ISSN2477-9458
BOLETÍN DEL
CENTRO DE
INVESTIGACIONES
BIOLÓGICAS
Fenotipo de la resistencia a MLSB y la tipificación estructural
del cassette cromososmal mec (SCCmec) en Staphylococcus
aureus resistentes a meticilina procedentes de manos de
manipuladores de alimentos.
Víctor Pico-Bracho, Jhoandry Rivera-Salazar, Velina Aranaga-
Natera, Isabel Mujica de Fernández, Yoalaimis La Paz-Delgado e
Irene Zabala-Díaz…...……………………………………………….
1
Paracymus de Venezuela (Coleoptera: Hydrophilidae:
Laccobiini), Parte VII: Registro de seis nuevas especies.
Mauricio García Ramírez…….……..………………….......................
20
Influencia del régimen hidrológico sobre la composición de
sedimentos de manglares en la Bahía de El Tablazo (Sistema
de Maracaibo).
Flora Barboza, Ana Marta Francisco, Jacinto Sánchez y Ernesto
Medina………………………………………………………………………
45
Discovery of two new genera of detritivorous aquatic beetles
Toneroides, in the Venezuelan Amazon (Coleoptera:
Noteridae: Noterinae).
Mauricio García Ramírez…………….……………………….……..…....
67
Notas científicas.
Nuevas observaciones y ampliación del rango altitudinal del
gabán Mycteria americana (Linnaeus, 1758) en los andes
de Venezuela, sugieren desplazamientos entre
biorregiones.
Luis A. Saavedra, Alexis Araujo-Quintero y Carla I. Aranguren…..
112
Notes on the genera Suphisellus Crotch, 1873 and Suphisellus
Zimmermann, 1919, a cocktail of encrypted Suphiselloides
genera (Coleoptera: Noteridae: Noterinae: Noterini).
Mauricio García Ramírez…………………………..…………..……….
124
Instrucciones a los autores……………….…..…………………………
138
Instructions for authors………………….………………………….…
148
Vol. 58, N0 1, Pp. 1-157, Enero-Junio 2024
UNA REVISTA INTERNACIONAL DE BIOLOGÍA PUBLICADA
POR
LA UNIVERSIDAD DEL ZULIA, MARACAIBO, VENEZUELA
Boletín del Centro de Investigaciones Biológicas
Vol.58. Nº 1, Enero-Junio 2024, Pp. 67-111
67
http://zoobank.org/urna:lsid:zoobank.org:pub:9CAA4AD9-B59C-4D42-83EE-6F5A887FDF9F
Discovery of two new genera of detritivorous aquatic beetles Toneroides,
in the Venezuelan Amazon (Coleoptera: Noteridae: Noterinae)
Mauricio M. García Ramírez1, 2
1Laboratorio de Taxidermia y preparados anatómicos “Ramón de Jesús Acosta”,
Centro de Investigaciones Biológicas. Facultad de Humanidades y Educación de La
Universidad del Zulia, Edif. De Postgrado, Zulia 4001-A, Apartado. 526,
Maracaibo, Zulia, Venezuela: http://orcid.org/0000-0003-3238-9527.
²Museo de Artrópodos de La Universidad del Zulia, Facultad de Agronomía. LUZ-
Maracaibo 4002-A, Apartado 526, Zulia, Venezuela.
E-mail: liocanthydrus@yahoo.es
ABSTRACT
Analysis of specimens of Tonerus wheeleri (MALUZ) reveals two distinct
lineages. This finding led to the description of Tonermorpha gen. nov. with two
species, T. unguilonga sp. nov. and T. submersa sp. nov. and Tonerinus gen. nov.
with eight species: Tonerinus amazonas sp. nov., T. amazonensis sp. nov, T.
amazonicus sp. nov., T. hypognathus sp. nov., T. pemonus sp. nov., T. sazhnevi sp.
nov., T. spangleri sp. nov. and T. toboganensis sp. nov. Tonerinus gen. nov. differs
from Tonerus Miller, 2009, in having serrate metatibial spurs, and other diagnostic
features. Tonermorpha gen. nov. with long, slightly curved nails on each tarsus for
a better grip on rocky stream substrates distinguishes it from both genera in addition
to other diagnostic characters of its own. Tonerinus gen. nov. and Tonermorpha
gen. nov. are distinguished by: Spines on the basal mandible margin and lacinia;
microspinous margins on femurs and tibiae (very marked on the protibia of
Tonerinus); a dorsoventral spur on the protibia; crenulate protarsal nails; serrate
metatibial spurs (the posterior one doubly so); bitruncate (Tonerinus) or doubly
sinuate (Tonermorpha) apical margin of the proventral process. A detritivorous and
scavenging behavior is proposed for adults in all three genera, different from the
predatory behavior generally associated with the family Noteridae, based on the
spiny characteristics of the mandible and lacinia. All species are described and
illustrated in detail, including habitus, genital terminalia, genital sclerites, sclerites
DOI: https://doi.org/10.5281/zenodo.13288786
Dos nuevos géneros Tonerini.
García Ramírez.
68
and cephalic appendages, locomotion sclerites. Finally the spatial distribution of
Toneroides species in Venezuela is presented.
Key words: Amazonas, Coleoptera, new species, Tonerus, taxonomy,
Venezuela.
Descubrimiento de dos nuevos géneros de escarabajos acuáticos detritívoros
Toneroides, en el Amazona venezolano (Coleoptera: Noteridae: Noterinae)
RESUMEN
El análisis de ejemplares de Tonerus wheeleri (MALUZ) reveló dos linajes
distintos. Este hallazgo dio lugar a la descripción de Tonermorpha gen. nov., con
dos especie, T. unguilonga sp. nov., y T. submersa sp. nov.; y Tonerinus gen. nov.,
con ocho especies: Tonerinus amazonas sp. nov., T. amazonensis sp. nov., T.
amazonicus sp. nov., T. hypognathus sp. nov., T. pemonus sp. nov., T. sazhnevi sp.
nov., T. spangleri sp. nov. y T. toboganensis sp. nov. Tonerinus gen. nov. difiere de
Tonerus Miller, 2009, por presentar los espolones metatibiales serrados, y otras
características diagnósticas. Tonermorpha gen. nov., con uñas largas, ligeramente
curvadas en cada tarso, para un mayor agarre en los sustratos rocosos de corrientes,
lo distingue de ambos géneros; además de otros caracteres diagnósticos propios.
Tonerinus gen. nov. y Tonermorpha gen. nov. se distinguen por: Espinas en el
margen basal mandibular y lacinia; márgenes microespinosos en fémures y tibias
(muy marcados en la protibia de Tonerinus); un espolón dorsoventral en la protibia;
uñas protarsales crenuladas; espolones metatibiales serrados (el posterior
doblemente); margen apical bitruncado (Tonerinus) o doblemente sinuado
(Tonermorpha) de la apófisis proventral. Se propone un comportamiento detritívoro
y carroñero para los adultos en los tres géneros, diferente al comportamiento
depredador generalmente asociado a la familia Noteridae, basado en las
características espinosas de la mandíbula y lacinia. Todas las especies se describen
e ilustran en detalle, incluyendo habitus, terminalia genital, escleritos genitales,
escleritos y apéndices cefálicos, escleritos de locomoción. Finalmente se presenta la
distribución espacial de las especies Toneroides en Venezuela.
Palabras clave: Amazonas, Coleoptera, nuevas especies, taxonomía, Tonerus,
Venezuela.
Recibido / Received: 12-02-2024 ~ Aceptado / Accepted: 30-05-2024.
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 69
INTRODUCTION
The tribe Tonerini, created by Miller in 2009, was based on a single species of
aquatic beetle: Tonerus wheeleri endemic to the Amazon region of Venezuela.
However, later studies (Baca et al. 2017) synonymized it, considering it a
monotypic tribe and genus. The species was described from specimens collected at
Tobogán de la Selva, on the banks of the Coromoto River, Amazonas State. In this
paper, we present the description of two new genera that, although they share
characteristics with the tribe Tonerini, they differ from the genus Tonerus.
The characterization of these new taxa is supported by a meticulous
comparative study of the external morphology, both dorsal and lateral,
complemented by a thorough analysis of the individual genitalia of each of the ten
species involved in the new genera (García y Briceño 2023). The genital terminalia
of both generic taxa are described in detail, as well as the cephalic sclerites, mouth
appendages and locomotion sclerites (García y Briceño 2023). The described
species represent new morphological stages with characteristic of both generic taxa,
which justifies their separation from the genus Tonerus.
The new species are clearly distinguished from Tonerus wheeleri by a number
of external and internal morphological features. Among these features are the shape
of the cephalic, dorsal and ventral sclerites, the structure of the genitalia of the male
and female, the morphology of the terminalia, the shape of the metatibial spurs, and
the shape of the tarsal claws. The research assumes a detritivorous and scavenging
behaviour for the adults of the genus, based on the spiny characteristics of the
mandible and lacinia (Guzmán-Soto and Tamaris-Turizo 2014, Chará-Serma et al.
2010) which differs from the predatory behaviour generally associated with the
family Noteridae. The objective of this research is the description of two new
genera and ten new species brings the number of known species to eleven and the
number of genera within the Tonerus group to three. This discovery represents a
significant advance in the knowledge of the biodiversity of this group of beetles in
Venezuela.
MATERIAL AND METHOD
The material was collected in the locality designated as “El Tobogán de la
Selva”, a park or recreational resort located in the Ature municipality of Amazonas
70
Dos nuevos géneros Tonerini.
García Ramírez.
state, south of Puerto Ayacucho (1A y 1B). This region is characterized by the
presence of a tropical rainforest with a mean annual temperature of 25 °C and an
annual rainfall of 2500 mm. Eleven specimens were analyzed.
Figure 1A. Distribution of Tonerus wheeleri species group in Venezuela. Geographical
location in the Amazonas state of the locality of the "Tobogán de la Selva" beach camp.
Figure 1B. Extraction of the local area or macrohabitat in which the three species of
Tonerini are distributed: Tonerinus gen. nov., Tonermorpha gen. nov. and Tonerus wheeleri
Miller, 2009; A) Collection area of the eight Tonerinus along the vegetation above the runoff
on the rock on the left bank of the Río Coromoto, “Tobogán de la Selva” and collection site
of the two Tonermorpha species in the “Tobogán de selva” S of Pto Ayacucho [Photo:
Mauricio García, January-2003], the satellite image illustrates the point of collection of
Tonerinus and Tonermorpha by the author(satellite image Google Earth, January, 2003); B)
Tobogancito de la Selva. Tonerus specimens were collected in 2007 and provided on loan to
Andrew Short and Kevin Miller.
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 71
Nine of the specimens were discovered in various natural habitats, including
decomposing log debris, a Madi microhabitat, and superimposed on runoff from a
hygrotrophic microhabitat, among the nascent grass accumulated on the north bank
of the Coromoto River (García et al. 2016). The other two were collected on the
rocky bank of the Rio Coromoto stream. All specimens were collected using a hand-
held suction device at both habitat sites.
The identification and separation of the ten morphospecies was performed
using a LEICA M10 stereo microscope with an 80x resolution objective, and a
binocular compound microscope OPTIMA XSZ-207 was employed for microscopic
observation. The genital sclerites were extracted according to the established
protocol, which involves cleaning and removal of connective tissue with 10%
potassium hydroxide (KOH) solution, followed by washing with distilled water. The
sclerites were preserved in a double-mounted cardboard triangle containing the type
specimen, using an alcohol-based entomological glue. The sclerites were then
illustrated and enhanced in the digital design program Inkscape version 1.3.0. A
combination of drawing and shading techniques was employed to create accurate
and detailed illustrations of the sclerites. Satellite imagen was obtained from Google
Earth Pro. A taxonomic key was constructed to differentiate the genus Tonerus
from the newly identified taxa Tonerinus and Tonermorpha, as well as a map
illustrating the zonation of the species within the tribe.
RESULTS
Taxonomic
Coleoptera Linnaeus, 1758
Adephaga Schellenberg, 1806
Noteridae Thomson, 1860
Noterinae Thomson, 1860
Tonerini Miller, 2009
Tonerinus García, gen. nov.
http://zoobank.org/urna:lsid:zoobank.org:act:9660A569-2E01-4C1E-92CA-D798F75A2817
(Figs. 2-5)
Type species. Tonerinus amazonas Garcia, sp. nov. by original designation.
Dos nuevos géneros Tonerini.
García Ramírez.
72
Differential diagnosis. The genus Tonerinus gen. nov. is characterized by a
robust body shape of the species, with a broad head, prognathous and hypognathous
depending on the species, small eyes and a long labrum. The genus Tonerinus gen.
nov. is characterized by a robust body shape of the species, with a broad head,
prognathous and hypognathous depending on the species, small eyes and a long
labrum.
Description. Species broad, greater width in humeral region. Lateral outline
continuous between pronotum and elytron, with uniform curvature both anterior and
posteriorly. Integral surface shiny and evenly covered with very fine isodiametric
(squamous or microreticulate) epithelial cells. Pronotal ridge with lateral margin
extending narrowly at posteroangular base, expanding and widening evenly towards
anteroangular margin and along anterior margin. Head broader than long with
labrum prostratum and retroflectum depending on species, broad and long, usually
with flat surface (Fig. 2A). Mandible broad with two premolars; first premolar stout
and broad, second premolar short with shaped surface of small teeth (molars); two-
thirds of basal length with series of long and short, thick and thin spines in a row
(Fig. 2B). Maxilla robust with coarsely spiny lacinia; maxillary palpus with the first
three palpomeres similar, the fourth palpomere is long and rectangular with two
small tactile sensors at the apex (Fig. 2C). Lip with two palpiferous openings in
prementum; labial palp with three spatial palpomeres and basal palp inserting
through palpiferous openings adjacent to the dorsal prementum; apical labial
palpomere long and slender with two broad tactile sensoria, one preapical and one
apical (Fig. 2D). Antenna short with bilobed scape, pedicel short, remaining
rectangular antennomeres slightly shorter or longer than pedicel, each with
superficial sensory; apical antennomere long and attenuate at apex with three
sensory micropelons (Fig. 2E). Prosternum broad anteriorly, convex medially.
Apophysis proventral extremely broad, flat and glabrous with bitruncate apex (Fig.
3). Metaventrite glabrous, medially broad and rounded. Platform noteroidal broad,
glabrous, extended anteriorly towards metaventrite; metacoxal lobes with glabrous
apices. Metacoxa and metafurca laterally fused to form a ring. Abdominal ventrites
with glabrous surface; ventrite III+IV very broad; ventrites V and VI slender;
ventrite VII glabrous, broad and long with rounded apex. Fore leg with broad, short
femur, posterior margin with a row of short, thick spines bordering it; tibia very
short and broad bordered with strong short, thick spines at the base and long, stout
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 73
spines in the apical half; protibial spur thick and slender, sparsely curved, pointed at
the apex; protarsus long and thick with a medium-sized discoidal sucker at apex;
tarsomere II long and thick with a small discoidal sucker at apex; tarsomeres III and
IV slender with fine ventral spines; tarsomere V long and slender with a pair of long
apical claws with crenulate ventral edge (Fig. 4A). Middle leg with broad femora
slightly longer than femora, with two parallel rows of thick, short spines bordering
anterior margin; mesotibia broad, one-third longer than protibia, anterior and
posterior margins with long, stout spines bordering it, apex crowned with short,
long, thin spines; tarsomere I very thick and long with a medium-sized discal sucker
at the apex; tarsomere II short and broad with a small discoidal sucker at the apex;
tarsomeres III and IV short with a thick, long spine at their apices; tarsomere V long
with a pair of long, thin spines cleft in the middle (Fig. 4B).
Hind leg with femur broad at the base and attenuated at the apex, slightly
longer than the mesofemur, bordered anteriorly with short, thick spines; metatibia
broad one-third larger than mesotibia, bordered anteriorly and posteriorly with long,
short, coarse spines and apex crowned with medium and coarse spines with the
anterior spur long and coarse with the apical margin doubly serrate with coarse teeth
at the apex, posterior spur short and broad with a row of short, fine serrations at the
apex; metatarsus with the first tarsomere thick, long and cylindrical with spines at
the apex, the remaining tarsomeres are short and cylindrical with spines at the apex
and a pair of thin, straight single claws (Fig. 4C). Genital terminalia with the VIII
abdominal sternite with slight sexual dimorphism, broad, long, longitumedially with
a broad depression on dorsal face projecting on ventral face; dorsal surface densely
setose; in anterior and posterior view forms an arc (Fig. 5); IX abdominal sternites
short, broad and pseudo-rounded with varied rectangular basal margin (see species);
gonocoxoesternites symmetrical and with sexual dimorphism (see species). Genital
sclerite of female with short L-shaped laterotergites; laterotergal base extending
beyond base along laterodorsal margin; gonocoxa flat with pointed apex. The
median lobe on the genital sclerite of the male has a deep ventral groove that curves
along its surface; viewed laterally, it shows a broad, bifurcated phallobase.
Etymology. The new epithet is a diminutive reference to the name Tonerus,
preserving the root of the genus.
Dos nuevos géneros Tonerini.
García Ramírez.
74
Tonerinus amazonas García, sp. nov.
(Figs. 1A-B, 2A, 6A-7A-8A-G)
http://zoobank.org/urna:lsid:zoobank.org:act:709009BC-A929-4ECD-B766-E432A87D54D7
Diagnosis. Seen dorsally Tonerinus amazonas sp. n. is widest at the humeral
margin. In addition, the anterior margin of the pronotum in this species is greater
than the cephalic width, making it wider anteriorly than posteriorly.
Location type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♂, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W), 125 m (MALUZ06649).
Description. Broad oval body shape with rounded anterior and posterior
margin and apex as broad as anteriorly (Fig. 6A). In lateral view the dorsal
convexity is slightly discontinuous between head and pronotum, with lobulation of
the elytral margin between the first and second third followed by arching in the
apical third (Fig. 7A). Length 2.15 mm and width 1.10 mm, widest at humeral
margin. Dorsal colouration dark fulvus (brown) with dark longitudinal and
transverse bands crosswise but diffuse in the central area which is less dark and a
very diffuse arcuate band on the anterior elytral margin. Ventrally the colouring is
fuliginosus (brownish black). Head prognathous wider than long with small eyes 3x
its diameter apart; in lateral view narrowly convex; labrum long, convex and vertical
or prostratum (Figs. 2A, 7A). Pronotum long, 1.5x head length, sparsely convex;
lateral margin narrow with border almost straight 0.7x medial length, and
continuous with elytral border (Fig. 7A). Elytra with dorsal surface fine and sparsely
punctate; lateral margin slightly sinuous; viewed laterally the dorsum is convex
andundulate at the lateral margin (Fig. 7A). IX abdominal sternite short and broad,
subrectangular with very short, broad base and bilobed at basal margin with two
micro-setae on each side; one membranous ventral side with sclerotised margins and
one sclerotised dorsal side (Fig. 8F). Gonocoxoesternites bilobed with broad,
rounded posterior margin and narrow anterior margin with rounded dorsal and
ventral apodemas (Fig. 8G). IX sternite broad and short with rounded apex and
broad base and with bilobed margin and several short fine setae on each lobe (Fig.
8F).
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 75
Figure 2. Cephalic sclerites and mouth appendages in species of Tonerinus gen.
nov. A. Labrum, B. Left mandible, 1m = first molar, 2m = second molar, hsp =
irregular row of mandibular spines, C. Left maxilla, lc = lacinia, pmx = maxillary
palpus in anterior view, 4p = apex of fourth maxillary palpomere in lateral view, st =
tactile sensory, D. Lip, abp = palpal aperture, pmt = prementum, slp = lateral setae
of prementum, plb = labial palpus in anterior and lateral views, plpg = palpiger
(basal palpomere of labial palpus fused behind prementum), E. Antenna, sp = scape,
pd = pedicel.
Dos nuevos géneros Tonerini.
García Ramírez.
76
Figure 3. Apophysis proventralis of Tonerinus species gen. nov.
Gonocoxosternite with broad posterior end almost rounded and narrow anterior
margin with rounded apodemes and series of short fine setae on basal lateral margin
(Fig.8G). Male genital sclerite with median lobe curved from middle, attenuating to
apex; broad bilobed phallobase with sinuous margin; right paramere short and broad
with acute end and left paramere curved with acute L-shaped base and several apical
setae (Figs. 8A-E).
Female. Unknown.
Etymology. The epithet refers to the local region in Venezuela and dedicates
the species to the state of Amazonas.
Habitalogy. The species was collected in a microhabitat of the limnetic/lentic
hydroecological system, "Madis" (García et al. 2016).
Distribution. Restricted to the locality of the Tobogán de la selva recreational
park located in the riparian corridor of the Río Sipapo and Río Coromoto, Amazonas
State.
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 77
Tonerinus amazonensis García, sp. nov.
(Figs. 1A-B, 2A, 6B-7B, 9A-C)
http://zoobank.org/urna:lsid:zoobank.org:act:C410B933-55DA-45F7-A228-390D3E37C469
Differential diagnosis. Tonerinus amazonensis sp. nov. resembles T.
amazonas, but with a slightly shorter and less broad pronotum and a non-sinuate
elytral lateral margin. However, T. amazonensis sp. n. differs in the elytral pattern.
Instead of longitudinal and transverse bands, it has four very diffuse corner maculae
with a central fulvus settlement darker than the rest of the elytral surface. In addition,
T. amazonensis has a more extended head, similar to T. amazonas. Viewed laterally,
the habitus of T. amazonensis is wavy with the lateral edge of the pronotum rounded
and without elytral lobulation. In contrast, T. amazonas has a slightly rounded,
almost straight elytral margin and an elytral lobulation almost in the middle.
Localidad tipo. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♀, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W), 125 m (MALUZ06648).
Description. Body shape broad oval with discontinuous convexity between
head and pronotum; rounded anterior and posterior margin and broad apex; with
extended cephalic prognathism (Fig. 6B). In lateral view it is ventrally undulated
without lobulation of the elytral margin followed by an arching in the apical third
(Fig. 7B). Length 2.20 mm and width 1.10 mm, widest at humeral margin. Dorsal
colouration fulvous with four slightly obscured corner maculae separated by a
distinctly reddened blurred central area and a blurred arc on the anterior margin.
Head broader than long, prognathous, with small eyes 3x its diameter apart; sparsely
convex when viewed laterally with discontinuity between labium and clypeus;
labium long convex and prostratum (Fig. 2A). Pronotum long 1.5x cephalic length,
sparsely convex; broad lateral margin viewed laterally with rounded edge, of equal
medial length and discontinuous with elytral margin. Elytra convex with fine,
slightly punctate surface with thick ventral margin, and seen laterally straight in the
first two elytral thirds, arcuate in the apical third (Fig. 7B). Gonocoxoesternite broad
pseudo rectangular with rounded apodemes and a sharp lateral apodeme; dorsal mar-
Dos nuevos géneros Tonerini.
García Ramírez.
78
gin with a series of basal setae (Fig. 9A). Female genital sclerite with very short
laterotergite forming a basal lobe extending along laterodorsal margin of gonocoxa;
small gonocoxa with broad apex with a row of setae extending to almost apical half
of lateroventral margin, lobed anteriorly (Figs. 9 B, C).
Male. Unknown.
Etymology. The new epithet is dedicated to the gentilicio of origin or
belonging of the species to the state of Amazonas.
Habitalogy. Similar to the previous species.
Distribution. Similar to the previous species.
Tonerinus amazonicus García, sp. nov.
(Figs. 1A-B, 2A, 6C-7C, 10A-G)
http://zoobank.org/urna:lsid:zoobank.org:act:B5D9896A-2E19-4039-B408-70F19757FD15
Differential diagnosis. This species is distinguished from those described
above by unique characteristics. Firstly, the pronotal margin is narrower in the
transverse half. This is because the lateral margins are not arched, but taper in the
middle, almost forming a straight line. Secondly, the elytral pattern of perpendicular
bands is very thick, more concise and not very blurred.
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♂, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W), 125 m (MALUZ06617).
Description. Body shape broad oval, convex with rounded anterior and
posterior margins: viewed laterally there is discontinuity between head and
pronotum (Fig. 6C). Lateral view convex with sinuous ventral margin (Fig. 7C).
Length 2.25 mm and width 1.15 mm, widest at humerus. Dorsal colouration dark
fulvus with elytral bands fuliginosus; ventrally colouration fuliginosus. Head large
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 79
prognathous, broader than long with small eyes 3x their diameter; labrum long,
convex and prostratum discontinuous with clipeal margin (Fig. 2A); lateral view
slightly flat surface. Pronotum long 1.5x cephalic length, surface slightly convex;
lateral view pronotal margin equal to 0.8 x medial length with margin narrowing
posteroangularly and widening posteriorly to anteroangular apex, rounded edge
discontinuous with elytral margin. Elytra convex with finely punctate surface;
ventral margin arcuate only in apical third. IX abdominal sternite broad, ovoid
rounded with very short, lobed base (Fig. 10F). Gonocoxosternite short, broad,
narrow anterior margin with broad, triangular apodemes; posterior margin very
broad and sinuous with rows of lateral setae (Fig. 10G). Genital sclerite of male with
broad median lobe, curved in apical half attenuated at apex; falobase broad and
bilobed. Right paramere broad, turned in middle and left paramere long with two
apical thirds broad and one basal third attenuate (Figs. 10A-E).
Female. Unknown.
Etymology. The epithet "amazonicus" is an adjective of Latin origin meaning
"belonging to or relating to the Amazon" found in the Amazon basin.
Habitalogy. Similar to the previous species.
Distribution. Similar to the previous species.
Tonerinus hypognathus García, sp. nov.
(Figs. 1A-B, 2A, 6D-7D, 11A-G)
http://zoobank.org/urna:lsid:zoobank.org:act:485A5BFF-BD90-4952-A206-2D59A33A8BC8
Differential diagnosis. Tonerinus hypognathus sp. n. is distinguished from
other species of the genus by a number of unique characteristics. It has a
hypognathous head, i.e. it has a very downward sloping head. This means that only
the vertex is visible in dorsal view, and the surface of the head is flattened in lateral
view. Tonerinus hypognathus has a more robust and convex body shape from the
labrum. The width and length of the pronotum are greater than in other species of
the genus. The elytral banding pattern is more diffuse in the elytral half. In lateral
view, the marginal margin of the pronotum is almost straight and is in line with the
Dos nuevos géneros Tonerini.
García Ramírez.
80
elytral margin. These distinctive features allow differentiation of T. hypognathus
from the other species of the genus Tonerinus.
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♂, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W) 125 m (MALUZ06650).
Description. Body shape robust, broad oval and very convex, with the anterior
margin rounded, slightly discontinuous with the pronotal margin the apical end in
less rounded and less broad (Fig. 6D); in lateral view it is very convex with the
labrum completely inclined; the ventral margin is almost straight except for the
somewhat arched elytral apex (Fig. 7D). Length 2.26 mm and width 1.15 mm,
widest at humerus and posterior margin of pronotum. Testaceous colouration on
dorsum with the cross pattern of elytral bands very blurred except for the apex of the
longitudinal band. Head as broad as long, hypognathous, flattened in lateral view
and long, convex labrum and prostratum below the longitudinal axis of the body
(Fig. 2A); small eyes 4x its diameter apart. Pronotum long and broad, 2x cephalic
length, convex; anteriorly with a row of small circles or beads along it and the
posterior margin with a short C-shaped row of small ovals above the scutellar
margin; in lateral view the pronotal margin is broad with straight edge, in line with
the elytral margin and equals the medial length. Elytra very convex with finely
punctate surface with ventral margin straight in first two thirds and arched at apex.
IX abdominal sternite broad with sinuous apical margin and rounded lateral margins
with short, bilobed base (Fig. 11F). Gonocoxosternite short and broad with narrow
anterior margin and broad, short, rounded apodemes (Fig. 11G). Genital of male
with broad median lobe of bilobed phalaobase with apical margin curved and
attenuate at apex; left paramere broad in two apical thirds with one of the margins
sinuous and one basal third attenuate and right paramere broad pseudo rectangular
turned in the middle, apex and base attenuate (Figs. 11A-E).
Female. Unknown.
Etymology. The species is named "hypognathus" because of the inclined
position of the head with the jaws vertical, different from the prognathous position
of the previous species.
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Figure 4. locomotion sclerites of Tonerinus gen. nov. species: A. Propata, B.
Mesopata and C. Metapata. Metapata: spt = protibial spur, ven = protarsal and
mesotarsal disc suckers, uc = protarsal nail with crenulated ventral edge and uh =
mesotarsal nail with cleft ventral edge, spp = posterior spur doubly serrate at apical
margin and spa = anterior spur serrate at apical margin, 2s = double row of teeth and
1s = single row of teeth.
Habitalogy. Similar to previous species.
Distribution. Similar to previous species.
Dos nuevos géneros Tonerini.
García Ramírez.
82
Figure 5. VIII abdominal sternite in Tonerinus species gen. nov.: vd = dorsal view of VIII
abdominal sternite of male and female specimens, van = anterior view, dplg = dorsal
longitudinal depression, vap = posterior view, ablg = ventral longitudinal bulge.
Tonerinus pemonus García, sp. nov.
(Figs. 1A-B, 2A, 6E-7E, 12A-G)
http://zoobank.org/urna:lsid:zoobank.org:act:3B977CA8-B3FC-4E35-BBE1-99D65730F620
Differential diagnosis. Tonerus pemonus sp. n. is related to T. spangleri sp. n.,
but differs from it in a number of key features. Tonerinus pemonus has bicoloured
colouration on the head and pronotum, which is distinct from the elytral coloura
tion. The design of the elytral bands in T. pemonus is different from that of T.
spangleri.
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In T. pemonus the elytral bands form a V-shape between the transverse maculae
and the apical macula. In addition, the basal band is arched and joins longitudinally
with the anterior transverse band. In contrast, T. spangleri has a cross-shaped
pattern. In lateral view, T. pemonus is very broad and convex from labrum to apex,
with a sinuous ventral margin. This characteristic distinguishes it from the other
species described above.
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♂, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W), 125 m (MALUZ06647).
Description. Robust, highly convex oval shape with rounded anterior and
posterior margins (Fig. 6E); in lateral view shows prominent arching without dorsal
discontinuities and a doubly sinuous ventral margin (Fig. 7E). Length 2.20 mm and
width 1.16 mm, greater width on humerus. Colouring of dorsum bicoloured with
head and pronotum testaceous yellowish and elytra testaceous reddish with elytral
bands forming a dark V in apical half a crescent in basal half connected
longitudinally. Ventrally only the proventral process, metaventrite and noteroidal
plate are fuliginosus, abdominal ventrites reddish testaceous and the rest of the
sclerite yellowish testaceous. Head broader than long, robust and convex, slightly
hypognathous or inclined with labrum below longitudinal axis of body; labrum
long, convex and prostratum (Fig. 2A); eyes small, 3x their diameter apart.
Pronotum long, 1.5x cephalic length; anterior margin with series of small circles or
beads along it with some setose points and posterior margin with short row of small
C-shaped ovals in front of scutellar margin; in lateral view convex and pronotal
margin broad and rounded edge not in line with elytral margin, and equal to its
medial length. Elytra with finely punctate surface and some setae spaced at apex; in
lateral view very convex on dorsum and ventral margin sinuous, with a small broad
lobulation between the first elytral thirds. IX abdominal sternite (Fig. 12F).
Gonocoxosternite (Fig. 12G). Genitalia of male with median lobe broad in basal
half and attenuate in apical half with curved, acute apex; falobase very broad with
sinuous, bilobed margins; right paramere S-shaped twisted with bilobed base and
left paramere of equal length to right with two broad apical thirds with long, thick
setae at apex and attenuate basal third (Figs. 12A-E).
Dos nuevos géneros Tonerini.
García Ramírez.
84
Female. Unknown.
Etymology. The new species is dedicated to the Pemon ethnic group.
Habitalogy. Similar to previous species.
Distribution. Similar to previous species.
Tonerinus sazhnevi García, sp. Nov.
(Figs. 1A-B, 2A, 6F-7F, 12A-G)
http://zoobank.org/urna:lsid:zoobank.org:act:20D5AB31-2C96-4493-9197-737B6D0A8B2C
Differential diagnosis. The species is similar to T. amazonas and differs from
it only in lateral view, as it is wider and much more arched dorsally, has no dorsal
discontinuity and the head is more inclined with respect to T. amazonas. Another
difference is observed in the lateral margin of the pronotum which is pseudo-
rounded at the edge in contrast to the straight edge of T. amazonas. This condition
separates it from the rest of the species of the genus.
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♀, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W) 125 m (MALUZ06646).
Description. Body shape oval and broad with rounded anterior and posterior
margins and lateral margin with a strong discontinuity between head and pronotum
and slightly sinuous in the elytral region (Fig. 6F); in lateral view the dorsum is
very convex and the ventral margin is doubly sinuous (Fig. 7F). Length 2.30 mm
and width 1.18 mm, greatest width at the level of the humerus. Colouring of dorsum
fulvus with diffusely darkened perpendicular elytral bands. Ventrally only the
proventral process, metaventrite and noteroidal plate are fuliginosus, the abdominal
ventrites reddish testaceous and the rest of the ventral sclerites yellowish testaceous.
Head broader than long slightly inclined and slightly stout; labrum long and convex,
prostratum (Fig. 2A); seen from side is convex, labrum and clypeus in line; eyes
small 3x separated by 3x their diameter Pronotum long and broad; anterior margin
with row of small circles or beads along it and posterior margin with short C-shaped
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row of small ovals in front of scutellar margin; in lateral view margin narrow and
pronotal margin lobed anteriorly and straight posteriorly in discontinuity with
ventral elytral margin, and equal to medial length. Elytra with finely punctate
surface with some scattered setose points at the apex; seen laterally it has a strong
dorsal arching and the ventral margin is doubly sinuous with a long narrow
lobulation between the two basal thirds, arched in the apical third. Gonocoxosternite
very broad with the dorsal apodeme very broad relative to the ventral apodeme and a
long rounded apodeme on the ventral lateral margin and a series of long setae on the
dorsal margin (Fig. 13C). Genitalia of female with laterotergite very short, sinuous,
with broad, rounded apex and base extending along laterodorsal margin in a thin,
curved blade at end. Gonocoxa small and broad anteriorly and attenuate posteriorly
with rounded apex, with a row of small, fine spines bordering the apex and
extending almost to the middle of the lateroventral margin (Figs. 12A, B).
Male. Unknown.
Etymology. The name of the new species is dedicated to an invaluable scientist
and scholar of aquatic coleoptera Dr. Alexey S. Sazhnev.
Habitat. Similar to the previous species.
Distribution. Similar to previous species.
Tonerinus spangleri García, sp. nov.
(Figs. 1A-B, 2A, 6G-7G, 14A-E)
http://zoobank.org/urna:lsid:zoobank.org:act:D9032DCD-04D2-48DD-B489-D2F8FC1D72CD
Differential diagnosis. In T. spangleri sp. nov., the elytral bands form a cross
with diffusion in the centre. Tonerinus spangleri differs from the species described
above by the curved shape of the pronotal margin and the elytral banding pattern. In
lateral view, T. spangleri has a much broader and more convex head and no
discontinuity in the dorsal convexity.
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♂, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W) 125 m (MALUZ06645).
Dos nuevos géneros Tonerini.
García Ramírez.
86
Description. Body shape broad oval and convex, with rounded anterior and
posterior margins (Fig. 6G); in lateral view it is observed as a perfect arch and a
lobed ventral margin at pronotum level and slightly arched at level of elytra (Fig.
7G). Length 2.20 mm and width 1.10 mm, widest at humerus level. Dorsal colouring
testaceous and a dark cross-banded pattern blurred in the centre. Ventrally
proventral process, metaventrite and noteroidal plate fuliginosus, head, proventrite,
abdominal ventrites reddish testaceous and legs yellowish testaceous. Robust head
broader than long, prognathous with long, prostratum and convex labrum (Fig. 2A);
small eyes 3x their diameter apart; lateral view convex and continuous with
pronotum. Pronotum long and convex, 1.5x cephalic length; anterior margin with a
series of small circles or beads along margin with some fine setose points; posterior
margin with a short row of small C-shaped tegumental ovals above scutellar margin;
lateral margin broad with rounded edge not continuous with elytral margin, and
equals 0.8x medial length. Elytra convex with the elytral surface finely punctate
with scattered setose points at the apex; in lateral view the ventral margin is
slightly curved in the first two thirds somewhat accentuated in the last elytral third.
IX sternite very broad and short pseudo rectangular with very short and sinuous base
with several short fine setulae on margin (Fig. 14E). Gonocoxosternite short and
slightly widened posteriorly and slightly narrowed at anterior margin with dorsal
apodemes triangular and ventral apodemes rounded (Fig. 14D). Genital of male with
median lobe widened in basal two-thirds and attenuated in apical one-third with
acute apex; phallobase broadly bilobed (Figs. 14A-C).
Female. Unknown.
Etymology. The new of the species is dedicated to Dr. Paul Spangler (†) a
great researcher of the aquatic Coleoptera of the world.
Habitalogy. Similar to previous species.
Distribution. Similar to previous species.
Tonerinus toboganensis García, sp. nov.
(Figs. 1A-B, 2A, 6H-7H, 15A-G)
http://zoobank.org/urna:lsid:zoobank.org:act:57B14576-750B-4321-A278-B5037E3EC851
Differential diagnosis. Although this species is remarkably similar to T.
amazonicus, it is distinguished by a clearer colouration and a less defined and more
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Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 87
diffuse pattern of elytral bands. In lateral view, there is a greater width and a more
pronounced arched profile, with a more robust head. The pronotal margin is more
sloping and the ventral margin is lobed in the basal two thirds of the elytra. This
arched feature clearly differentiates it from the rest of the species of the genus.
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♀, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W), 125 m (MALUZ06640).
Description. Body shape broad oval with anterior and posterior margins
rounded (Fig. 6H); viewed laterally strongly convex on dorsum and doubly sinuous
on ventral margin with a pronounced arcuate point between pronotum and elytra
(Fig. 7H). Length 2.25 mm and width 1.15 mm, widest at humerus. Colouring of
dorsum testaceous yellowish with diffuse elytral bands forming a Y in the apical
half and a crescent in the basal half. Head broader than long, prognathous; in lateral
view narrowly convex not in line with labrum, long and convex, prostratum (Fig.
2A); eyes small, 3x their diameter apart. Pronotum long and broad with small bead-
like circles along anterior margin; posterior margin with a row of small ovals
forming a C in front of scutellar margin; in lateral view convex with pronotal margin
very broad and margin slightly rounded in discontinuity with ventral margin and
equals medial length. Elytra with the surface finely punctate with some scattered
setae; in lateral view very convex with the ventral margin gently lobed in the first
two thirds of the elytra sloping towards the apex. IX abdominal sternite very broad
and rounded with short base and straight margin (Fig. 15F). Gonocoxosternite short
axe-shaped, anteriorly rounded with angular and very short apodemes almost
imperceptible; posterior margin curved and acute at apex (Fig. 15G). Genitalia of
male much widened at bilobed phallobase with attenuate apical one-third and acute
apex; right paramere half-turned, with attenuate ends and rectangular left paramere
with acute base and rows of long hairs on two apical thirds (Figs. 15A-E).
Female. Unknown.
Habitat. Similar to the previous species.
Distribution. Similar to the previous species.
Dos nuevos géneros Tonerini.
García Ramírez.
88
Figure 6. Dorsal habitus of Tonerinus species. A. Tonerinus amazonas sp. nov., B.
T. amazonensis sp. nov., C. T. amazonicus sp. nov., D. T. hypognathus sp. nov., E.
T. pemonus sp. nov., F. T. sazhnevi sp. nov., G. T. spangleri sp. nov., and H. T.
toboganensis sp. nov.
Figure 7. Habitus dorsal de las especies de Tonerinus. A. Tonerinus amazonas
sp. nov., B. T. amazonensis sp. nov., C. T. amazonicus sp. nov., D. T.
hypognathus sp. nov., E. T. pemonus sp. nov., F. T. sazhnevi sp. nov., G. T.
spangleri sp. nov., y H. T. toboganensis sp. nov.
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Figure 8. Male genitalia of Tonerinus amazonas sp. nov.: A. median lobe in right
lateral view, B. median lobe in ventral view, C. median lobe in left lateral view, D.
right paramere, E. left paramere, F. IX abdominal sternite in dorsal view, stb = two
basal lateral setae and G: left lateral and right lateral male Gonocoxoesternites, apd
= dorsal apodeme and apv = ventral apodeme.
Dos nuevos géneros Tonerini.
García Ramírez.
90
Figure 9. Female genitalia of Tonerinus amazonensis sp. nov.: A. Laterotergite in
left lateral view, B. Gonocoxa in left lateral view, C. Gonocoxoesternite in dorso-
ventral lateral view, ad = dorsal apodeme and av = ventral apodeme, lb =
laterotergal basal lobe, a = laterotergalapex, ld = laterotergal dorsal lobe, mlt =
laterodorsalmargin, ap = gonocoxalapex.
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Figure 10. Male genitalia of Tonerinus amazonicus sp. nov.: A. Median lobe in
right lateral view, B. Median lobe in ventral view, C. Median lobe in left lateral
view, D. Right paramere, E. Left paramere, F. IX abdominal sternite in dorsal view,
cd = dorsal face and cv = ventral face, lb = basal lobe and G. left lateral and right
lateral male Gonocoxosternites, apd = dorsal apodeme and apv = ventral apodeme.
92
Dos nuevos géneros Tonerini.
García Ramírez.
Figure 11. Male genitalia of Tonerinus hypognathus sp. nov.: A. median lobe in
right lateral view, B. median lobe in ventral view, C. median lobe in left lateral
view, D. left paramere, E. right paramere, F. IX abdominal sternite in dorsal view,
cd = dorsal face and cv = ventral face, lb = basal lobes and G. left lateral and right
lateral male gonocoxosternites, apd = dorsal apodeme and apv = ventral apodeme.
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Figure 12. Male genitalia of Tonerinus pemonus sp. nov.: A. median lobe in right
lateral view, B. median lobe in ventral view, C. median lobe in left lateral view, D.
right paramere, E. left paramere, F. IX abdominal sternite in dorsal view, stb = three
basal lateral setae and G: left lateral and right lateral male gonocoxosternites, apd =
dorsal apodeme and apv = ventral apodeme.
94
Dos nuevos géneros Tonerini.
García Ramírez.
Figure 13. Female genitalia of Tonerinus sazhnevi sp. nov.: A. Laterotergite in left
lateral view, B. Gonocoxa in left lateral view, stgcx = laterodorsalgonocoxal setae,
dgcx = lateroventral and apical gonocoxal spines, C. Gonocoxosternite in right and
left lateral view, ad = dorsal apodeme and av = ventral apodeme, apl = lateral
apodeme, stegon = gonocoxosternal setae.
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Figure 14. Male genitalia of Tonerinus spangleri sp. nov.: A. median lobe in right
lateral view, B. median lobe in ventral view, C. median lobe in left lateral view, D.
male left lateral and right lateral gonocoxosternites, E. IX abdominal sternite in
ventral view, apd = dorsal apodeme and apv = ventral apodeme, E. IX abdominal
sternite in ventral view, cv = ventral face and cd = dorsal face, stb = basal setae.
96
Dos nuevos géneros Tonerini.
García Ramírez.
Figure 15. Male genitalia of Tonerinus toboganensis sp. nov.: A. median lobe in
right lateral view, B. median lobe in ventral view, C. median lobe in left lateral
view, D. right paramere, E. left paramere, F. IX abdominal sternite in ventral view,
cv = ventral face and cd = dorsal face, G: left lateral and right lateral male
gonocoxosternites, apd = dorsal apodeme and apv = ventral apodeme.
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Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 97
Tonermorpha García, gen. nov.
(Figs. 1A-B, 2A-B, E, 16-19)
http://zoobank.org/urna:lsid:zoobank.org:50A1AAAF-7A39-4E56-A449-6951FB01806E
Especie tipo. Tonermorpha unguilonga García, sp. nov. por designación
original.
Differential diagnosis. The new taxon Tonermorpha gen. nov. differs from the
genera Tonerus Miller and Tonerinus gen. nov. by a number of distinctive features.
In terms of claw length, Tonermorpha has claws as long as tarsomere five on the
anterior and middle tarsi and as long as the fourth and fifth together on the posterior
tarsi, unlike Tonerus and Tonerinus, where the claws are shorter than tarsomere five.
The shape of the apophysis is also distinctive: in Tonermorpha, the apophysis is
doubly sinuate at the apex, whereas in Tonerus it is slightly curved with two slightly
straight sides at the apex and in Tonerinus the apical margin of the apophysis is
distinctly bitruncate. As for the serration of the metatibial spurs, Tonermorpha
presents both metatibial spurs serrated, while in Tonerus the metatibial spurs are
simple. Finally, the anterior and posterior margins of the protibia are less spiny in
Tonermorpha, and strongly spiny in Tonerinus.
Description. Species broad, greater width in humeral region. Lateral outline
continuous between pronotum and elytron, with uniform curvature both anteriorly
and posteriorly. Integral surface shiny and evenly covered with very fine
isodiametric (squamous or microreticulate) epithelial cells. Pronotal fold with lateral
margin extending narrowly at posteroangular base, expanding and widening evenly
towards anteroangular margin and along anterior margin. Head broader than long
with labrumprostratum, broad and long, usually with flat surface (Fig. 2A).
Mandible broad, with two premolars (Fig. 2A). Short second premolar with profiled
surface of small teeth (molars); two-thirds of basal length with series of long and
short, thick and thin spines in a row (Fig. 2B). Maxilla robust with coarsely spiny
laciniae; maxillary palpus with the first three palpomeres similar, the fourth
palpomere is long and rectangular with two small tactile sensors at the apex; its
anterior margin is outlined by a thick carina along its entire length (Fig. 17).
Labium with two palpiferous openings in the prementum; labial palpus with three
spatial palpomeres and palpifer inserting through palpiferous openings and adjacent
to the dorsal prementum; apical labial palpomere, one preapical and the other apical
98
Dos nuevos géneros Tonerini.
García Ramírez.
separated by a convex space with the anterior margin outlined in a thickened carina
along its entire length (Fig. 17). Antenna short with bilobed scape, pedicel short,
remaining rectangular antennomeres slightly shorter or longer than pedicel, each
with superficial sensory; apical antennomere long and attenuate at apex with three
sensory micropelons (Fig. 2E). Proventrite broad and long longitumedially with
anterior margin with a small apical cleavage convex medially; proventral apophysis
extremely broad, flat, short and glabrous with bisinuate apex (Fig. 17). Metaventrite
glabrous, medially broad and rounded. Platform noteroidal broad, glabrous,
extended anteriorly towards metaventrite; metacoxal lobes with glabrous apices.
Metacoxa and metafurca laterally fused to form a ring. Abdominal ventrites with
glabrous surface; ventrite III+IV very broad; ventrites V and VI slender; ventrite VII
glabrous, broad and long with rounded apex. Anterior femur short, broad, with
narrow apex; anterior margin with series of short, thick spines bordering on apical
half; posterior margin with series of very fine setae along entire length. Tibiae small,
pseudo-rectangular with fine, short, sparse spaced setae; anterior margin with some
long, thick spines on apical half; posterior margin with rows of short, thick spines on
basal two-thirds and long, thick spines on apical third; apex slightly rounded with
crown of short, thick spines and two long, slender, very sharp spurs, one at each end;
protibial spur long and slender slightly curved at apex. Short tarsi gradually
elongating with the fifth very long equivalent to the fourth and third combined;
tarsal nail very long almost the length of the fifth tarsomere (Fig. 18a).
Mid femur long, broad and rectangular with glabrous surface except for a series
of short, thick spines bordering the anterior margin along the entire length and a
series of long, thin, widely spaced setae on the posterior margin; femoral base
attenuated to form a narrow lobe and rounded femoral apex. Median tibia cylindrical
with glabrous disc surface and margins strongly and thickly spiny, with retractile
spines; apical margin with crown of thick, short, long spines, with long sharp spurs;
median tarsi with broad tarsomeres I and II; long fifth tarsomere with widened apex
of greater length than fourth and third combined, with pair of thin, very long, almost
straight nails of similar length to fifth tarsomere (Fig. 18b). Hind femur short, broad,
attenuate at base, broadened and rounded at apex; disc surface glabrous with a short
series of short, fine spines on apical half of anterior margin. Hind tibia broad,
pseudo-triangular with broad base and bitruncate apex bordered with short, coarse
spines; disc surface glabrous with long, coarse spines on anterior and posterior
margins; posterior metatibial spurs thick and long; anterior spur long with lateral
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Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 99
margin serrate at apex and posterior spur short and doubly serrate at apex (Fig. 18d).
Posterior tarsomeres cylindrical, with the fifth less in length than the fourth and third
combined, and a pair of very long claws greater in length than the fifth and fourth
combined (Fig. 18c). VIII abdominal sternite with surface with scattered short,
coarse setae; lateral margins with slight broad depressions; marginal margin sinuate;
longitumedially depressed (Fig. 19A). Laterotergite very short, curved and thin at
base and furrowed and broad at apex (Fig. 19B). Gonocoxa with laterodorsal
lateroventral margins arched; apical lateroventral margin with series of short spines
and rounded apex (Fig. 19B).
Etymology. “Tonermorpha” this generic name originates from the root “Toner-
“ this root comes from the name of the tribe Tonerini to which the genus belongs
and the suffix ‘-morpha’ indicating distinctive shape or appearance, which in context
means “shaped like Tonerus”.
Tonermorpha submersa García, sp. nov.
(Fig.1A-B, 16A-B)
http://zoobank.org/urna:lsid:zoobank.org:act:C4B782D3-D34F-47D5-9D4E-6C0DA3AED0EA
Diagnosis. This species present a greater diffusion of dark colouration on the
elytral disc and by having the greatest width at the level of the humeral margin,
below the pronotal base, in T. unguilonga the greatest width is at the level of the
basal pronotal margin.
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela
Material type. Holotype ♂, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W), 125 m (MALUZ06660).
Description. Wide oval shape, very convex, with rounded anterior and
posterior margin; seen laterally the dorsum is slightly discontinuous between the
margin of the head and pronotum and very discontinuous between the lateral margin
of head, pronotum and elytra. Length 2.25 mm and width 1.20 mm, greater width at
the level of the humeral margin. Fulvus coloration with dark shades on the lateral of
eyes, pronotum discal zone and on the elytra three rectangular dark spots, diffuse, on
the scutellar margin and medial sides. Head wider than long; small eyes 3x diameter
100
Dos nuevos géneros Tonerini.
García Ramírez.
apart; labrum curved, broad and short. Pronotum long 1.3x head length; lateral
margin with slightly straight edge, 0.8x its total length. Elytra with sparsely punctate
surface and widely spaced setae at level of apical third; in lateral view ventral
margin slightly straight in first two thirds, with sinuosity in apical third. Genital
sclerites and genital terminalia misplaced.
Female. Unknown.
Etymology. The new epithet refers to the aquatic environment based on the
dimension of the long claws used for grasping the rocky substrate of currents:
“Submersus” comes from the classical Latin “submergere, submersus” and means
“submerged” or “sunken” the species does not have swimming hairs on the legs and
it is thought that they are not adapted to swimming but walk on the rocky substrate
of the Coromoto River littoral.
Habitalogy. The specimen was collected together with the other species in the
group on the rocky bank of the Río Coromoto, a hygropetric horizontal run off
microhabitat of a limnic/lotic hydroecological system (García et al. 2016).
Distribution. Restricted to the locality of the “Tobogán de la selva”, a river
side corridor of the Coromoto River, in the state of Amazonas.
Tonermorpha unguilonga García, sp. nov.
(Fig. 1A-B, 16C-D)
http://zoobank.org/urna:lsid:zoobank.org:act:8E078D37-A81F-4A38-86B7-C75792EB8429
Diagnosis. Pronotum broad with oval basal margin, widening markedly
towards the humeri. Elytral surface of diffuse dark colouration on the central disc,
forming four small square bands at the distal ends, significantly darker than the
central region.
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♀, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W), 125 m (MALUZ06659).
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 101
Locality type. Tobogán de la selva, Ature, Amazonas, Venezuela.
Material type. Holotype ♀, from Venezuela, Amazonas, Ature, Tobogán de la
selva, 20.i.2003, M. García and M. Balke cols., (5º 23'12.01" N and 67º 36'57.34"
W), 125 m (MALUZ06659).
Description. Description. Broad oval shape with rounded anterior and posterior
margins and continuous lateral margins between pronotum and elytra, with a small
discontinuity at head and pronotal margin; in lateral view of uniform convexity from
head to elytral apex. Length 2.20 mm and width 1.30 mm, greater width at pronotal
and humeral margin. Coloration of the head fulvous with dark shades distributed on
the inner margins of the eyes. Pronotum fulvus with dark shading on pronotal disc.
Elytra fulvous with pseudo rectangular dark shading on lateral and apical ends
illuding a V in elytral half; and a larger triangular dark shading on anterior margin,
joined to dark shading of pronotal posterior margin. Ventrally with apophysis,
metaventrite and abdomen blackish with fulvus shades; legs with light fulvous and
dark shades. Head wider than long with small eyes separated by three times its
diameter. Labrum flat, long, broad, retroflectum (Fig. 2); in lateral view continuous
convexity with pronotum. Pronotum long 1.2x cephalic length; convexity
continuous with elytra; in lateral view equivalent length 0.8x of central disc;
rounded margin slightly discontinuous with elytral margin. Elytra convex; lateral
view with ventral margin doubly sinuate to apex.
Male. Unknown.
Etymology. The new epithet is a combination of two words relating to the
length of the tarsal nails: ‘Unguis’ meaning nail and ‘Longus’ meaning long, so the
meaning in context is ‘with long nails’.
Habitalogy. The specimen was collected together with the other species in the
group on the rocky bank of the Río Coromoto, a hygropetric horizontal runoff
microhabitat of a limnic/lotic hydroecological system (García et al. 2016).
Distribution. Restricted to the locality of the ‘Tobogán de la selva’, a riverside
corridor of the Coromoto River, in the state of Amazonas.
102
Dos nuevos géneros Tonerini.
García Ramírez.
Taxonomic key to separate genera of Toneroides group.
(Fig. 20)
1.-Tarsal claws of the protarsus and mesotarsus equal in length to the fifth
tarsomere and longer than the fifth and fourth combined on the metatarsus (Fig.
18)…………………………………………………………Tonermorpha gen. nov.
-Tarsal claws of equivalent length shorter than the fifth tarsomere (Fig. 4)........ 2
2.-Metatibial spurs with serrate margins (Fig. 4c)...................Tonerinus gen. nov.
- Simple metatibial spurs………....................................................Tonerus Miller.
Figure 16. Dorsal and lateral habitus of Tonermorpha gen. nov. A. Dorsal and B.
Lateral views of Tonermorpha submersa sp. nov. C and D. Tonermorpha
unguilonga sp.nov.
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 103
Figure 17. Cephalic and ventral sclerites of Tonermorpha gen. nov. A. maxilla
palpus, B. labial palpus and C. Proventral apophyisis.
Figure 18. Locomotion sclerites of Tonermorpha species gen. nov. gen. nov. A.
Propata, B. Mesopata, C. Metapata and D. Metatibial spurs, sptb = protibial spur.
Dos nuevos géneros Tonerini.
García Ramírez.
104
Figure 19. VIII abdominal sternite female genital sclerite of Tonermorpha
unguilonga sp. nov. A. VIII sternite: as = lateral loop, pl = lateral pore, dpl = lateral
depression, dp = apical depression, slgt = longitumedial groove, B. Genital sclerite:
gcx = gonocoxa, lto = laterotergite, altg = laterotergal apex, agcx = gonocoxal apex.
Figure 20. Toneroides group (Tonerini). A. Tonerinus amazonicus García, gen.
etsp. nov., B. Tonerus wheeleri Miller and C. Tonermorpha unguilonga García, gen.
etsp. nov.
Boletín del Centro de Investigaciones Biológicas
Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 105
DISCUSSION
Morphological features in both new genera suggest a primitive condition: the
presence of cephalic characters such as mouth sclerites adapted to a passive diet
(non-predatory) as a collector of organic particles or scraper of plant surfaces
(Guzmán-Soto and Tamaris-Turrizo 2014; Chará et al. 2010), small eyes, legs with
distinctive features, visible abdominal sternites, and a particular genital terminalia
(sternites VIII, IX and gonocoxosternites) indicate an early evolutionary state.
A thorough analysis reveals significant differences between the two new genera
and the genus Tonerus. Coloration patterns, dorsal and lateral view morphology
allowed for the clear distinction of the nine species. Interspecific characters
suggested the presence of two new genera. Tegumentary similarities point to a tribal
distinction between the three genera: all species present flat isodiametric epithelial
cells, similar to those of squamous tissue, which gives them a microrreticulated
texture on their surface. This shared feature confirms that the three genera belong to
a tribal taxon different from Noterini.
The Tonerini Tribe is retributable
According to Baca et al. (2017), DNA-based phylogenetic analyses do not
place the Tonerini and Noterini tribes as separate monophyletic clades. This implies
that they share an exclusive common ancestor and that both tribes possess numerous
similar morphological features, suggesting synonymy between them. However,
while Tonerini and Noterini share some similarities, there are distinctive
morphological features that refute their synonymy.
The mandibles of Tonerinus gen. nov. and Tonermorpha gen. nov. (the author
assumes they share the same character with the genus Tonerus) are not specialized
for predation like those of Noterini. Instead, Tonerinus gen. nov. and Tonermorpha
gen. nov. possess a robust apical molar for scraping or cracking shells, and a square
second molar with a rough or granulated surface for grinding. Additionally, at the
base of the mandible, Tonerinus gen. nov. and Tonermorpha gen. nov. have a row of
fine spines on the basal third for scraping or picking over the surface of rocks,
decaying wood, and even dead leaves. Lastly, Tonerinus gen. nov. and Tonermorpha
gen. nov. possess a lacinia with coarse spines for scraping or collecting fine
particles.
Dos nuevos géneros Tonerini.
García Ramírez.
106
The differences in jaw structure and the presence of specific features such as
spines and the spiny lacinia demonstrate that Tonerini and Noterini are not
synonymous.
The species of Tonerinus gen. nov. and Tonermorpha gen. nov., as well as
Tonerus wheeleri, are characterized by a robust build with large, wide heads.
However, unlike members of Noterini, they have small eyes relative to their body
size. This characteristic is not favorable for predation, as it limits visual capacity
and, therefore, hunting accuracy and success. In contrast, the large eyes of Noterini
provide them with better vision, allowing them to detect and pursue prey more
effectively.
The tibiae of the genera Tonerinus, Tonermorpha, and Tonerus are
considerably wide and have a prominent apicodorsal margin. This margin is
bordered by long, thick spines, the number and size of which vary among the three
genera: Tonerinus has strong spines, Tonerus has moderate spines, and
Tonermorpha has sparse spines. At the apex of the tibia, a slightly curved spur is
observed, in contrast to the robust, curved hook-shaped spur present in noterines.
This latter spur has a very sharp apex, with the exception of the genus Mesonoterus
Sharp (Miller 2009).
Regarding their legs, Tonerinus and Tonermorpha share short and wide femurs
and tibias, with a spiny appearance and retractable spines on the tibias. However, the
tarsal claws show notable differences between the two genera. In Tonerinus, the
claws are short and curved in the pro- and mesotarsus, while in the metatarsus they
are straight and cover the length of the fifth tarsomere. In Tonermorpha, the claws
are long, equivalent to the combined length of the fifth and fourth tarsomere. In both
genera, they are slightly curved in the pro- and mesotarsus and straight in the
metatarsus.
Tonerinus species have ridges on the ventral margin of the front claws, an
invagination on the middle tarsal claws and a simple shape on the metatarsal claws.
In contrast to Tonermorpha species, all claws on the tarsi are simple. It is
noteworthy that Miller (2009) does not mention any features on the tarsal claws of
Tonerus wheeleri. In contrast, the claws of noterinids are serrate and have different
formulae depending on the genus: (2-3-3), (2-2-3), (3-3-3), (3-3-2) or (1-1-1). The
shape of the teeth on each claw also varies.
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The metatibial spurs of the genus Tonerinus are characterized by their distinctly
serrated edges. The anterior claw exhibits double parallel serrations, while the
posterior claw only displays a serrated margin with long, slender teeth. In the case of
Tonermorpha, only the posterior spur possesses a serrated edge. In contrast, Tonerus
wheeleri exhibits simple metatibial spurs. Unlike Noterini, the metatibiae of
Tonerinus, Tonermorpha, and Tonerus lack swimming hairs on their anterior and
posterior margins, indicating their unsuitability for swimming. Conversely, Noterini
does possess swimming hairs on its metatibiae, which facilitates its locomotion in
water. The species of Tonerinus, Tonermorpha, and Tonerus are more adapted to
locomotion on soils with a thin film of moisture. It is presumed that, due to the
condition of long claws, the species of Tonermorpha are better adapted to walking
clinging to rocky ground in streams.
The genital terminalia of Tonerinus are primitive in structure compared to
Noterini. The VIII sternite is simple, while the IX sternite is short and broad. The
gonocoxosternites have sparsely lobed apodemes and are absent in some species. In
the genitalia of the female, the laterotergites are short, slender and L-shaped bilobed.
The gonocoxae, on the other hand, are very small, flattened with pointed apices (in
Tonerus they are thick with rounded apices). In contrast, the Noterini have a large
pointed or lobed gonocoxal apex, which markedly differentiates their genitalia from
those of Tonerini.
Tonerus, Tonerinus and Tonermorpha species are endemic and found in a
single region of the Venezuelan Amazon, while Noterini has a much wider
geographical distribution. This notable difference in distribution is an additional
factor supporting the distinction between the two taxa.
In conclusion, the morphological differences and geographical distribution of
Tonerini and Noterini suggest that they are not synonyms. The synonymy proposal
of Baca et al. (2017) has not been accepted by all aquatic beetle specialists. Some
authors argue that the molecular analysis presented is not conclusive and that the
two tribes can still be distinguished based on specific morphological characters.
Specialized jaws ??
Tonerinus maxillae are characterized by robust laciniae with spines of variable
size, giving them a rough and rugged texture. This morphological particularity sug-
Dos nuevos géneros Tonerini.
García Ramírez.
108
gests one adaptation to scrape surfaces such as rocks or decaying wood present in
surface runoff. Its use extends to biofilm, an algal film that forms on wet rocks, in
areas where the water layer varies between 5 and 10 mm thick (Guzmán-Soto and
Tamaris-Turrizo 2014; Chará et al. 2010). The combination of robust lacinia and
strong spines would allow Tonerinus, Tonermorpha, and presumably, Tonerus
wheeleri species to rasp these surfaces efficiently, accessing an alternative food
source to predation.
The jaws of Tonerinus and Tonermorpha have a series of morphological
characteristics that differentiate them from those of predatory species and adapt
them to a diet based on organic detritus. In the apical region, we find a robust and
specific molar for shredding or scraping coarse and hard particles. This feature
contrasts with the sharp and strong molars present in predatory species. In the
middle region, a coarse second molar is observed with an unrounded, rough or
granulated rectangular apical surface as a series of small coarse teeth. These teeth
are presumably used to demolish or pulverise coarse particles of detritus. On the
basal part of the mandible, there is a long, compact row of fine spines distributed in
a comb-like pattern. These spines could be used to collect small particles on the
surface of moist soil or other soft surface.
In general, the morphological features of the mandibles of Tonerinus and
Tonermorpha suggest an adaptation to a non-predatory diet based on detritus
ingestion. This adaptation would allow species of both genera to exploit an
ecological niche different from that of predatory species, expanding their success in
environments with low prey availability.
All species of Tonerinus and Tonerus wheeleri were collected in specific
habitats on the water film running over the rocks under decaying wood or under the
vegetation layer on the runoff, while Tonermorpha species were collected in a
sweep of the rocky substrate in the littoral zone of the Coromoto River stream. A
striking feature of these species is the absence of swimming hairs on their
metatibiae. Instead, they have large, thick, retractable spines, an adaptation for life
in the leaf litter. This morphological feature indicates that they are not suitable for
swimming, unlike other aquatic beetles.
The retractile spines of Tonerinus and metatibiae allow them to cling to leaf
litter and other substrates present in their microhabitat. They move easily over the
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Vol. 58. Nº 1, Enero-Junio 2024, Pp. 67-111 109
surface of the water film and hold their position against the current of runoff.
Tonerinus metatibiae represent a morphological adaptation to their particular
lifestyle on leaf litter and water film. This adaptation allows them to exploit a unique
ecological niche in shallow water environments with little current.
CONCLUSION
This study represents a significant advance in the knowledge of aquatic
Coleoptera in Venezuela, particularly with regard to the systematics of the Tonerini
tribe. The discovery of two new taxa, along with the genus Tonerus, has allowed for
the recognition of new character states that fit the original taxonomic characteristics
of the tribe. Thus, the reinstatement to its original status as a tribe distinct from
Noterini is justified. These findings open up new avenues of research to better
understand the diversity and evolution of aquatic Coleoptera in Venezuela and the
Neotropical region.
In 2009, only one species of Tonerus wheeleri had been identified from more
than 90 specimens within the then Tonerini tribe. The material analyzed in this
research, collected in 2004, was not part of the original study of Tonerus. The
analysis of coloration and habitus, characteristics that show great variability among
the three genera, allowed us to recognize that the nine species described later were
nothing more than different forms of Tonerus wheeleri, that is, hidden species within
what was until then considered a single species. This hidden diversity led to the
proposal of two new genera: Tonerinus gen. nov. and Tonermorpha gen. nov. These
two new taxa, although less abundant than Tonerus, present greater morphological
diversity. Overall, this evidence highlights the need for a thorough revision of the
Tonerus wheeleri species in all available collections.
The findings of this study are highly relevant to the reinstatement of the
original status of the Tonerini tribe in the near future. This finding significantly
expands the knowledge on the taxonomy of the tribe Tonerini and its relevance for
understanding the biodiversity of aquatic beetles of the family Noteridae. The novel
information obtained on the diversity and distribution of the Tonerus genus group in
Venezuela will significantly strengthen the understanding of the systematic and
evolution of these aquatic beetles within the suborder Adephaga.
110
Dos nuevos géneros Tonerini.
García Ramírez.
Note: Specimens of Tonerus wheeleri are not available at the Arthropod Museum
of the University of Zulia (MALUZ) in Venezuela because they have not yet been
returned.
ACKNOWLEDGEMENT
I would like to express my deepest gratitude to the following people for their
invaluable collaboration in this research. To Alfredo Briceño, digital design
specialist and entomological researcher at the Centro de Investigaciones Biológicas,
for the elaboration of the illustrations and photographs (except for the satellite
images of the Google Earth programme) that enrich this work. To María Leal,
lecturer at the Faculty of Science, for sharing her knowledge of the mandibular
structure of Ephemeroptera, which was fundamental to the development of this
research. To Teresa Martínez, professor and researcher at the Centro de
Investigaciones Biológicas, for her meticulous reading and correction of the
manuscript, which contributed significantly to its improvement. Her support and
collaboration were essential for the completion of this study.
LITERATURE CITED
BACA, S. M., E. F.A, TOUSSAINT, K. B. MILLER y A. E. Z. SHORT. (2017).
Molecular phylogeny of the aquatic beetle family Noteridae (Coleoptera: Adephaga)
with an emphasis on data partitioning strategies. Molecular Phylogenetics and
Evolution.107 (2017): 282–292.
CHARÁ-SERMA, A. M., J. D. CHARÁ, M. C. ZUÑIGA, G. X. PEDRAZA y L. P.
GIRALDO. (2010). Clasificación trófica de insectos acuáticos en ocho quebradas
protegidas de la ecorregión cafetera colombiana. Universitas Scieniarum. 15(1): 27-
36.
GARCÍA, M y A. BRICEÑO. (2023). Revision of the burrowing beetle genus
Llanoterus García and Camacho, 2018 (Coleoptera: Noteridae: Noterini). Part I. Bol.
Centro Invest. Biol. 57(2): 204-271.
GARCÍA M., A. VERA, C. BENETTI y L. BLANCO. (2016). Identificación y
Clasificación de los microhábitats de agua dulce. Acta Zoológica Mexicana. 32(1):
12-31.
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GUZMÁN-SOTO, C. J. y C. E. TAMARIS-TURIZO. (2014). Feeding habits of
inmature individual of Ephemeroptera, Plecoptera and Trichoptera from middle
reaches of a tropical mountain stream. Revista de Biología Tropical, San José.
62(Suppl. 2): 169-178.
MILLER, K. B. (2009). On the systematics of Noteridae (Coleoptera: Adephaga:
Hydradephaga): phylogeny, description of a new tribe, genus and species, and
survey of female genital morphology. Systematics and Biodiversity. 7: 191–214.
ISSN2477-9458
BOLETIN
DEL CENTRO DE INVESTIGACIONES BIOLÓGICAS
AN INTERNATIONAL JOURNAL OF BIOLOGY
PUBLISHED BY THE UNIVERSITY OF ZULIA, MARACAIBO, VENEZUELA
Vol.58, No1, Pp. 1-157, January-June 2024
Phenotype of resistance into MLSB and structural typing of the
Staphylococcal Cassette Chromosome mec (SCCmec) in
methicillin-resistant S. aureus coming from hands of food
handlers
Víctor Pico-Bracho, Jhoandry Rivera-Salazar, Velina Aranaga-
Natera, Isabel Mujica de Fernández, Yoalaimis La Paz-Delgado e
Irene Zabala-Díaz…...……………………………………………….
1
Paracymus from Venezuela (Coleoptera: Hydrophilidae:
Laccobiini), Part VII: Record of six new species.
Mauricio García Ramírez…….……..………………….......................
20
Influence of the hydrological regime on the composition of mangrove
sediments in El Tablazo Bay (Maracaibo System).
Flora Barboza, Ana Marta Francisco, Jacinto Sánchez y Ernesto
Medina……........................................................................................
45
Descubrimiento de dos nuevos géneros de escarabajos acuáticos
detritívoros Toneroides, en el Amazona venezolano
(Coleoptera: Noteridae: Noterinae).
Mauricio García Ramírez…………….……………………….……..…....
67
Scientific Notes.
New observations and expansion of the altitudinal range of wood
stork Mycteria americana (Linnaeus, 1758) in the Venezuela
andes, suggest movements between bioregions.
Luis A. Saavedra, Alexis Araujo-Quintero y Carla I. Aranguren…..
112
Notes on the genera Suphisellus Crotch, 1873 and Suphisellus
Zimmermann, 1919, a cocktail of encrypted Suphiselloides
genera (Coleoptera: Noteridae: Noterinae: Noterini).
Mauricio García Ramírez………………………..…………..………
124
Instrucciones a los autores……………….…..…………………………
138
Instructions for authors……………….…..………………………...……
148
