars: 0.05 mm (18, 19); 0.02 mm (20).
Female. Figures
Vestiture.
first antennal segment without long ciliated setae (Figs
Quasitagalis afonsoi gen. et sp. nov., female paratypes 25 upper portion of head and pronotum and antennal segment I, lateral view 26 antennal segment II 27–29 median portion of antennal segments 27 segment I 28 segment II 29 segment III 30 distal and basal portions of antennal segments III and IV (III and IV, respectively) 31 posterior portion of head and prothorax, lateral view 32 proepisternal process, lateral view. Scale bars: 0.2 mm (25, 26, 31); 0.1 mm (27–30); 0.05 mm (32).
Quasitagalis afonsoi gen. et sp. nov., female paratypes 33 meso- and metathorax (wings moved down), lateral view 34–36 forewing 35 basal portion 36 apical half of pterostigma and subjacent portion 37 hind wing. Abbreviations: sc scutellum, sp spine of metanotum, tb tuberculum of metanotum. Scale bars: 0.5 mm (34, 37); 0.2 mm (33, 35, 36).
Structure.
Venation of both wings as shown in Figs
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.,
Quasitagalis afonsoi gen. et sp. nov., female paratypes, fore leg 40–43, 45–47 lateral view 40, 41 outer face 41 coxa, trochanter and base of femur 42, 43 inner face 42 coxa and basal portion of trochanter 43 trochanter and femur 44 right femur, dorsal view 45, 46 segment of ventral portion of femur 47 tibia, outer face. Abbreviations: ls longer seta, lsp long spine, ssp short spine. Scale bars: 0.5 mm (44); 0.2 mm (40, 43, 45–47); 0.1 mm (41, 42).
Quasitagalis afonsoi gen. et sp. nov., female paratypes 48, 49 fore tibia, lateral view, inner face 49 portion of the double row of spines; at the center a pair of the latter in which the posterior spine is larger and longer than the anterior one 50 fore tarsus, ventral view, the arrow points to the scopula 51 genitalia, posterior view. Scale bars: 0.2 mm (48); 0.1 mm (51); 0.05 mm (50); 0.02 mm (49).
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
Measurements (mm) of nymph specimens of Quasitagalis afonsoi gen. et sp. nov.
Earlier instar nymph | Later instar nymph | |
---|---|---|
Body length to tip of abdomen | 3.50 | 4.90 |
Head length (excluding neck) | 0.50 | 0.60 |
Anteocular portion length | 0.20 | 0.20 |
Postocular portion length | 0.25 | 0.30 |
Head width across eyes | 0.40 | 0.50 |
Interocular distance (synthlipsis) | 0.30 | 0.30 |
Transverse width of eye | 0.05 | 0.10 |
Length of eye | 0.10 | 0.15 |
Antennal segment I length | 2.10 | 2.60 |
Antennal segment II length | 1.00 | 1.40 |
Antennal segment III length | 1.20 | 1.50 |
Antennal segment IV length | 0.90 | 1.00 |
Labial segment II length | 0.35 | 0.40 |
Labial segment III length | 0.20 | 0.30 |
Labial segment IV length | 0.15 | 0.20 |
Pronotum length | 0.50 | 0.60 |
Pronotum maximum width | 0.60 | 0.60 |
Wing pad length | 0.70 | 1.30 |
Fore coxa length | 0.50 | 0.60 |
Fore femur length | 1.40 | 1.70 |
Fore tibia length | 1.20 | 1.50 |
Fore tarsus length | 0.25 | 0.25 |
Mid femur length | 1.80 | 2.40 |
Mid tibia length | 2.30 | 3.10 |
Mid tarsus length | 0.20 | 0.20 |
Hind femur length | absent | 3.40 |
Hind tibia length | absent | 4.20 |
Hind tarsus length | absent | 0.20 |
Abdomen length | 1.00 | 2.50 |
Abdomen maximum width | 0.40 | 0.80 |
They are shown to have bi-segmented tarsi (Figs
Quasitagalis afonsoi gen. et sp. nov., later instar nymph, fore leg, lateral view 55, 56 inner face 55 entire leg 56 approximately distal two thirds of femur and tibia; the setae point to the larger setigerous spines on inner surface of fore tibiae, close to its dorsal surface 57 tarsus 58 basal portion of the ventral armature of femur 59 portion of tibia in which the second lateral spine (pointed by black arrows) is inserted; adjacent long submedian spine pointed by a white arrow. Scale bars: 0.2 mm (55, 56); 0.04 mm (57) 0.02 mm (58, 59).
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 Position of the Gruta da Gia [Gia’s Cave], the type locality of Quasitagalis afonsoi sp. nov., in relation to Brazil and the State of Tocantins 62 the entrance to Gruta da Gia [Gia’s Cave] 63 Position of the Gruta da Gia [Gia’s Cave] in relation to the municipality of Lavandeira, other caves identified in the region and the delimitation of carbonate rocks (the Bambuí Geological Group in light gray). Abbreviations: Brazilian States: BA Bahia, GO Goiás, MA Maranhão, MG Minas Gerais, MT Mato Grosso, PA Pará, PI Piauí, TO Tocantins.
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 (
Currently, the State of Tocantins has 939 caves registered in the official government databases (
It is noteworthy that, while Paratagalis and Quasitagalis have in common the presence of two pairs of setigerous spines on the gula (Figs
Regarding the male genitalia, it is noteworthy that the shape of the pygophore, its process, the endosoma portions (articulatory apparatus, dorsal phallothecal sclerite, curved elongated processes of endosoma) (Figs
More importantly, in all species of Paratagalis and Tagalis in which the paramere was described (
The presence of the scopula on the apex of the third tarsomere on all the legs in Q. afonsoi (Fig.
The three larger lateral setigerous spines on the inner surface of the fore tibiae, close to its dorsal surface, present in the nymphs of Q. afonsoi (Figs
Future phylogenetic analyses should be carried out to assess all these similarities and differences and the taxonomic validity of the genera and their systematic positions, and the relationships among species of New World genera of Saicinae.
Several species of Emesinae, a group considered as close related to Saicinae (e.g.,
Based on
1 | Foreleg without spines, at most with erect setae | 2 |
– | Fore femur with two or three rows of spines, fore tibiae either with setae or with spines | 5 |
2 | Posterior pronotal lobe with upward projecting spines or tubercles; mesoscutum (scutellum) and metanotum apically with vertical spines or tubercles | 3 |
– | Pronotum generally unarmed, but sometimes with humeral spines; apex of mesoscutum produced into a long horizontal tapering spine, metanotum without spine or tubercle | Oncerotrachelus Stål, 1868 |
3 | Opposed surfaces of labium and head with spine-like setae or bristles; forewing with two to three cells; metapleura without a tubercle near coxal cavity | 4 |
– | Opposed surfaces of labium and head without spine-like setae or bristles; forewing with four cells; metapleura with a tubercle near coxal cavity | Saicireta Melo & Coscarón, 2005 |
4 | Process on lower anterior angle of prothorax acute to subacute; second antennal segment approximately half as long as first antennal segment; medial process of male pygophore bifurcate; posterior margin of seventh abdominal sternite in females vertical or subvertical | Saica Amyot & Serville, 1843 |
– | Process on lower anterior angle of prothorax subconical; second antennal segment approximately 3/4 as long as first antennal segment; medial process of male pygophore a single, erect spine; posterior margin of seventh abdominal sternite in females sloping ventrocephalad | Pseudosaica Blinn, 1990 |
5 | Humeral angles of pronotum without processes, rounded | 6 |
– | Humeral angles of pronotum with spine-like processes | 7 |
6 | Ventral portion of the head below (between) the eyes spineless; fore tibiae with a three or four (T. femorata) stronger, setigerous spines implanted on external border of inner surface, close to dorsal surface | Tagalis Stål, 1860 |
– | Head with a ventral pair of spines below (between) the eyes; fore tibiae with a single or double longitudinal row of numerous short spines on median portion of inner surface | Quasitagalis gen. nov. |
7 | Fore coxae and anterior pronotal lobe unarmed | Bagriella McAtee & Malloch, 1923 |
– | Fore coxae spined, anterior pronotal lobe with four spines or rounded humps | 8 |
8 | Fore lobe of pronotum with four spines | Paratagalis Monte, 1943 |
– | Fore lobe of pronotum with four humps | 9 |
9 | Two first (visible) labial segments spiny; only apterous females known | Kiskeyana Weirauch & Forero, 2007 |
– | Only the first (visible) or all three labial segments spiny; females macropterous | 10 |
10 | Only the first (visible) labial segment spiny; forewings with four closed cells | Buninotus Maldonado, 1981 |
– | All three (visible) labial segments spiny; forewings with two closed cells | Caprilesia Gil-Santana, Marques & Costa, 2006 |
We are grateful to João Paulo Sales Oliveira Correia (LNIRTT,