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Studies on Neotropical Fauna and Environment
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A new species of Epectinaspis Blanchard
(Coleoptera: Scarabaeidae: Rutelinae) from
Guatemala and rediscovery of Balanogonia
Andrés Ramírez Ponce , Leonardo Delgado & Yoshua Soto
To cite this article: Andrés Ramírez Ponce , Leonardo Delgado & Yoshua Soto (2021): A new
species of Epectinaspis Blanchard (Coleoptera: Scarabaeidae: Rutelinae) from Guatemala
and rediscovery of Balanogonia�constricta Paucar-Cabrera, Studies on Neotropical Fauna and
Environment, DOI: 10.1080/01650521.2020.1859874
To link to this article: https://doi.org/10.1080/01650521.2020.1859874
Published online: 01 Jan 2021.
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A new species of Epectinaspis Blanchard (Coleoptera: Scarabaeidae: Rutelinae)
from Guatemala and rediscovery of Balanogonia constricta Paucar-Cabrera
Andrés Ramírez Ponce
, Leonardo Delgado
and Yoshua Soto
CONACYT—Instituto de Biología. Laboratorio Regional de Biodiversidad y Cultivo de Tejidos Vegetales, Ex Fábrica San Manuel de Morcóm,
Universidad Nacional Autónoma de México, Santa Cruz Tlaxcala, Mexico;
Red de Biodiversidad y Sistemática, Instituto de Ecología A. C.,
Instituto Tecnológico Superior de Zacapoaxtla, Carretera Acuaco-Zacapoaxtla Km. 8, , Zacapoaxtla, Mexico
Epectinaspis canoi new species is described and illustrated, and its similarities with the other
species of the genus are discussed. A modification of the taxonomic key of this genus is provided.
In addition, Balanogonia constricta Paucar-Cabrera is rediscovered and illustrated with data on its
geographical distribution and habits.
Received 19 August 2020
Accepted 1 December 2020
Anomalini; Central America;
The genus Epectinaspis Blanchard (Coleoptera: Scarabaeidae:
Rutelinae: Anomalini) is a small group of diurnal phytopha-
gous beetles that includes 10 species distributed from central
and southern Mexico to the north of Venezuela (Paucar-
Cabrera 2003; Morón & Ramírez-Ponce 2012; Ramírez-
Ponce & Curoe 2014). The species of this genus occur in
montane forest, humid oak forest, deciduous tropical forest,
coffee plantations and secondary forest (Paucar-Cabrera
2003), with the highest species richness in southern Mexico
(six species), and Guatemala (five species).
Another genus of Anomalini that occurs in Mexico
is Balanogonia Paucar-Cabrera. This genus, including
two species, is endemic to Mexico and was previously
confused with Epectinaspis (Paucar-Cabrera 2003;
Jameson et al. 2003; Ramírez-Ponce & Curoe 2014).
One of the species of Balanogonia is B. constricta
Paucar-Cabrera, described with a single male, and
without precise locality, since this specimen was col-
lected at beginning of the previous century.
The purposes of this work are to describe a new
species of Epectinaspis from Guatemala, and to provide
data on the geographical distribution and natural his-
tory of Balanogonia constricta Paucar-Cabrera.
Material and methods
The morphological characters, terms, and criteria used are
those of Paucar-Cabrera (2003) and Ramírez-Ponce and
Curoe (2014). All specimens and characters were observed
with a Carl Zeiss Stemi 305 stereomicroscope (Jena,
Germany) with a Pǀ10×/23 eyepiece. Measurements were
obtained with an ocular micrometer and photographs
were taken with a Carl Zeiss Axio Zoom V.16 multifocal
stereomicroscope with Pǀ10×/23 eyepieces, a Plan Neo
Fluar Z (Jena, Germany) 1×/0.25, FWD 56 mm objective
and an associated Axiocam 506 color camera (Jena,
Germany). Images were processed and edited in Adobe
Photoshop CS5 (San José, CA, USA). The map was cre-
ated by the SimpleMappr website (Shorthouse 2010). The
textual data for the labels is quoted in quotation marks,
and subsequent letters in closed parentheses indicates
The specimens studied are deposited in the following
collections: Andrés Ramírez Ponce Collection, Tlaxcala,
Mexico (ARPC), Arthropod collection of the Universidad
del Valle de Guatemala, Guatemala City, Guatemala
(UVGC), Carsten Zorn Collection, Gnoien, Rostock,
Germany (CCZ), Colección de Coleoptera, Universidad
Autónoma del Estado de Hidalgo, Pachuca, Mexico (CC-
UAEH), Daniel Curoe Collection, Mexico City, Mexico
(DDC), José Luis Huerta Collection, Xalapa, Veracruz,
Mexico (JLSH-XAL), Leccinum García-Morales collec-
tion, Ciudad Victoria, Tamaulipas, Mexico (CLGM),
Leonardo Delgado Collection, Xalapa, Veracruz, Mexico
(LLDC), Matthias Seidel Collection, Prague, Czech
Republic (MSPC), Naturhistorisches Museum Basel,
Basel, Switzerland (NMB), Oscar Galindo Collection,
CONTACT Andrés Ramírez Ponce email@example.com
This article has been republished with minor changes. These changes do not impact the academic content of the article.
STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT
© 2020 Informa UK Limited, trading as Taylor & Francis Group
Published online 01 Jan 2021
Texcoco, Mexico (GALC), Fernando Zagury Vaz de
Mello, Universidade Federal de Mato Grosso, Cuiabà,
Brazil (CEMT) and Yoshua Soto Collection, Puebla,
Mexico (EUSY). The holotype of Balanogonia constricta
Paucar-Cabrera was revised in the Georg Frey Collection,
at the Naturhistorisches Museum Basel, Switzerland
Epectinaspis canoi Ramírez-Ponce & Delgado, new
(Figures 1a–h, 2b, 5)
Type material. Holotype male labeled: a) “GUATE-
MALA: A.V. 20 km al N de Sn Pedro Carchá Fco:
Chamtaca 1200 m. 10 viii 1991. H. Krees (manta gol-
peteo)”; b) “En cafetal”; c) “Col.: B. Jolae Terrestres”; d)
“HOLOTYPE ♂ Epectinaspis canoi Ramírez-Ponce
et al., 2020.” Holotype deposited in UVGC.
Diagnosis. Dorsal surface glabrous, shining black; prono-
tum weakly convex in lateral view; elytron with 10 striae,
epipleural expansion extends to the second visible ventrite;
male genitalia with parameres distinctive (Figure 1g, h).
Description of holotype. Length 8.83 mm; width
4.68 mm. Color: Completely shiny black. Head (Figure
1a): Interocular width equals 6.4 transverse eye diameters.
Clypeus with apex widely rounded, weakly reflexed; sur-
face rugopunctate, punctures smaller and rounded toward
middle frontoclypeal disc; frontoclypeal disc raised.
Maxilla with lacinia setose, terminal palpomerus as long
as 3 basal segments, and 1.5 times wider than 3 basal
segments. Prementum surface irregular, punctures shape-
less; scarcely setosus, setae long, leaning forward; mentum
surface smoother than prementum, punctures not defined;
long setae only near of the outer edge; disc almost glabrous.
Pronotum (Figure 1b): Lateral and posterior margins
beaded; posterior angles in right angle. Surface weakly
convex (Figures 1c, 2a), disc uniformly punctate, punctures
small, rounded, and shallow, not contiguous, at sides twice
the size than those of the disc. Scutellum: Slightly wider
than long (1:0.8), shape subtriangular. Surface punctate,
punctures as in the disc of pronotum. Elytron (Figure 1b,
1c): Surface with 10 punctate striae; 1 next to elytral suture
well defined, 5 on disc reaching apex and base (the second
not clearly defined in the middle), 1 on the humerus, 2
laterad of humerus, and 1 along lateral margin; punctures
ocellate, moderately in size. Apex of suture spiniform.
Intervals 1–6 subequal in width, (4 times width of
stria 1), 7–9 broad (4 times width of stria). Elytral margin
with only 1 or 2 small setae and epypleura remarkably
developed, reaches the end of the second visible sternite
(Figure 1c). Propygidium: Apex partially exposed, surface
punctate, punctures elongated, not closed and sparse on
anterior half, becoming strigate toward apex. Pygidium:
Twice as long as the length of propygidium. Shape sub-
triangular in caudal view, evenly convex in lateral view.
Surface strigate, with 8–10 small disperse setae on each
side. Venter (Figure 1d): Sternites 2–4 subequal in length,
0.55 times length of sternite 5; sternites 2–6 irregularly
punctate, and with a transverse row of sparse setae on
the first half; last sternite with a row of setae on the edge.
Legs: Protibia with 2 teeth and subapical spur (Figure 1e),
spur 0.8 times length of protarsomere 1. Protarsomere 5 as
long as protarsomeres 1–4 combined, with internomedial
protuberance absent; inner claw with internal ramus 3
times wider than external ramus (Figure 1f). Mesotibia at
apex with 7 spinules; external edge with developed, oblique
carina at middle with 6 short spines. Mesotarsal claw split,
ventral ramus 2.3 times wider than dorsal ramus. Metatibia
apex with 10 spinules, external edge with developed, obli-
que carina at middle with 6 short spines. Parameres: In
lateral view wide and round at the base, drastically slim-
ming toward the apex; apex remarkably curved, hook-like,
sharp (Figure 1g); in frontal view noticeably and gradually
thinned to its middle part, then markedly thickened, with
the edges of the thickest area projected externally, spine-
like, and rounded and progressively little thinned toward
the apex; apex widely rounded (Figure 1h).
Etymology. This species is named after Enio Cano
(Universidad del Valle de Guatemala), enthusiastic ento-
mologist interested in the study of the diversity and
conservation of Guatemala, and author of several pub-
lications on the families Passalidae and Scarabaeidae.
Distribution. This species is known only by the holotype
collected in the central part of Guatemala (20 km N Sn
Pedro Carchá), in an area of coffee crops (Figure 5).
Remarks. This species is similar to E. chalconota Bates.
Differs by the pronotum weakly convex (Figures 1c, 2a)
(strongly convex in E. chalconota; Figure 2b), the interno-
medial protuberance of the protarsomere 5 hardly per-
ceptible (well-developed interno-medial protuberance in
E. chalconota), and the genitalia with parameres
2A. R. PONCE ET AL.
Figure 1. Habitus and morphological structures. Epectinaspis canoi sp. nov., holotype: (a) head; (b) habitus in dorsal view; (c) habitus
in lateral view; (d) habitus in ventral view; (e) foretibia; (f) foretarsus; (g) male genitalia in lateral view; (h) parameres in fronto-distal
STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT 3
strongly narrowed and abruptly expanded in distal
view, with outer edge toward apex entire, broadly
rounded (continuous with a lateral protuberance,
and outer edge toward apex sinuated, with tightly
rounded in E. chalconota).
Modification to the key of Ramírez-Ponce and Curoe
(2014) for the males of the species of Epectinaspis
1. Epipleural expansion extends from base of ely-
tron to middle of metacoxa. Frons with disc
convex or flat ......................................................... 3
Epipleural expansion extends from base of ely-
tron to ﬁrst or second abdominal sternite. Frons
disc ﬂat to weakly concave .................................. 4
2. Frons with disc setose. Pronotum black, rugo-
punctate (punctures dense), and densely setose.
Pronotum and venter with red setae. Genitalia
with parameres setose; in dorsal view weakly cur-
ving inward to rounded apices; in side view
weakly curved ventrally. Guatemala and Belize . . .
...................................................... E. ambigens Bates
Frons with disc glabrous. Pronotum testaceous,
with or without castaneous macula on middle,
punctate (punctures moderately dense). Pronotum
glabrous or moderately setose. Pronotum and ven-
ter with tawny setae. Genitalia with parameres glab-
rous; in dorsal view, not rounded nor curved, in
lateral view, not bent downward .......................... 11
3. Length of clypeus in ventral view about 1/2
length of mentum. Genitalia strongly bent at
phallobase–tectum joint (side view); tectum
short; parameres (dorsal view) narrowing to
rounded apices, with preapical constrictions; in
side view, dorsal margin broadly rounded to
acute, recurved apices. Mexico . . . . . . . . . . . . . . . . . .
........................................ E. mexicana (Burmeister)
Length of clypeus in ventral view about 1/3 to 1/4
length of mentum. Genitalia not strongly bent at
phallobase–tectum joint (side view) ................... 5
4. Pronotum with setae on entire disc, at apical
angles or at apical angles and margins . . . . . .... 6
Pronotum glabrous (with or without setae at lateral
margins) or with 1–10 setae either side of disc .. 7
5. Elytron with ﬁrst 3 intervals punctate in basal
half. Base of clypeus in lateral view with weak
protuberance. Elytra with opaque surface.
Genitalia with tectum long, thin (side view); para-
meres (dorsal view) inner margins touching along
entire length, narrowing to blunt apices, with
preapical constrictions; in side view, apices long,
acute, directed downward. México, Guatemala, El
Salvador, Nicaragua, Panama . . . . . . . . . . . . . . . . . .
...................................... E. moreletiana (Blanchard)
Elytron with impunctate intervals. Base of cly-
peus in lateral view without protuberance. Elytra
with shiny surface. Genitalia with tectum thick
(side view); parameres (dorsal view) inner mar-
gins overlapping 1/3 of length, strongly narrow-
ing to expanded apices; inside view, narrowing to
recurved apices. Mexico . . . . . . . . . . . . . . ................
..................................................... E. opacicollis Bates
6. Pronotum glabrous. Elytron with 10 striae . . . . . .
Pronotum with setae. Elytron not with 10 striae . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7. Pronotum strongly convex in lateral view. Genitalia
with parameres more or less continuous with
a lateral protuberance in fronto-distal view, and
outer edge toward apex sinuated, tightly rounded.
Figure 2. Pronotum in lateral view. (a) Epectinaspis canoi sp. nov. (Modified from Paucar-Cabrera 2003); (b) Epectinaspis chalconota
4A. R. PONCE ET AL.
Mexico and Guatemala. . . . . . . . . . . . . . . .
............................ ............................ E. chalconota Bates
Pronotum weakly convex in lateral view.
Genitalia with parameres strongly narrowed
and abruptly expanded (denticle-like) in
fronto-distal view, and outer edge toward
apex entire, broadly rounded. Guatemala. . . .
. . . . . . . . ............................................................................
E. canoi Ramírez-Ponce, Delgado & Soto, sp. n.
8. Elytron with 11–13 striae. Parameres with notch
in apical 1/3. Mexico . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . ................... E. pictipennis Bates
Elytron with 9 striae. Parameres without notch in
apical 1/3................................................................. 10
9. 10. Pronotum with about 10 whitish setae at
anterior sides of disc. Genitalia with parameres
(dorsal view) overlapping to middle, becoming
apically broad with narrow gap between them;
inner margins nearly straight; in side view, ven-
tral margin weakly convex, apices hook-like. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................
....................................... E. bisyrica Paucar-Cabrera
Pronotum with or without 1–8 tawny setae at
anterior sides of disc. Genitalia with parameres
(dorsal view) overlapping in basal third, becom-
ing apically slender with wide gap between them;
inner margins curved; in side view, ventral mar-
gin straight, apices hook-like . . . . . . . . . . . . . . . . . . .
...................... ..................... E. guatemalensis Ohaus
10. Frons with disc convex. Pronotum sparsely to
moderately setose, basal angles obtuse. Metatibial
apex with 12–15 spinules. Genitalia with para-
meres (dorsal view) inflexed outward and diver-
ging to reflexed apices; in side view margins
subangular, abruptly narrowing apically. Panama ..
........................................................... E. chelifera Bates
Frons with disc flat. Pronotum glabrous, basal
angles acute. Metatibial apex with 18–19 spinules.
Genitalia with parameres (dorsal view) with tips
straight and apically rounded; in side view margins
continuous, very broadly rounded, simple. Costa
Rica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...............
................ E. costaricensis Ramírez-Ponce & Curoe
Balanogonia constricta Paucar-Cabrera, 2003
(Figures 3a–d, 4, 5)
Holotype male labeled. a) “Truqui Mexico” (typewritten);
b) “Fry Coll. 1905–100” (typewritten); c) “Epectinaspis picti-
pennis Bates” (handwritten); d) “Balanogonia constricta
Paucar-Cabrera, HOLOTYPE, male symbol, det. A. Paucar-
Cabrera 2001” (typewritten). Holotype revised at MGFT.
The description of this species was made by Paucar-
Cabrera (2003) with only one single male specimen
from Mexico with no other data about the distribution
or natural history. Several male and female specimens
were collected in recent expeditions to the Sierra
Negra, Puebla, Mexico, and other specimens were
revised in other collections of Mexico, all of which
were determined as B. constricta by comparison with
the type, whose data are detailed below, as well as the
description of the female and the variation in males.
a) “MEX. Pue. Tlatlahuquitepec. Macuilquila, Río La Soledad.
Mesófilo. 13-iii-18. 720 m. 19.963105°, −97.471521° En C.
coelestinum (L.). Soto, Y.”; b) “Balanogonia constricta
Paucar-Cabrera 2003 Homotipo. Ramírez-Ponce, A. Det.
2019” (10 ♂♂; ARPC (7), CCZ (1), MSPC (1); a) “MÉX.
Pue. Tlatlauquitepec. Chicuaco, río La Soledad. 23/iii/2018.
730 m. Mesófilo. En flores de C. coelestinum (L.). Yoshua
Soto, col.”; b) “Balanogonia constricta Paucar-Cabrera
2003 Homotipo. Det. Ramírez-Ponce, A. 2019” (29 ♂♂,
3 ♀♀: ARPC (12 ♂♂, 3 ♀♀), CC-UAEH (1 ♂), CLGM (2
♂♂), DCC (2 ♂♂), JLSH-XAL (3 ♂♂), LLDC (2 ♂♂),
NMB (1 ♂), GALC (2 ♂♂), CEMT (2 ♂), EUSY (2 ♂♂); a)
“México: Puebla Tlatlauquitepec Macuilquila 800 m 24-VI
-17 Y. Esquivel”; b) “Balanogonia constricta Paucar-
Cabrera 2003. Homotipo Det. Ramírez-Ponce, A. 2019”
(1♂, 1♀; MXAL); a) “MÉXICO, PUEBLA Juan António
Rayón Jonotlán 12-03-2005 1308 ‘?’ m Colecta Directa
C. O. Tlapanco col”; b) “Balanogonia constricta Paucar-
Cabrera 2003. Homotipo Det. Ramírez-Ponce, A. 2019”
(1♀; MXAL); a) “MÉXICO, PUEBLA Barranca de Patla.
15-iii-2020. Galindo, O. Col.”; b) “Balanogonia constricta
Paucar-Cabrera, 2003. Homotipo Det. Ramírez-Ponce,
A. 2019” (1♂; GALC); a) “MÉXICO, PUEBLA Xicotepec
de Juárez, La Ceiba. 15-iii-2020. Galindo, O. Col.”; b)
“Balanogonia constricta Paucar-Cabrera, 2003. Homotipo
Det. Ramírez-Ponce, A. 2019” (1♂; GALC); a) “MÉXICO:
Hidalgo, Río Atlapezco 120 m. Selva baja pert. UTM
2324–439, 14Q0548842. Sobre plantas arbustivas. 15-IV
-2003. L. Toledo. J. Asiain y J. Márquez cols.”; b)
“Strigoderma teapensis Bates L. Delgado det. 2004”; c)
“CC-UAEH/SC438”; d) “Balanogonia constricta Paucar-
Cabrera 2003. Homotipo. Det. Ramírez-Ponce, A. 2019”
(1♂; CC-UAEH); a) same data; b) same data except
“CC-UAEH/SC440” (1♂; CC-UAEH); a) same data; b)
same data except “CC-UAEH/SC441” (1♂; CC-UAEH).
Description of female. Five specimens (Figure 3b).
Similar to the male, except in the following characters:
Length 9.50–10.5 mm; width 4.70–5.25 mm. Color:
STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT 5
Completely reddish brown or dark reddish brown with
intense greenish reflections on pronotum, or reddish
brown on pronotum and legs, and dark reddish brown
on elytra and venter, with greenish reflections on frons,
pronotum and elytra, more intense on venter. Head:
Clypeus subrectangular or subtrapezoidal, weakly reflexed;
surface rugopunctate. Frons rugopunctate, supraocular
area with 5–7 long erect setae, interocular width equals
8.45–8.60 transverse eye diameters. Pronotum: surface with
2–3 erect setae on each side of disc. Venter: Sternites 2–5
Figure 3. Balanogonia constricta Paucar-Cabrera. (a) Habitus of male; (b) habitus of female; (c) male genitalia in lateral view; (d)
male genitalia in fronto-distal view.
6A. R. PONCE ET AL.
Figure 4. Balanogonia constricta Paucar-Cabrera feeding on flowers (Picture Soto, 2018).
Figure 5. Distribution of Epectinaspis canoi sp. nov. (black circle), and Balanogonia constricta Paucar-Cabrera (red circles).
STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT 7
0.3 times longer. Legs: distal denticle long and curve out-
ward, protarsomere 1 longer than protarsomeres 2–4, pro-
tarsomere 5 subequal to protarsomeres 2–4. Inner
protarsal claw split; ventral ramus 1.2 times as dorsal
ramus, dorsal ramus longer. Mesotibia with 7–9 spinules
at apex; last spine thinner and twice as long as the pre-
vious; external edge with oblique carina with 7–8 spines,
the two located at the ends remarkably long. Metatibia
with 12–14 spinules at apex, oblique carina with 9 spines,
the last in the dorsal side, as long as 4 times than the
others. Female genital plates: triangular, abundantly setose
in the distal half, setae thin, long, decumbent.
Variation in males. Forty-six specimens (Figure 3a, c,
d). Length 6.23–9.23 mm; width 3.23–4.53 mm. Color:
two general patterns; a) head, apex of tibiae and tarsi
reddish brown; frons, disc of pronotum, venter and legs
(except metafemurs) coppery brown with intense green-
ish reflections (brighter and iridescent in pronotum);
elytra, borders of pronotum, disc of scutellum and
some parts of metafemurs yellowish, and laterad of
humerus and elytral margins dark brown; or b) body
completely reddish brown or dark reddish brown with
intense greenish reflections more intense and bright on
frons, pronotum, venter and femurs and tibiae. Clypeus
subquadrate or slightly subtrapezoidal, punctuation
moderately or strongly rugopunctate. Frons with
supraocular area with 4–7 long erect setae. Pronotum
surface punctate or semi rugopunctate, without or with
2–3 erect setae on each side of disc. Protibia with sub-
apical spur as long as the second protarsomere or
shorter; mesotibia with oblique carina with 7–9 spinules,
and apex with 8–10 spinules; metatibia with oblique
carina with 8–13 spinules, and apex with 14–16 spinules.
Biological data. Adults are active in the months of March
(46), April (3), June (2), and have been collected in shrub
vegetation and mainly in flowers of Conoclinium coelestinum
(Linnaeus) and Salix Linnaeus, in sites located on the river-
banks (Figure 4).
Distribution. This species inhabits the cloud mountain
forest of the Sierra Madre Oriental and tropical forest
in the coast of the Gulf of Mexico in the states of
Hidalgo and Puebla, located in regions between 150
and 1170 m in elevation (Figure 5).
We are especially grateful to Enio Cano (Universidad de San
Carlos de Guatemala) for the loan of the specimen of the new
species here described. The senior author expresses thanks to the
Consejo Nacional de Ciencia y Tecnología (CONACYT) for the
“Cátedras CONACYT” program and to the Instituto de Biología,
Universidad Nacional Autónoma de México, for the technical and
scientific support given to the project No. 59: “Laboratorio
Regional de Biodiversidad y Cultivo de Tejidos Vegetales
(LBCTV) del Instituto de Biología (IBUNAM), sede Tlaxcala”;
to CONACYT for the Laboratorios Nacionales CONACYT pro-
gram that allowed us to obtain microscopy equipment, and to the
Secretaría de Fomento Agropecuario del Estado de Tlaxcala
(SEFOA) for a support grant as part of the institutional collabora-
tion agreement, and to Jorge Cortés Flores (UNAM) for the
taxonomic determination of the plant specimen.
No potential conflict of interest was reported by the author(s).
Andrés Ramírez Ponce http://orcid.org/0000-0003-4742-
Jameson ML, Paucar-Cabrera A, Solís A. 2003. Synopsis of
the New World genera of Anomalini (Coleoptera:
Scarabaeidae: Rutelinae) and description of a new genus
from Costa Rica and Nicaragua. Ann Entomol Soc Am.
Morón MA, Ramírez-Ponce A. 2012. Mesoamerican genera
of Anomalini (Coleoptera: Melolonthidae: Rutelinae):
a brief review. Trends Entomol. 8:97–114.
Paucar-Cabrera A. 2003. Systematics and phylogeny of the
Genus Epectinaspis Blanchard (Coleoptera: Scarabaeidae:
Rutelinae) and description of a new genus of Anomalini
from Mexico. Coleopt Soc Monogr. 2:3–60.
Ramírez-Ponce A, Curoe D. 2014. Description of two new
species in the genera Epectinaspis Blanchard and Strigoderma
Burmeister (Coleoptera: Scarabaeidae: Rutelinae: Anomalini).
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8A. R. PONCE ET AL.