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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
Hélcio R. Gil-Santana1, Jader Oliveira2, Robson de A. Zampaulo3
1 Laboratório de Diptera, Instituto Oswaldo Cruz, Av. Brasil, 4365, 21040-360, Rio de Janeiro, RJ, Brazil
2Laboratório de Parasitologia, Universidade Estadual Paulista “Julio de Mesquita Filho”, Faculdade de Ciên-
cias Farmacêuticas UNESP/FCFAR, Rodovia Araraquara Jaú, KM 1, 14801-902, Araraquara, SP, Brazil
3VALE SA, Gerência de Licenciamento e Espeleologia (COI), Av. de Ligação, 3580, prédio 1, 1° andar, Mina
de Águas Claras, 34000000, Nova Lima, MG, Brazil
Corresponding author: Hélcio R. Gil-Santana (helciogil@uol.com.br; helciogil@ioc.ocruz.br)
Academic editor: Laurence Livermore |Received 5 April 2020|Accepted 31 July 2020|Published 9 September 2020
http://zoobank.org/16B82020-77F1-4690-AD03-2F9629C31A88
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
Abstract
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 characteris-
tics from two nymphs of dierent instars of the same species are also recorded. An updated key to the New
World genera of Saicinae is provided.
Keywords
Heteroptera, male genitalia, Neotropics, Paratagalis, Tagalis
ZooKeys 966: 9–39 (2020)
doi: 10.3897/zookeys.966.52930
https://zookeys.pensoft.net
Copyright Hélcio R. Gil-Santana et al. 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.
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Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
10
Introduction
ere are ten genera of Saicinae in the Neotropical region, ve of which are currently
monotypic (Gil-Santana et al. 2015). A summary of the taxonomy of this group and a
key to the genera of the New World were provided by Gil-Santana et al. (2015).
Little is known of the biology and natural history of Saicinae (Gil-Santana et al.
2010). Specimens have been most commonly collected at lights (Schuh and Weirauch
2020) or swept and beaten from vegetation (Gil-Santana et al. 2015). Gil-Santana et
al. (2010) included a synopsis of the biological and ecological information available
for New World Saicinae, recording Tagalis evavilmae Gil-Santana, Gouveia & Zeraik,
2010 as an inhabitant of birds’ nests, a rst for Saicinae.
Among Neotropical Saicinae, sexual dimorphism consisting of larger eyes and
longer, ciliated setae on the rst antennal segment of males was observed in Paratagalis
spinosus Monte, 1943 (Gil-Santana and Costa 2009), and it was conrmed in the fol-
lowing species of Tagalis Stål, 1860: T. evavilmae, and T. seminigra Champion, 1899
(Gil-Santana et al. 2010). In T. inornata inornata Stål, 1860, however, the male eyes
were shown not to be much larger than those of females and the long ciliated setae on
the rst antennal segments were shorter than those of some other species (Gil-Santana
2011). Large eyes and long ciliated setae on the rst antennal segments of males have
also been recorded in the following species of Tagalis: T. baenai Gil-Santana, 2011,
T. grossii Gil-Santana, 2011, T. marquesi Gil-Santana, 2011 (Gil-Santana 2011), T.
dichroa Castro-Huertas & Forero, 2014 (Castro-Huertas and Forero 2014) and T. dr-
akkar Varela & Melo, 2017 (Varela and Melo 2017). However, since no females of
these species are known, the possible sexual dimorphism could not be veried. Longer
ciliated setae on the rst third of the second antennal segment of males was also ob-
served in P. spinosus (Gil-Santana and Costa 2009), while they were recorded on both
the rst and the second antennal segments in males of the species of Oncerotrachelus
Stål, 1868 studied by Gil-Santana (2013).
On the other hand, McAtee and Malloch (1923) recorded that the length of the
spines on the fore femora for Tagalis was sexually dimorphic and, to some extent, also
exhibited intraspecic variation, and therefore would not seem to be of taxonomic im-
portance. e armature of the fore femora in T. i. inornata was shown to have smaller
spines in males when compared to those of the females (Gil-Santana 2011). However,
the apparent sexual variation in length of the spines of the fore femora (McAtee and
Malloch 1923) may be related to the size of the individual, since the females are usually
larger. is characteristic and individual variation would be better evaluated through
examination of more specimens, including other species.
A scopula, previously documented for some taxa of Emesinae (Wygodzinsky 1966),
was recently recorded in several Saicinae (Weirauch and Forero 2007a; Weirauch 2007,
2008a). In Saicinae, the scopula is present as a hairy attached structure on the ventral
surface of the apex of the third tarsomere of all pairs of legs. It was recorded in sev-
eral species of Saicinae (Weirauch 2007), among which only two were from the New
World, Saica recurvata (Fabricius, 1803) (Weirauch 2007) and Kiskeyana palassaina
Quasitagalis afonsoi a new genus and a new species of Saicinae 11
Weirauch & Forero, 2007 (Weirauch and Forero 2007a, b). Later, Castro-Huertas and
Forero (2014) recorded the presence of the scopula on the apex of the third tarsomere
in all the legs on both species of Tagalis described by them. Regarding its function,
Weirauch and Forero (2007a) speculated that the scopula might be a structure that
assists in movement on smooth (e.g., plant) surfaces, while Weirauch (2007) although
arguing that the scopula in Saicinae could be exclusively used for locomotion or that
it could play a certain role in prey capture, concluded that its primary function is
probably during locomotion. Yet, it was considered as a synapomorphy of part of the
Saicinae by Weirauch (2008a).
Our knowledge of immature stages of Saicinae is very limited (Rédei 2004). Among
the Saicinae from the New World, nymphal stages have only been described in one
species, Tagalis evavilmae (Gil-Santana et al. 2010). On one hand, the latter authors
recorded that the nymphs of T. evavilmae show common features found in immature
Heteroptera, such as a bi-segmented tarsi and smaller eyes in younger instars (Schuh
and Weirauch 2020; Rédei 2004). On the other hand, Gil-Santana et al. (2010) also
recorded the presence of dierent patterns of features with taxonomical signicance in
nymphs when compared with adults (like an additional spine on the ventral side of the
head, more than three spines on tibiae) and argued that these characteristics which are
observed in adults of other related genera, might help to understand the relationships
among the genera of Saicinae in future studies.
Quasitagalis afonsoi gen. et sp. nov. of Saicinae (Hemiptera, Reduviidae) is de-
scribed 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 dierent
instars of the same species are recorded too. An updated key to the New World genera
of Saicinae is provided.
Materials and methods
All eldwork, including the collection of the specimens inside a cave, was undertaken
by the third author (RAZ), who also provided Figs 60–63.
Photographs of the male holotype and a female paratype of Quasitagalis afonsoi gen.
et sp. nov. (Figs 1, 21) were taken by João Paulo Sales Oliveira Correia (“Laboratório
Nacional e Internacional de Referência em Taxonomia de Triatomíneos” (LNIRTT),
Instituto Oswaldo Cruz (IOC), Rio de Janeiro, Brazil), with a Leica DMC 2900 cam-
era attached to a Leica M205C stereomicroscope. Several images were stacked using
the LAs software version 4.9.
Scanning electron microscopy images (Figs 23–28, 31, 32, 38, 40–43, 45–59, 64,
65) were obtained by the second author (JO). A female, two nymphs of dierent instars
of the new species, and a female of Tagalis inornata inornata Stål, 1860 were cleaned in
an ultrasound machine. Subsequently, the samples were dehydrated in alcohol, dried in
an incubator at 45 °C for 20 min, and xed in small aluminium cylinders with trans-
parent glaze. Sputtering metallisation was then performed on the samples for 2 min at
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
12
10 mA in an Edwards sputter coater.After this process, the samples were studied and
photographed using a high-resolution eld emission gun scanning electron microscope
(FEG-SEM; JEOL, JSM-7500F), similarly as described by Rosa et al. (2010, 2014).
All remaining gures were produced by the rst author (HRG-S). e xed adults,
microscopic preparations, and genitalia were photographed using a digital cam era
(Sony DSC-W830). Drawings were made using a camera lucida. For clarity, the gen-
eral vestiture (setation) in several ink drawings was completely (Figs 2, 3, 6, 7, 11, 22,
33, 34, and 44) or almost completely (Figs 4, 5, and 39) omitted. Images were edited
using Adobe Photoshop CS6. Dissections of the male genitalia were made by rst
removing the pygophore from the abdomen with a pair of forceps and then clearing it
in 20% NaOH solution for 24 hours. e dissected structures were studied and pho-
tographed or drawn in glycerol.
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 Lent and Wygodzinsky
(1979) and Schuh and Weirauch (2020). e (visible) segments of labium are num-
bered as II to IV, given that the rst segment is lost or fused to the head capsule in
Reduviidae (Weirauch 2008b, Schuh et al. 2009). Gil-Santana et al. (2010) and Gil-
Santana (2011) were mostly followed in the case of the terms applied to structures that
are characteristic or peculiar to Saicinae.
e 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 (MNRJ) and the female paratype, the nymphs and a female of Tagalis
i. inornata used for obtain SEM images were deposited in the Dr Jose Maria Soares
Barata Triatominae Collec tion (CTJMSB) of the São Paulo State University Julio de
Mesquita Filho, School of Pharmaceutical Sciences, Araraquara, São Paulo, Brazil. All
measurements are in millimetres (mm).
Results
Taxonomy
Subfamily Saicinae
Quasitagalis gen. nov.
http://zoobank.org/1549BE8A-08FE-455F-A9BB-FB42A97E373A
Type species. Quasitagalis afonsoi sp. nov., by present designation.
Diagnosis. 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
Quasitagalis afonsoi a new genus and a new species of Saicinae 13
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 (be-
tween) 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).
Description. 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 pos-
teroventrally 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. An-
tennal segments slender; segment I longest, clothed with long ne (ciliated) setae in
males; segment II and III longer than half or half as long as the rst segment; segment
IV approximately one third as long as the rst 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 rst third, where another
pair of stout spines are located; segment IV slender, tapering. orax: prothorax divid-
ed 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 nely 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, an-
terodorsal spine moderately curved, posteroventral almost straight. Supracoxal lobes of
propleura prominent. Prosternum larger on anterior margin; stridulitrum long, nar-
row. Mesosternum larger than prosternum and metasternum; meso and metasternum
with a longitudinal, thin, shallow median keel. Fore legs stouter and shorter than oth-
ers; fore coxa elongated, cylindrical, with a long spine on basal third of anterior surface,
and three or ve 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
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
14
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 approxi-
mately 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; rst 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.
Distribution. Brazil, State of Tocantins.
Etymology. e 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. e gender is feminine.
Quasitagalis afonsoi sp. nov.
http://zoobank.org/8006DD5A-3868-4742-B152-86B2640437E2
Figures 1–51
Type material. 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).
Description. Male. Figures 1–20. Measurements are given in Table 1.
Coloration. General coloration testaceous; approximately distal half of rst an-
tennal 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 in-
ner 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; abdo-
men 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 rst 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 ne (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
Quasitagalis afonsoi a new genus and a new species of Saicinae 15
Table 1. 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
or slightly curved thin setae. orax: dorsal portion mostly glabrous, except in the an-
terior and lateral portions of mesoscutum which are covered by setae; median portion
of meso- and metasternum covered with numerous short, erect setae forming a pubes-
cence. 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 sub-
median basal, submedian distal and apical positions (Fig. 5); right fore coxa with ve
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, ve 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
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
16
Figures 1–3. Quasitagalis afonsoi gen. et sp. nov., male holotype 1 lateral view 2 head and pronotum,
dorsal view 3 head, lateral view. Scale bars: 2.0 mm (1); 0.5 mm (2); 0.2 mm (3).
Quasitagalis afonsoi a new genus and a new species of Saicinae 17
Figures 4–7. Quasitagalis afonsoi gen. et sp. nov., male holotype 4 rst antennal segment (general ves-
titure 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 7left fore femur, dorsal view. Scale bars: 0.5 mm (4, 5 , 7 ); 0.2 mm (6).
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
18
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 seg-
ment, 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). e 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). Seg-
ment 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 (trans verse) narrow bridge; margins of anterior opening subround-
ed; margins of posterior opening sinuous; in lateral view (Fig. 10): ventral margin
rounded; dorsal margin rounded at approximately basal half and almost straight at dis-
tal half. Medial process of pygophore (mpp) (Figs 8, 10–12) narrow, somewhat elon-
gated, 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 prolon-
gation (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 phal-
Quasitagalis afonsoi a new genus and a new species of Saicinae 19
Figures 8–13. Quasitagalis afonsoi gen. et sp. nov., male holotype 8, 9 apical margin of abdominal seg-
ment 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 scle-
rite, mpp medial process of pygophore, pa paramere, VIII abdominal segment VIII. Scale bars: 0.2mm
(8–11); 0.05 mm (12, 13).
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
20
lothecal sclerite, numerous, elongated, sclerotised, curved, laminar processes (lp) (Figs
14–18, 20). ese 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. e recorded dierences in size are given in Table 1.
Vestiture. rst 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 mar-
gin 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 ve shorter spines;
at approximately distal third, four to eight shorter spines, similar in size or progres-
sively 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 fth 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 ve 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: pos-
terior view of external genitalia as in Fig. 51.
Distribution. Brazil, State of Tocantins, Lavandeira municipality, Gruta da Gia
[Gia’s Cave], 12°49'42"S, 46°20'43"W.
Etymology. e 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.
Comments. 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 rst antennal segment of males,
only the latter was undoubtedly recorded in the specimens of Quasitagalis afonsoi ex-
amined here (Fig. 4). e 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
Quasitagalis afonsoi a new genus and a new species of Saicinae 21
Figures 14–17. Quasitagalis afonsoi gen. et sp. nov., male holotype, phallus 14 dorsal view 15–17lateral
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).
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
22
and ventral margin of inner face) (e.g., Figs 5, 38, 39). Yet, in the females, the arma-
ture 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 conrmation that
the eyes are of similar size in both sexes as well as if the dierences in the armature of
the fore legs are part of sexual dimorphism or a part of individual variation in Quasita-
galis afonsoi would be better evaluated through examining more specimens.
Additional material examined. Quasitagalis afonsoi gen. et sp. nov. Brazil, To -
cantins, 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, 1.049m, 01 female,
09.xii.1997 (CTJMSB, 862).
Remarks on nymphs. Two nymphs of Quasitagalis afonsoi were collected with
the adults. ey were from dierent and undetermined instars and were not in good
condition for formal or complete descriptions. However, the head of the smaller (ear-
lier instar) nymph and fore legs of both of them were well conserved and were used to
obtain SEM images (Figs 52–59). eir measurements are given in Table 2.
ey 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). e arma-
ture 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
Figures 18–20. 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).
Quasitagalis afonsoi a new genus and a new species of Saicinae 23
Figures 21–24. Quasitagalis afonsoi gen. et sp. nov., female paratypes 21 lateral view 22 head and pro-
notum, dorsal view 23, 24 head and anteroinferior portion of prothorax 23 lateral view 24 posterolateral
view. Scale bars: 2.0 mm (21); 0.5 mm (22); 0.1 mm (23, 24).
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
24
Table 2. 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
basal to midpoint and on swollen portion, respectively (Figs 52, 53). e 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). e
armature of the fore trochanter is similar, but with small dierences: 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). e 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. e armature
of the fore tibia, however, is conspicuously dierent from that of the adults, with a
submedian (somewhat lateral to the median portion of the segment) row of six (earlier
Quasitagalis afonsoi a new genus and a new species of Saicinae 25
Figures 25–32. Quasitagalis afonsoi gen. et sp. nov., female paratypes 25 upper portion of head and pro-
notum 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).
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 setiger-
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
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Figures 33–37. 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).
ous spines (Figs 54–56); at the apical portion of the dorsal surface, two or three small
curved spines (Figs 54, 56, 57).
Habitat. e 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. e climate in
Quasitagalis afonsoi a new genus and a new species of Saicinae 27
Figures 38, 39. Quasitagalis afonsoi gen. et sp. nov., female paratypes 38, 39 fore leg, inner face, lateral
view 39 setae omitted, except the longer setae of femur, pointed by arrows. Scale bars: 0.5 mm.
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
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Figures 40–47. 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 a new genus and a new species of Saicinae 29
Figures 48–51. 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
51genitalia, posterior view. Scale bars: 0.2 mm (48); 0.1 mm (51); 0.05 mm (50); 0.02 mm (49).
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). e predomi-
nant 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).
e carbonates present in the region are located in the Speleological Province of
the Bambuí Group (São Domingos District) and distributed in a north-south direc-
tion. At this portion, elevations vary between 400 and 600 m, while elevations below
400 m dominate the northwestern portion of the study area. e Bambuí Group con-
stitutes 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 ocial government
databases (CECAV 2019), with approximately 350 known caves in the southeastern region.
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Figures 52–54. Quasitagalis afonsoi gen. et sp. nov., earlier instar nymph 52–54 lateral view 52 head
53labium 54 fore leg, inner face, the setae point to the larger setigerous spines on inner surface of fore
tibiae, close to its dorsal surface. Scale bars: 0.1 mm.
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,
Quasitagalis afonsoi a new genus and a new species of Saicinae 31
Figures 55–59. Quasitagalis afonsoi gen. et sp. nov., later instar nymph, fore leg, lateral view 55, 56in-
ner 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 ar-
rows) is inserted; adjacent long submedian spine pointed by a white arrow. Scale bars: 0.2 mm (55, 56);
0.04mm (57) 0.02 mm (58, 59).
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
32
Figures 60–63. 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
identied 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.
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. e
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. ey have a
predominantly at oor, and few have underground lakes or rivers.
Quasitagalis afonsoi a new genus and a new species of Saicinae 33
Figures 64, 65. Tagalis inornata inornata Stål, 1860, female, fore leg, lateral view, inner face; the setae
point to the larger setigerous spines on inner surface of fore tibiae, close to its dorsal surface 64 entire leg
65 portion of femur and tibia. Scale bars: 0.1 mm.
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34
Discussion
Gil-Santana et al. (2010) argued that the revised description of Tagalis when compared
to the one of Paratagalis Monte, 1943 (Gil-Santana and Costa 2009) showed many
similarities, suggesting that they were closely related genera, what was reinforced by
Gil-Santana (2011), while studying additional species of Tagalis. A similar situation is
revealed in relation to Tagalis and Quasitagalis, both sharing many common character-
istics, suggesting that they are also closely related genera.
It is noteworthy that, while Paratagalis and Quasitagalis have in common the pres-
ence of two pairs of setigerous spines on the gula (Figs 3, 23, 24), all other dierential
characteristics of the latter genus presented in relation to Tagalis, are also valid as such
in relation to Paratagalis. On the other hand, some additional characteristics of Qua-
sitagalis afonsoi sp. nov. were not recorded in any species of Paratagalis or Tagalis so
far. ose are the additional row of spines (intermediate between those of the upper
margin and ventral margin of the inner face) of the fore femora in the females (Figs 38,
39, 43) and the shape of parameres in the male (Fig. 13).
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 10–12, 14–20) have many similarities
to what was recorded in other species of Tagalis (e.g., Gil-Santana et al. 2010; Gil-San-
tana 2011). However, while in some species of the latter genus, the pygophore showed
a conspicuous lateral apophysis, it was not recorded in Q. afonsoi.
More importantly, in all species of Paratagalis and Tagalis in which the paramere
was described (Blinn 2008; Melo 2008; Gil-Santana and Costa 2009; Gil-Santana et
al. 2010; Gil-Santana 2011; Castro-Huertas and Forero 2014; Varela and Melo 2017),
it showed to be elongated, strongly curved apically, or with an apical large teeth, which
was implanted laterally, forming a right to an acute angle with the body of the paramere,
while in Q. afonsoi, the paramere is short, rounded at apex, in which the strong apical
acute spine is implanted in the same direction of the body of the paramere (Fig. 13).
e presence of the scopula on the apex of the third tarsomere on all the legs in Q.
afonsoi (Fig. 50) is in accordance with the statements of Weirauch and Forero (2007a)
and Weirauch (2008a) that this character seems to be widespread among Saicinae and
is a synapomorphy of part of this subfamily, respectively. As such, it has been recorded
in species belonging to several genera of Saicinae (Weirauch 2007; Weirauch and Fore-
ro 2007a, b), including two species of Tagalis (Castro-Huertas and Forero 2014), genus
which seems to be closer to Quasitagalis
e 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 54–56, 59) are striking,
because they are very similar to those recorded in almost all species of Tagalis (Figs 64,
65) (T. femorata Melo, 2008 has four spines) and also in Paratagalis (Gil-Santana and
Costa 2009), while they are completely absent in the adults of the new species (Figs 5,
38, 39, 48, 49). Similarly, Gil-Santana et al. (2010) recorded features in the nymphs of
T. evavilmae not present in adults of Tagalis but observed in adults of other related gen-
era, what they argued might help to understand the relationships among the genera of
Quasitagalis afonsoi a new genus and a new species of Saicinae 35
Saicinae in future studies. It is noteworthy that the mentioned features in the nymphs
of T. evavilmae included two pairs of spines on the ventral side of the head as present
both in adults and nymphs of Q. afonsoi sp. nov. (Fig. 52) and ve (T. evavilmae) to six
(Q. afonsoi) (Fig. 54) strong long spines on the inner face of the fore tibiae recorded in
the nymphs of earlier stage of both species.
Future phylogenetic analyses should be carried out to assess all these similarities
and dierences 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.,
Wygodzinsky 1966; Weirauch 2008a; Schuh and Weirauch 2020), are occasionally
found or consistently live in caves (Wygodzinsky 1966; Pape 2013; Gil-Santana and
Ferreira 2017). On the other hand, as far as it seems, this is the rst record of a spe-
cies of Saicinae, at least in the New World, found inside or as a possible inhabitant
of caves. e presence of a male, females and immature forms make believe that the
presence of Q. afonsoi inside the cave was not an incidental nding, but more prob-
ably their life cycle was, at least, partially being carried out there, possibly the species
might be reproducing in such habitats, as it occurs with some emesines (Wygodzinsky
1966; Pape 2013). However, more than a dozen caves were sampled in the region and
few specimens were found in only one of them (Gia’s Cave), which gives this species
a certain rarity. In any case, a more extensive study would be necessary, including
more frequent collecting and searches around the cave in the epigean environment to
determine the importance of caves as a habitat for Q. afonsoi. On the other hand, for
similar reasons, and also because of the fact that part of the surrounding landscape is
made up of pasture areas, which tend to extend, the distribution of the new species was
considered here strictly restricted to the Gia´s Cave (Figs 62, 63), which should be also
considered its type locality.
Key to the New World genera of Saicinae
Based on Weirauch and Forero (2007a, b), Gil-Santana and Costa (2009), and Gil-
Santana et al. (2015)
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; mesoscu-
tum (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
Hélcio R. Gil-Santana et al. / ZooKeys 966: 9–39 (2020)
36
4 Process on lower anterior angle of prothorax acute to subacute; second an-
tennal segment approximately half as long as rst 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 seg-
ment approximately 3/4 as long as rst 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 por-
tion 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 rst (visible) labial segments spiny; only apterous females known ..........
......................................................... Kiskeyana Weirauch & Forero, 2007
– Only the rst (visible) or all three labial segments spiny; females macropter-
ous ........................................................................................................... 10
10 Only the rst (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
Acknowledgments
We are grateful to João Paulo Sales Oliveira Correia (LNIRTT, IOC) for the photos
presented here as Figs 1 and 21. We would also like to thank the Central Scanning
Electron Microscopy Laboratory of the Faculty of Dentistry of the Araraquara Cam-
pus for the opportunity to use its Scanning Electron Microscopy. JO thanks Brazilian
CAPES (“Coordenação de Aperfeiçoamento de Pessoal de Nível Superior 001”) for
nancial support and Dr João Aristeu da Rosa for the support and for providing the
structure of the Parasitology Laboratory. RAZ thanks the Brazilian Society of Speleol-
ogy (Sociedade Brasileira de Espeleologia, SBE), Prefeitura de [City Hall of] Aurora
do Tocantins and the Espeleo Grupo [Group] Dolinas for logistical support during
Quasitagalis afonsoi a new genus and a new species of Saicinae 37
the eldwork, especially to Linda Gentry El Dash, Marcos Enoque de Leite Lima and
Sibele F. de O. Sanchez. We are also very grateful to Wanzhi Cai (China Agricultural
University, China), an anonymous reviewer, Laurence Livermore, and Nathalie Yonow
for their valuable comments and suggestions.
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