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Accepted by R. Constantino: 21 Jun. 2011; published: 15 Jul. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2967: 61–68 (2011)
www.mapress.com/zootaxa/Article
61
Key to the soldiers of Angularitermes Emerson with a new species from Brazilian
Amazonia (Isoptera: Termitidae: Nasutitermitinae)
TIAGO F. CARRIJO1,2,4, MAURICIO M. ROCHA1, CAROLINA CUEZZO3 & ELIANA M. CANCELLO1
1Museu de Zoologia da Universidade de São Paulo, Cx. Postal 42.494, 04218-970 São Paulo, SP, Brazil.
E-mail: tiagocarrijo@gmail.com; mrocha@usp.br; ecancell@usp.br
2Faculdade de Filosofia Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Av. Bandeirantes 3900, 14040-901,
Ribeirão Preto, SP, Brazil
3CONICET- Instituto Superior de Entomología “Dr. A. Willink”, Facultad de Ciencias Naturales e Instituto Miguel Lillo, UNT, Miguel
Lillo 205, T4000JFE, San Miguel de Tucumán, Argentina. E-mail: carolinacuezzo@csnat.unt.edu.ar
4Corresponding author. E-mail: tiagocarrijo@gmail.com
Abstract
An identification key based on characters of the soldier caste is provided for species of Angularitermes. Soldiers of pre-
viously described species in the genus, A. clypeatus, A. nasutissimus, A. orestes, A. pinocchio and A. tiguassu, are illus-
trated along with a new species, Angularitermes coninasus, n. sp., that is described and illustrated from soldier and worker
castes. Samples of the new species were collected from epigeal nests at the Brazilian Amazon rainforest. The soldier of
A. coninasus, n. sp. is distinguished from its congeners by having a short conical frontal tube, much wider at its base.
Key words: termites, taxonomy, Neotropical region, indentication key
Introduction
The Nasutitermitinae genus Angularitermes was described by Emerson (1925) as a new subgenus of Nasutitermes
Dudley, based on imago and soldier castes from the rain forest of Kartabo, Guyana (South America). Later, Snyder
(1949) raised Angularitermes to generic level and Mathews (1977) considered it as a senior synonym of Tintermes
Araujo.
Angularitermes is a well defined genus, whose soldiers are characterized by bilobed postclypeus, the intricate
microsculpture on the frontal tube base (Cancello et al. 1996) and the relatively large mandible “points”, while
workers have mandibles with reduced marginal teeth. The genus ranges from Trinidad to southeastern Brazil (Con-
stantino, 1998).
Five species were included in Angularitermes until now. A. nasutissimus Emerson 1925, the type-species, is
known from Amazonia and Trinidad. Emerson (1925) did not mention the microhabitat of the type material. One
sample from the Museu de Zoologia da Universidade de São Paulo (MZUSP) from Amapá, Brazil, has an observa-
tion made by the collector as “in rotten wood”.
A. clypeatus Mathews 1977 was described from Mato Grosso State and was found as an inquiline in nests of
Cornitermes silvestrii. A. orestes (Araujo 1970) shows a distribution in the Cerrado vegetation in central Brazil,
and was found by Araujo (1970) as an inquiline of nests of Armitermes euamignathus and by Mathews (1977) in
nest of Embiratermes neotenicus. A. pinocchio Cancello & Brandão 1996 is known from the Cerrado region in the
State of Goiás (Brazil), and also from the Atlantic Forest (Reis & Cancello, 2007); this species was found by D.
Brandão (Cancello et al. 1996) as an inquiline in nests of C. silvestrii. Finally, A. tiguassu Cancello & Brandão
1996 is known only from the State of Goiás, and was collected in the soil drawn up by the raised roots of a fallen
tree, in a semi-deciduous tropical forest (Cancello et al. 1996) and also found as an inquiline in nests of Corni-
termes cumulans by Costa et al. (2009).
CARRIJO ET AL.
62 · Zootaxa 2967 © 2011 Magnolia Press
Herein, Angularitermes coninasus, n. sp., from Brazilian Amazonia, is described and illustrated, based on
characters of soldier and worker castes. Also, for the first time an epigeal nest is characterized for a species of
Angularitermes. A key for the identification of the soldiers of the six species of Angularitermes is provided.
Material and methods
The examined material is deposited in the Isoptera collection of the Museu de Zoologia da Universidade de São
Paulo (MZUSP) and comprises the following samples of each species: A. nasutissimus: 3460, paratype (Guyana,
Kartabo; A.E. Emerson coll.; 11.viii.1920), and 9433 (Brazil, Amapá, Macapá; R. Constantino coll.; 28.x.1989); A.
clypeatus: 7388, holotype (Brazil, Mato Grosso, 12o49’S 51o46’W; A.G.A. Mathews coll.; 11.i.1968); A. orestes:
4708, holotype (Brazil, Minas Gerais, Serra do Curral; R.L. Araujo coll.; 2.xii.1956); A. pinocchio: 10040, holo-
type (Brazil, Goiás, Goiânia; D. Brandão coll.; x.1987); A. tiguassu: 10041, holotype (Brazil, Goiás, Goiânia; D.
Brandão coll.; x.1987) and two samples of the new species described herein (9425 and 13445, detailed below).
The morphometric characters used in this paper and their correspondence with Roonwal’s system (Roonwal 1970)
are indicated in parentheses when possible, as follows: LT, length of hind tibia (85); maximum width of head (17,
Fig. 1D), LN, length of frontal tube (figure 1A); LH, length of head to lateral base of mandibles (5, Fig. 1B); WN,
maximum width of frontal tube (Fig. 1C); and LN/WN, LN/LH, and LH/LT indices. All measurements, provided in
millimeters, were taken with a micrometric reticule adapted to a binocular stereomicroscope. The number of speci-
mens measured for each caste of Angularitermes coninasus n. sp. is provide in descriptions; also three soldiers of
A. nasutissimus, three of A. tiguassu and the type of A. clypeatus had the LN/WN index measured to be used into
the key.
Line drawings were prepared with a camera lucida coupled to a stereomicroscope and then scanned. Photo-
graphs of soldiers were taken with a digital camera adapted to a stereomicroscope at different focal points of each
individual and then combined using Combine ZP software (Hadley 2010). The worker enteric valve was mounted
on a microscope slide with alcohol gel and photographed under an optic microscope Axio Imager A1 and digital
camera ICc 1, Zeiss.
FIGURE 1. Morphometric characters of soldier head. In lateral view (left) and in dorsal view (right): A, length of frontal tube;
B, length of head to lateral base of mandibles; C, maximum width of frontal tube; and D, maximum width of head.
The terms used for pilosity are comparative: “thick bristles” are erect hairs with well-marked bases and thicker
than the “bristles”; bristles are thinner than the first (see the differences between them comparing the bristles in the
soldier head: figures 2A—A. coninasus n. sp. with 2D—A. tiguassu); “hairs” are shorter than bristles, less rigid
and with inconspicuous bases; “microscopic hairs” are very short and only visible under high magnification, usu-
ally 50× or more. The term epicranial suture is used as explained by Grassé (1982, p. 7). The terminology adopted
for the worker mandibles follows Fontes (1987), and the terminology for the digestive tube is after Noirot (2001).
Zootaxa 2967 © 2011 Magnolia Press · 63
KEY TO ANGULARITERMES, WITH A NEW SPECIES
Key to species of Angularitermes
1. Frontal tube very conical with a wide base, almost half of the length of frontal tube (Fig. 2A) . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. coninasus Carrijo & Rocha, n. sp.
- Frontal tube longer, not as conical as above, width of base much shorter than half of the length of frontal tube (Figs. 2B–F) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Short hairs on frontal tube, when present, only in the first basal third. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Short hairs along the length of frontal tube. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
3. Postclypeus very large and projected forward; length of frontal tube proportionally longer compared to cephalic capsule size
(Fig. 2E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pinocchio Cancello & Brandão
- Postclypeus smaller; length of frontal tube proportionally shorter compared to cephalic capsule size (Fig. 2C) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. orestes (Araujo)
4. Antennae with 14 articles; ratio LN/WH more than 1.85 (Figs. 2B and 2F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Antennae with 15 articles; ratio LN/WH less than 1.55; (Figs. 2D). . . . . . . . . . . . . . . . . . . . . . A. tiguassu Cancello & Brandão
5. Postclypeus very prominent and upward oriented (Fig. 2F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. clypeatus Mathews
- Postclypeus short (Fig. 2B). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nasutissimus Emerson
FIGURE 2. Angularitermes soldier head, in profile: A, A. coninasus n. sp. (paratype); B, A. nasutissimus; C, A. orestes (para-
type); D, A. tiguassu (holotype); E, A. pinocchio (paratype); and F, A. clypeatus (holotype).
CARRIJO ET AL.
64 · Zootaxa 2967 © 2011 Magnolia Press
Angularitermes coninasus Carrijo & Rocha, new species
Holotype. Soldier, part of lot MZUSP 13445, Brasil, Rondônia, Porto Velho, Jaci-Paraná, Ilha da Pedra module
from the Fauna Monitoring Program of Santo Antônio UHE, transect two, 3850 meters, 09o09’S 64o35’W; T. F.
Carrijo & R. G. Santos coll.; 14.ix.2010.
Paratypes. The remain specimens from holotype’s sample (workes and soldies) and Brazil, Amazonas,
Humaitá, 13.ix.1990, R. Constantino coll. (MZUSP 9425), part of the sample from the Museu Paraense Emílio
Goeldi, (MPEG 3680).
Etymology. The specific epithet coninasus is a combination of the Latin term conus (meaning cone) and fron-
tal tube as a reference to the conical shape of the soldier frontal tube.
Description. Imago. Unknown.
Soldier (Figs. 2A and 3). Monomorphic. Head capsule pyriform in dorsal view. frontal tube conical, with a
slight to strong constriction at the base (Fig. 3A), pointing upward in profile. Postclypeus well-developed; pro-
jected upward and forward (Fig. 3B). Mandibular blade reduced to conspicuous “points”. Labrum shorter than
wider; anterior margin of labrum rounded. Antennae with 14 articles; 1st article is the largest with at least twice the
length of 2nd; 3rd longer than 2nd; from 4th to 7th or 8th slightly longer than the anterior; from the 8th to the last, one
shorter than the next, the last is as long as the 2nd. Anterior lobe of pronotum trapezoidal, almost rectangular, with
anterior margin slightly emarginated. Legs very long; tibiae longer than head capsule length. Frontal tube with a
microsculpturing of irregular pits concentrated at the first half, some pits may reach near the apex. Surface of the
head capsule covered by thick bristles and hairs. Frontal tube surface with a few sparse bristles and a dense layer of
decumbent short hairs; frontal tube apex with four rigid bristles smaller than those of head. Labrum glabrous. Bris-
tles of the antennae concentrated at the apex of each article with short hairs on their surface; first article with bris-
tles restricted to the apex. Pronotum with bristles on margins and two or three over its surface. Meso- and
metanotum with many thick bristles on lateral and posterior margins. Abdominal tergites covered by many thick
bristles of different sizes. Abdominal sternites with a line of thick, erected bristles and sparse short hairs oriented
backward. Legs with many thick bristles and hairs over their surfaces. Tibial spurs 2:2:2. Head capsule varying
from ferruginous orange to pale yellow, frontal tube brown/red, always darker than the head capsule. Body yellow
to pale yellow. Measurements of four soldiers from MZUSP 9425 and four from MZUSP 13445: LN (1.38–1.44),
LH (1.32–1.38), WN (0.61–0.67), WH (1.17–1.25), LT (2.27–2.40), LN/WH (1.11–1.18), LN/ LH (1.00–1.05),
LH/LT (0.57–0.60).
FIGURES 3. Angularitermes coninasus n. sp.: 3A, soldier head capsule, dorsal view. 3B, postclypeus, frontal view.
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KEY TO ANGULARITERMES, WITH A NEW SPECIES
FIGURES 4–6. Worker of Angularitermes coninasus n. sp.: 4, mandibles; 5, digestive tube in situ: A, dorsal; B, right; C, ven-
tral; and D, left views; 6, detail of mesenteric tongue and Malpighian tubules: A, profile; and B, internal views.
Worker (Figs. 4–8). Dimorphic. Differences among the dimorphic workers are restricted to their size. Head
capsule trapezoidal, fontanelle and epicranial suture inconspicuous. Antennae with 15 articles, if 14 the 3rd is
always subdivided. Postclypeus inflated. Anterior lobe of pronotum slightly emarginated. Tibial spurs 2:2:2. Head
capsule with many bristles. Pronotum with bristles on the anterior and posterior margins; meso- and metanotum
with bristles on posterior margins; abdominal tergites covered with many bristles and hairs; sternites with bristles
and a dense layer of short hairs. Legs covered by scattered bristles and hairs. Measurements of four major and four
minor workers from MZUSP 9425 and MZUSP 13445 are given as a range: major, WH (1.32–1.36), LT (1.87–
1.96); and minor, WH (1.17–1.19), LT (1.64–1.74).
CARRIJO ET AL.
66 · Zootaxa 2967 © 2011 Magnolia Press
FIGURE 7. Angularitermes coninasus n. sp.: A, gizzard; and B, detail of the pulvillus I. f1, f2 and f3: first, second and third
order folds, respectively, p1 and p2: pulvillus I and II respectively; and arrow: spines on the folds.
FIGURE 8. Enteric valve armature of the worker of Angularitermes coninasus n. sp., showing major cushions (large arrows)
intercalated by minor cushions (thinner arrows). Observation: when enteric valve was mounted major cushions adopted differ-
ent positions but there are no differences between them.
Mandibles (Fig. 4). Left mandible: apical tooth slightly larger than M1+2; angle between apical tooth and M1+2
tooth nearly acute; M3 with half size of M1+2; notable gap between M3 and molar prominence; molar tooth apex hid-
den by molar prominence; molar prominence very concave and without ridges. Right mandible: apical tooth
approximately twice the size of M1; M2 reduced or absent; molar plate very concave and without ridges.
Digestive tube (Figs. 5–8). Crop well developed. Gizzard armature complete, surface of folds I and II orna-
mented with small and sparse spines (Fig. 7A); pulvillus I well developed, covered with long aciculiform spines on
the superior portion and many lines with very small spines (4–6) on the inferior portion (Fig. 7B), pulvillus II
reduced, surface smooth. Anterior end of midgut with a distinct hump dorsal to the insertion of the gizzard. Mixed
segment very short, mesenteric tongue external to mesenteric arch (Fig. 6A). Malpighian tubules attached individ-
Zootaxa 2967 © 2011 Magnolia Press · 67
KEY TO ANGULARITERMES, WITH A NEW SPECIES
ually but closely adjacent in pairs at mesenteron-proctodeum junction; each tubule conspicuously dilated in its
proximal part (Fig. 6A and B). Proctodeal segment (P1) short and tubular throughout its length. Enteric valve (P2)
visible at left side (Fig. 5B). P2 with three major cushions intercalated with three minor ones (Fig. 8): major cush-
ions with shorter spines on its surface and, after a deflection on the distal portion, a row of 4–6 larger spines point-
ing inward to gut lumen (Fig. 8, large arrows); minor cushions with only sparse small spines (Fig. 8, thinner
arrows). Paunch (P3) with P3a and P3b dilated; proximal part of colon (P4a) dilated; U-turn short and visible at right
view.
Biology. All samples were collected in a primary rainforest. A. coninasus n. sp. type material was sorted out of
a domed shaped epigeal nest, measuring 29 cm in height and 55 cm in width at its base (Fig. 9). The surface of the
nest was covered by loose soil, which also formed the peripherical layer of about 5 cm in depth with noticeable gal-
leries larger than 1 cm in diameter. Galleries at the internal layer had 0.5 cm diameter and were occupied by inter-
twined roots extending almost 10 cm down in soil. There were no chimneys communicating interior with exterior.
The nest was surrounded by a shallow litter layer and some sticks. Neither inquilines nor real pair were found
within A. coninasus n. sp. nest.
FIGURE 9. Nest of Angularitermes coninasus n. sp. from Rondônia, Brazil.
Comparisons. Soldiers of Angularitermes coninasus n. sp., can be easily recognized among its congeners by
having a short and strongly conical frontal tube, with a wide base. The combination of characters as microsculpture
of the head capsule and frontal tube, pilosity and coloration distinguish A. coninasus n. sp. from the other species.
A. tiguassu is morphologically similar to the new species but has a much narrower frontal tube at the base; anten-
nae with 15 articles and LH/LT index 0.66–0.68. Workers are dimorphic. No other species of Angularitermes was
reported as nest-building.
Acknowledgements
We are grateful to “PROBIOTA Consultoria Ambiental and Santo Antônio Energia” for the execution of the Ento-
mological Monitoring Program from Santo Antônio UHE and for field support. We thank Rodrigo B. Gonçalves,
Gláucia Marconato and Rodrigo Feitosa for reading an earlier draft of this paper and offering helpful advice and
criticism; and Rafaella G. Santos who helped the first author in collecting the type material of A. coninasus. This
work was partially supported by a doctoral scholarship from Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior, CAPES, Brazil, to T. Carrijo and M. M. Rocha; a grant from Conselho Nacional de Desenvolvimento
Científico e Tecnológico (Proc. CNPq No. 304765/2007-4), Brazil, to E. M. Cancello; and a doctoral scholarship
from CONICET, Argentina, to C. Cuezzo.
CARRIJO ET AL.
68 · Zootaxa 2967 © 2011 Magnolia Press
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