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Zoqueichthys carolinae gen. and sp. nov. 735
A Cenomanian aipichthyoid sh (Teleostei, Acanthomorpha) from
America, Zoqueichthys carolinae gen. and sp. nov. from El Chango quarry
(Cintalapa Member, Sierra Madre Formation), Chiapas, Mexico
Jesús Alvarado-Ortega1,* and Bruno Andrés Than-Marchese2
1 Instituto de Geología, Universidad Nacional Autónoma de México; Circuito de la Investigación S/N,
Ciudad Universitaria, Coyoacán, D.F., 04510, Mexico.
2 Museo de Paleontología “Eliseo Palacios Aguilera” SEMAHN, Calzada de los Hombres Ilustres S/N,
Colonia Centro, Tuxtla Gutiérrez, Chiapas, Mexico.
*alvarado@geologia.unam.mx
ABSTRACT
Zoqueichthys carolinae gen. and sp. nov. is described based on three specimens collected at the El
Chango quarry, an outcrop of Cenomanian marine limestones belonging to the Sierra Madre Formation
located in the Municipality of Ocozocoautla de Espinosa, near Tuxtla Gutiérrez, Chiapas State, southern
Mexico. This species is identied as a new member of the superfamily Aipichthyoidea because it possesses
a combination of characteristics previously recognized as diagnostic for this group. This species differs
from other aipichthyoides in skeletal and meristic characteristics not previously reported in this superfamily
including a completely laminar supraoccipital crest involving the participation of the supraoccipital and
frontal bones, and lacking anterior thickening or spine; a pelvic n with eight rays, and pectoral n with
twelve rays. Zoqueichthys carolinae is a basal member of the family Aipichthyoididae because it shows
putative primitive characteristics including a comparatively high number of pelvic n rays, three epurals,
hypurals one and two are autogenous and in addition a complex supraoccipital crest. Zoqueichthys
carolinae is the rst aipichthyoid sh found in marine Cretaceous sediments deposited in the western
Tethys Sea of southern North America.
Key words: Aipichthyoidea, Aipichthyoidae, Aipichthyoididae, Cenomanian, Mexico.
RESUMEN
Zoqueichthys carolinae gen. y sp. nov. es descrito a partir de tres ejemplares colectados de la
cantera El Chango, un aoramiento Cenomaniano de calizas marinas pertenecientes a la Formación
Sierra Madre, ubicado dentro del Municipio de Ocozocoautla de Espinosa y próximo a Tuxtla Gutiérrez,
Chiapas, sureste de México. Esta especie es miembro de la superfamilia Aipichthyoidea porque presenta
los rasgos diagnósticos de la misma. Esta especie es diferente de otros aipichthyoideos en rasgos
merísticas y esqueléticos que anteriormente no habían sido reportadas en la superfamilia; su cresta
supraoccipital es totalmente laminar, está formada por los huesos supraoccipital y frontal, y carece de un
engrosamiento o espina anterior; además sus aletas pélvica y pectoral están formadas por ocho y doce
radios respectivamente. Zoqueichthys carolinae es un miembro basal de la familia Aipichthyoididae ya
que presenta un elevado número de radios en la aleta pélvica, tres epurales, sus dos primeros hipurales
Revista Mexicana de Ciencias Geológicas, v. 29, núm. 3, 2012, p. 735-748
Alvarado-Ortega, J., Than-Marchese, B.A., 2012, A Cenomanian aipichthyoid sh (Teleostei, Acanthomorpha) from America, Zoqueichthys carolinae
gen. and sp. nov. from El Chango quarry (Cintalapa Member, Sierra Madre Formation), Chiapas, Mexico: Revista Mexicana de Ciencias Geológicas,
v. 29, núm. 3, p. 735-748.
Alvarado-Ortega and Than-Marchese
736
INTRODUCTION
Otero and Gayet (1996) named the superfamily
Aipichthyoidea. This group of basal acanthomorphs in-
cludes highly specialized small marine shes, characterized
by their rounded bodies that bear large unpaired ns armed
with anterior spines and a pelvic n in thoracic position,
and by their short and high heads showing a hypertrophied
supraoccipital crest (“sagittal crest” of Otero and Gayet,
1996) lacking the antorbital bone. Currently this group
comprises two families, Aipichthyidae Patterson (1964)
and Aipichthyiodidae and Gayet (1980a).
Until now, the superfamily Aipichthyoidea comprised
primarily Cenomanian shes discovered in eastern Tethyan
outcrops of Europe and the Middle East, and included
ve species of Aipichthys: the type species A. pretiosus
Steindachner, 1850, from Comen, Croatia (Dalmatia in
Patterson, 1964: 303); A. minor Pictet, 1850, from Hakel
(=Haqil) and Hadjula, Lebanon; A. velifer Woodward,
1901, and A. oblongus Gayet, 1980b, from Hakel; and A.
nuchalis Dixon, 1850, from Sussex, England; as well as
Paraipichthys lusitanicus Gaudant, 1978, from Laveiras,
Portugal; two species of Aipichthyoides described by
Gayet (1980a) under the names A. galeatus and A. for-
mosus, from Ein Yabrud, Israel; Freigichthys elleipsis
Otero, 1997, from marine outcrops of Hadjula (=Hgula),
Lebanon; and Aspesaipichthys cavaensis Taverne, 2004,
from Campanian–Maastrichtian of Nardo, Italy.
Although the first phylogenetic analysis of
Aipichthyoidea performed by Otero and Gayet (1996)
supports the inclusion of Paraipichthys in the family
Aipichthyidae, relationships within the superfamily be-
came unclear after the discovery of Freigichthys (Otero,
1997, g. 5). The inclusion of Aspesaipichthys in family
Aipichthyiodidae, as Taverne (2004) intended, requires
further examination. Beyond these works, the phylogeny
of this superfamily has not been re-examined.
The aim of this paper is to describe and discuss
the relationships of Zoqueichthys carolinae gen. and sp.
nov., an aipichthyoid sh known from three beautifully
preserved specimens collected in Cenomanian strata
of El Chango quarry, Chiapas, southern Mexico. Prior
to the present paper, Aipichthyoidea were known as a
marine Late Cretaceous group restricted to the eastern
Tethys Sea domain of Europe and the Middle East. The
discovery Zoqueichthys has important implications because
son autógenos, y su cresta supraoccipital es compleja. Esta especie es el primer pez de la superfamilia
Aipichthyoidea encontrado en rocas marinas del Cretácico depositadas bajo el Dominio Occidental del
Mar de Tetis.
Palabras clave: Aipichthyoidea, Aipichthyoidae, Aipichthyoididae, Cenomanian, México.
it complements the taxonomic diversity of the group and
improves our knowledge of the temporal and geographic
distribution patterns of these primitive acanthomorphs. The
presence of Zoqueichthys in Mexico suggests that the early
evolution and distribution of superfamily Aipichthyoidea
took place within a wider and more complex geographical
setting that included both western and eastern domains of
the Tethys Sea.
El Chango quarry is an outcrop of limestones near
Tuxtla Gutiérrez, Chiapas State, southern Mexico (Figure
1). This site is located at 16º34’14.9”N and 093º16’12.7”W
within Ocozocautla de Espinosa Municipality. Since 2006
a well-preserved and diverse fossil assemblage has been
collected in the laminated and monotonous sequence ex-
posed in this outcrop, which pertains to the Sierra Madre
Formation (Ovalles-Damián et al., 2006; Alvarado-Ortega
et al., 2009; among others). According Vega et al. (2007)
these strata accumulated within an estuarine or salty lagoon
with ephemeral freshwater inux.
The fossil assemblage thus far collected at El Chango
includes shes, plants, mollusks, crustaceans, and insects.
Recently, Alvarado-Ortega et al. (2009) found that the shes
found at El Chango (and El Espinal) indicate a Cenomanian
age. The shes are the best-preserved fossils at El Chango.
The specimens are tightly laterally compressed, usually
complete and articulated, and often show parts of soft tissues
(muscles) and stomach contents phosphatized.
Böse (1905) described the Cretaceous platform
carbonate series that includes int-bearing dolomite and
limestone exposed along the Central Chiapas depression,
where El Chango quarry is found, as “Cretaceous limestones
with rudists”. Although Wiebe (1925) named these strata
the San Cristóbal Formation, Nutall (1929, in Salas, 1949)
suggested the name Sierra Madre Formation that is currently
in use. Salas (1949) grossly divided this formaton in two
sequences, an Albian–Cenomanian unit of limestones
having sugary aspect and apparently barren of fossils, and a
Turonian unit of limestones bearing int and rudists. Later,
González (1963) formalized this suggestion, recognizing
these units as the Cantelhá and the Jolpabuchil members
respectively.
The discovery of microfossils during petroleum ex-
ploration contributed to re-ordering the units of the Sierra
Madre limestones (Figure 1). Sánchez-Montes de Oca
(1969, 1973), Álvarez-Mena (1975), and others, suggested
that these actually comprise two formations, at the base the
Zoqueichthys carolinae gen. and sp. nov. 737
mens were examined for comparative purposes; all belong
to the Otto Haas Collection, housed in the Bayerische
Staatssammling für Paläontologie und Geologie, München,
Germany, and catalogued under the following numbers.
Aipichthys minor: specimens 177, 178, and 180, from
Hakel, Lebanon. Aipichthy sp.: specimen Nr. 1984 I 230,
from Hakel, Lebanon. Additional comparative data were
obtained from available literature.
SYSTEMATIC PALEONTOLOGY
Cohort Acanthomorpha Rosen, 1973
Order incertae sedis
Superfamily Aipichthyoidea Otero and Gayet, 1996
Family Aipichthyoididae Gayet, 1980a
Emended diagnosis. Aipichthyoidea with high, long, and
complex supraoccipital crest that extends over the midline
of the skull from the nasal area to occiput, with the anterior
and posterior edges rounded and convex; and the supraoc-
cipital and frontal bones forming the supraoccipital crest.
Genus Zoqueichthys gen. nov.
Type species. Zoqueichthys carolinae sp. nov.
Etymology. The generic name derives from “Zoque”, a
Sierra Madre Formation made up of the Cantelhá (Lower-
middle Albian) and Cintalapa (Cenomanian) members, and
at the top the Jolpabuchil Formation (Turonian–Santoniano).
Steele (1986) and Waite (1986) subdivided the Sierra Madre
limestones into lithologic and biostratigraphic units (herein
these are simplied as 1 to 4 in Figure 1), which include
Upper Albian–Santonian strata. Rosales-Domínguez et al.
(1994, 1997) carried out microfossil analyses along the
Suchiapa River and on the Ocozocuautla–Cintalapa road
that also indicate an Albian–Santonian age for these strata.
The fossiliferous strata at El Chango quarry represent a
section within the 700 to 1600 meter thick sequence of
the Cenomanian Cintalapa Member of the Sierra Madre
Formation.
MATERIAL AND METHODS
Preparation methods. The specimens were prepared in
baths of 5–10% acetic acid solution and hardened with
Plexygum. Pin vises and needles were used under a bin-
ocular microscope to remove matrix from the skeletons.
Anatomical abbreviations. Most of the abbreviations used
in Figures 2 through 7 follow Patterson (1964), Gaudant
(1978), Otero and Gayet (1995, 1996), and Stiassny and
Moore (1992).
Comparative material analyzed. The following speci-
Figure 1. Map of the Sierra Madre Limestone (Cretaceous) in the Ocozocoautla-Tuxtla Gutérrez area showing the position of El Chango quarry and El
Espinal quarry (another fossil sh locality in this geological unit) and the age suggested for different geological sections.
Alvarado-Ortega and Than-Marchese
738
native group of people in Chiapas, and “Ichthys” or sh in
Greek. It means “the sh of the Zoque people”.
Diagnosis. As in the type species, below.
Zoqueichthys carolinae sp. nov.
(Figures 2–6, Table 1)
Etymology. The species name honors Mrs. Carolina Elvira
Marchese y Tronkar, mother of second author of present
paper.
Holotype. IHNFG-2224, a complete specimen exposing the
right lateral side and having total and standard length (SL)
of 69 and 49 mm respectively (Figure 2).
Referred material. IHNFG-2225, a specimen in part and
counterpart, incompletely preserved, and exposing its right
lateral side; it lacks the whole dorsal area from the vertebral
column and the caudal n; its estimated SL is about 58
mm. IHNFG-2270, a specimen without the caudal n and
exposing its left lateral side, with an estimated SL of about
58 mm.
Diagnosis. Aipichthyoididea with an ovoid body, maximum
known standard length of 49 mm and maximum body height
representing 60% of SL; its supraoccipital crest is high and
long, projecting over the midline of the skull from the oc-
ciput to the mesethmoid area, with its anterior and posterior
edges rounded and convex, and formed by the supraoccipital
and frontal bones; the vertebral column includes 11 abdomi-
nal and 16 caudal vertebrae; dorsal n includes three anterior
spines and 24 segmented and branched rays supported by
26 pterygiophores; anal n is formed by three spines and 17
branched and segmented rays supporting 18 pterygiophores
including the rst pterygiophore that is transformed in the
hemaxanal complex; pectoral n includes 12 segmented
and branched rays; pelvic n in thoracic position has eight
rays; all pelvic rays are segmented and branched except for
the rst that is unbranched; 19 principal caudal n rays,
including two unbranched and 17 segmented and branched
rays plus 10 and eight dorsal and ventral procurrent rays
(10-I+9—8+I-8).
Locality. El Chango quarry, Ocozocoautla de Espinosa
Municipality, near Tuxtla Gutiérrez, Chiapas State, southern
Mexico.
Stratigraphic horizon. Cintalapa Member (Cenomanian)
Sierra Madre Formation.
Description.
General proportions. Table 1 summarizes the measurements
of the specimens described herein. Zoqueichthys carolinae
gen. and sp. nov is a centimetric diamond-shaped sh with a
deeply forked homocercal tail, a gently rounded dorsal trunk
prole continued by a large triangular supraoccipital crest,
and a ventral border also rounded but having an abdominal
horizontal at prole. In this species the maximum height of
the trunk is about 60% of the standard length (SL); the head
is about 1.75 times higher than long, with its length being
about 34.4–36% of the SL. Its trunk shows abdominal and
postabdominal regions with the same length. The pectoral
n is located in the middle of the body between the vertebral
column and the ventral body border. The pelvic n position
is thoracic. Anal and dorsal ns are long, the former being
50% and the latter being 26% of SL.
Figure 2. Holotype of Zoqueichthys carolinae gen. and sp. nov., specimen
IHNFG-2224, from El Chango quarry, near Tuxtla Gutiérrez, Chiapas,
Mexico.
IHNFG-2224 IHNFG-2225 IHNFG-2270
Total length 69 (138) ? ?
Standard length
(SL)
49 58 58?
Head length 18 (36) 20 (34.4) 21
Head height 28 (56) ?28
Dorsal length 25 (50) ? ?
Predorsal
length
20 (40) ? ?
Anal length 13 (26) 15 (25.8) ?
Preanal length 35 (70) 41(70) 36
Prepelvic
length
22 (44) 25 (43) 25
Maximum
body height
30 (60) ? 36
Caudal
peduncle
height
7 (14) ?11?
Vertebral
column
11abd+16(4)
ca
11+?(?) 11+3?(?)
Dorsal n III+24/(26) III+?/(26) ?
Anal n III+17/(I+17) III+17/(I+17) ?
Table 1. Meristic data for the specimens of Zoqueichthys carolinae gen.
and sp. nov. All measurements are in millimeters and the proportions
enclosed between parentheses are expressed as percentage of standard
length (SL). Counts of vertebrae include abdominal (abd) and caudal (ca)
vertebrae, as well as vertebrae with hemal and neural spines supporting the
caudal n skeleton (between parentheses and as part of caudal vertebrae).
Counts for dorsal and anal ns include spines (in Roman numbers), soft
rays (in Arabic numbers), plus pterygiophores (enclosed in parentheses).
Anal perygiophores are divided in the rst pretygiophore modied in the
hemaxanal complex (I) and normal pterygiophores (17).
10 mm
Zoqueichthys carolinae gen. and sp. nov. 739
Figure 3. Head and anterior part of the body of Zoqueichthys carolinae
gen. and sp. nov., specimen IHNFG-2224 with contrast heightened with
smoke of magnesium.
Skull. The skull in Zoqueichthys carolinae gen. and sp. nov.
is triangular and 1.5 times longer than high, with its ocular
diameter representing a third of its length (Figures 3). The
mesethmoid is a well ossied and complex bone; it has a
ventral arm that forms the anterior edge of the nasal capsule
and joins with the parasphenoid and probably the vomer. The
dorsal section of the mesethmoid has an anterior process,
and joins with the ventral anterior end of the frontal. The
nasal is a thick tubular bone; in IHNFG-2224 this bone is
located in front of the mesethmoid but in IHNFG-2225 it
is in its natural position over the nasal capsule. The lateral
ethmoid is another coarse bone. This rectangular stilt forms
the anterior and posterior walls of the orbit and nasal capsule
respectively, and joins with the frontal bone dorsally and
the parasphenoid ventrally.
The frontal is large and roofs the anterior two thirds
of the skull, including all the ethmoid and ocular areas.
Dorsally this bone has a triangular vertical laminar wing
that forms the anterior part of the supraoccipital crest and
joins with the supraoccipital section of the crest at the level
of the rear of the orbital (Figure 4). The supraoccipital bone
occupies the rest of the skull roof, separates the parietals
and joins the rear of the frontal bone. This bone also has a
vertical laminar wing that forms the posterior section of the
supraoccipital crest. This crest is triangular with a pointed
dorsal border forming an angle of about 90 degrees and
located near the occiput, at two thirds of its length from its
anterior end. The lateral surface of the central area of the
supraoccipital crest is ornamented with large, shallow, and
parallel ridges that run from the dorsal border of the crest to
the area where the supraoccipital and frontal meet.
The parietal is an elongate bone located parallel to and
below the supraoccipital (Figure 4). The posttemporal fossa
is located in the anterior end of this bone but is roofed by
the posterior expansion of the frontal. The lateral wall of the
skull is occupied mainly by the pterotic, which forms the
dorsal roof of the hyomandibular fossa. Part of the sphenotic
bone is hidden below Infraorbital 5. The limits and sutures
of other postocular skull bones are not discernible in the
available specimens.
The parasphenoid is a large and slightly upward-
curving stilt, toothless and probably triangular in cross
section (Figure 4). The orbital cavity is partially occupied
by the orbitosphenoid and pleurosphenoid bones that project
from its internal dorsal surface, and by anterior projections
of basisphenoid bones that join the ocular section of the
parasphenoid. In IHNFG-2224 a smooth laminar ovoid
bone of uncertain identity occupies the anterior orbital
cavity (Figure 4).
The supraorbital canal runs along the whole frontal
bone enclosed in tubes that open near the anterior end of this
bone as well as in wide conspicuous pores located above and
below the orbit. Some small pores of this canal also open
in the base of the supraoccipital crest at its highest point.
Lower jaw. The lower jaw involves the dentary, angular,
and dermarticular bones (Figure 4). The lateral prole of the
jaw is chicken-legged in shape, having an almost straight
alveolar border and the articular process for the quadrate
located below the middle of the ocular orbit. The dentary
bone is triangular with the posterior edge bifurcated, the
anterior end narrow, and the ventral border showing an
anterior hook-shaped ventral process. The dentary forms
the entire alveolar border of the jaw, which is tted with a
wide patch of uniform small rounded teeth.
The angulo-articular is a coarse triangular bone
strongly attached to the posterior bifurcated edge of dentary.
The coronoid process area in this bone is obscured in the
available specimens. The articular process rises from the
posterior vertical edge of the angulo-articular. The joint
between the lower jaw and the head of the quadrate is visible
laterally (Figure 4). The mandibular canal runs along the
ventral border of lower jaw, crossing and opening through
large pores near the ventral edge of the angulo-articular
bone, 5 or 6 of the pores being in the former and 2 or 3 in
the latter.
Upper jaw. The premaxilla is a long complex bone that
forms practically all the occlusal border of the upper jaw.
The height of its triangular anterior ascendant process is a
third of the length of its alveolar limb. The premaxilla has
a stout and rod-like articular process, the height of which is
2 mm
Alvarado-Ortega and Than-Marchese
740
about half of the ascending process. The alveolar surface of
this bone bears a patch of numerous uniform small rounded
teeth that resemble those on the lower jaw.
The maxilla is a toothless long rectangular bone with
an expanded rounded posterior end and an anterior stout
rod-like articular process. The maxilla articular process
joins the premaxilla articular process ventrally and probably
the vomer dorsally. A large long drop-shaped posterior
supramaxilla lies over at least the posterior half of the
premaxilla dorsal border (Figure 4). In IHNFG-2225 it is
clear that Zoqueichthys carolinae has two supramaxillae,
the anterior one being of a similar shape and size to the
posterior one.
Infraorbtal series. The ocular orbit is incompletely sur-
rounded by ve infraorbitals, which cover it from the eth-
moid area to the postocular lateral part of the skull, passing
over the hymandibula and entopterygoid (Figure 4). The
infraorbital 1 (also called the lacrimal bone by Gayet, 1980a:
24) is the largest in the series; it covers a large part of the
nasal area. Infraorbital 1 is subrectangular in lateral view,
Figue 4. Reconstruction of the head and anterior part of the body of Zoqueichthys carolinae gen. and sp. nov. based on Figure 3. Abbreviations:
ang,angulao-articular; br, branchiostegal; cl,cleithrum; cor, coracoid; den, dentary; dfr , dorsal n ray; dfsp, dorsal n spine; dpt, dorsal pterygiophore;
ecp,ectopterygoid; enp, endopterygoid; epl, epipleural; epn, epineural; fr, frontal; hy, hyomandibular; iop, infraopercle; le, lateral ethmoid; mes, mesethmoid;
mpt, metapterygoid; mx, maxilla; na, nasal; obs, orbitosphenoid; op, opercle; pa, parietal; pcld; postcleithrum dorsal; pclv; postclethrum ventral; pd,
predorsal; pfr, pectoral n ray; pmx, premaxilla; pop, preopercle; psp, parasphenoid; pt, posttemporal; pto, pterotic; pv; pelvic bone; q, quadrate; sca,
sapula; scl, supracleithrum; smx, supramaxilla; soc, supraoccipital; sop, subopercle; spo, sphenotic; stp, suprattemporal; v, vertebral centra (numbered in
anteroposterior order); 1-5; infraorbital bones; arrows enclose the radial bones.
Zoqueichthys carolinae gen. and sp. nov. 741
located below the infraopercle and behind the lower jaw.
Although the rst branchiostegal ray is thin, short, almost
straight, and thread-like, the others tend to be longer and
are at and spatula-shaped.
Axial skeleton. The vertebral column consists of 27 centra,
11 abdominal and 16 caudal; the caudal centra include 14
preural, the pseudurostylar complex (=fusion of preural 1
and ural 1), and ural 2. All of the centra are subrectangular
in lateral view, and slightly higher than long (Figure 5).
The lateral surfaces of the vertebrae are ornamented with
deep unordered cavities, which are conspicuous in the most
posterior vertebrae.
All neural and hemal arches and spines are fused to
their respective centra except for hemal arches on the last
two preural centra (Figure 6) and the hemal spine in the
rst caudal centrum (=preural centrum 14) (Figure 5). The
condition of the neural on the rst abdominal vertebra is
unknown. These spines are fused with their respective hemal
and neural arches. The posterior ve hemal spines and all
neural spines on caudal vertebrae are oar-shaped; they have
a central bar support of uniform thickness and a couple of
bony wings on the tip (Figures 2, 5-6). The large and sharp
neural spines on the rst eight abdominal vertebrae show a
noticeable groove that divides them in two halves. The ribs
are long curved bars that enclose just the dorsal half of the
abdominal cavity. Each rib has a conspicuous longitudinal
hollow. The ribs join the transverse processes on the last
eight abdominal vertebrae, with the height of the processes
increasing with a corresponding decrease of the ribs poste-
riorly. The rst transverse processes are very small whereas
the most posterior are about three times higher (Figure 2).
Epineurals are present in the abdominal and in three caudal
vertebrae; these thread-like bones are about as long as three
abdominal vertebrae and project upward and backward from
the bases of the neural arches (Figure 4). Epipleurals are
clearly associated with the four posterior abdominal and
two caudal vertebrae; these ne bones project backward
and downward from the transverse processes and hemal
spines (Figure 5).
Three high predorsal bones are placed between the
rst neural spine and the occiput (Figure 4). These trumpet-
shaped bones are about two times as long at the top than
at their base. The rst predorsal is the highest and the only
one that is curved; it projects from the dorsal border to
the vertebral column. The second and third predorsals are
practically straight and progressively shorter; the height
of the third predorsal is about two thirds that of the rst.
Regarding the predorsal bones, neural spines, and dorsal n
(see below), it is possible to recognize the predorsal formula
(Ahlstrom et al., 1976; Johnson, 1984) in Zoqueichthys
carolinae is 0+0+0/1+1/1.
Pectoral girdle and n. The supratemporal is a narrow, short
tube-like bone about as long as one abdominal vertebra; it
is placed at an angle between the back of the supraoccipital
crest base and the posttemporal dorsal process (Figure 4).
In lateral external view the posttemporal shows a rounded
almost as high as it is long, and bears a small dorsal process.
Infraorbitals 2 through 4 are rectangular, longer than they
are high, and border the orbit ventrally; whereas infraorbital
5, the smallest rectangular plate in the series, is higher than
it is long and covers the rear of the orbit. The infraorbital
canal runs under a thin rim of bone near the orbital border
of infraorbitals 1 through 5; it opens through small pores
except in infraorbital 1, where this canal branches into ve
to seven enclosed radiating tubes projecting downward.
Hyomandibular bones. Large parts of the suspensorium are
obscured by other bones in all available specimens (Figure
4). The hymandibular is a coarse T-shaped bone with a
wide stout articular head at the top, a large massive axis
ventrally projecting, a short opercular process projecting
backward near the top, and a wide rounded laminar anterior
wing. The opercular anterior process or wing of the hyo-
mandibular bone strongly attaches to the endopterygoid and
metapterygoid. The quadrate is triangular bone, about two
times higher than long, with a stout articular head inclined
forward. The symplectic is a clove-like structure embedded
near the rear of the quadrate.
The large lateral surfaces of the metapterygoid and
endopterygoid are wrinkled, perhaps because these bones
bear teeth internally, in contrast with the dermopalatine and
ectopterygoid that show smooth lateral external surfaces,
suggesting they are toothless. The articular head of the
dermopalatine is hidden below the infraorbital 1.
Opercular series and brachiostegals. The opercle is a at,
ovoid bone with a wrinkled external surface that extends
vertically to cover only the dorsal half of the body height,
below the vertebral column and between the pectoral girdle
and the rest of the head. Its anterior and ventral borders are
almost straight and form an angle of about 65 degrees. The
hyomandibular facet of this bone is located near the top of
its anterior border (Figure 4).
The preopercle is smooth and boomerang-shaped,
gently curved and two times higher than long (Figure 4). The
main axis of both limbs form an angle of 130–140 degrees.
A small rounded notch is present in the anterior edge of this
bone, in the area where two preopercular limbs are joined.
This notch is similar to the hook-like process described and
illustrated in Aipichthys velifer and A. minor by Otero and
Gayet (1996: 324, g. 8A). The preopercular canal is notice-
able; it runs along the middle part of both limbs and opens
through a number of elongate pores, four in the horizontal
limb and another two or three in the vertical limb.
The sickle-shaped subopercle is located all along the
underside of the opercle. In IHNFG-2225 the height of the
anterior ascending process in this bone is almost a quarter
of the opercle height, and its shows a similar wrinkled
external surface. The infraopercle is an inclined at bone,
in lateral view having an orange-slice shape, and barely
overhanging the preopercle posterior margin. The wrinkles
that ornament the infraopercle project almost in parallel
from its dorsal to its ventral margin (Figure 4). In IHFG
2224 there are at least eight branchiostegal rays preserved,
Alvarado-Ortega and Than-Marchese
742
central structure with two processes projecting forward
forming an angle of about 60°, in which the upper process is
just slightly shorter and thinner than the lower one (Figures
3 and 4).
The supracleithrum is at and smooth, and covers the
lateral surfaces of the rst vertebra; in lateral external view
this is an almost ovoid bone that overlaps the dorsal tip of
the cleithrum and joins the posttemporal central region
(Figure 4). The cleithrum is sinuous in shape and extends
from the vertebral column to the ventral edge of the body. In
IHNFG-2225 the cleithrum lateral surface shows the same
shallow parallel ridges that ornament the opercle.
The coracoid is O-shaped with a rounded foramen in
the center; this bone is located behind the cleithrum inec-
Figure 5. Pelvic girdle and anal ns of Zoqueichthys carolinae gen. and sp. nov.; a) and b), photo and line drawing of pelvic bones of specimen IHNFG-2270;
c) and d), photo and line drawing pelvic girdle of IHNFG-2224; e) and f), photo and line drawing of the anal n of IHNFG-2224. c) and e) are contrasted
with smoke of magnesium. Abbreviations: afpr, articular facet for pelvic n ray; afr, anal n ray; afsp, anal n spine; apt, anal perygiophore; cp; central
part of pelvic bone; edw, external dorsal wing; epl, epipleural; evw, external ventral wing; hapu, hemal arch of preural vertebra; heaxc, hemaxanal complex
(= anal pterygiophore 1); hsppu, hemal spine of preural vertebra; iw, internal wing; lp, lateral process; mp, median process; pp, posterior process; pvr,
pectoral n ray; v, vertebral centra (numbered in anteroposterior order); vsc, ventral scute.
Zoqueichthys carolinae gen. and sp. nov. 743
tion area (Figure 4). There is an internal foramen formed
between the ventral and dorsal joints between the coracoid
and cleithrum. There are two postcleithra; the dorsal one
is at, ovoid, and two times higher than long. The ventral
postcleithrum is long and tube-like, and projects from the
dorsal postcleithrum to the dorsal edge of the pelvic n
base. The pectoral n has 12 segmented rays that join the
four autogenous radials, which attach this n to the coracoid
(Figure 4). This n is triangular and covers two thirds of
the abdomen length.
Pelvic girdle and n. The pelvic n is triangular and thoracic
in position. It rises near the middle of the trunk at about
43–44% of the SL (the prepelvic length) and consists of
eight segmented rays. All of these rays except the rst one
are branched. The rst two of these rays are so long that
they reach the base of the anal n. The pelvic girdle con-
sists of a pair of long triangular pelvic bones. The anterior
tips of these bones are in contact with the pectoral girdle.
The pelvic n rays join directly on the external posterior
edge of these bones. There is a long spine-like posterior
process projecting backward from the internal poste-
rior area of each pelvic bone. The pelvic bones join each
other throughout their internal wings and median processes
(Figure 7).
Dorsal n. This n is a long and acuminate structure located
on the posterior half of the back between 40 and 90% of
the SL (Figure 2). It exhibits 24 branched and segmented
rays and three stout anterior spines that are supported by 26
pterygiophores. All spines and rays are articulated one to one
with the pterygiophores, except for the last ray that is not
supported. The rst and second pterygiophores are located
in the space between the rst two neural spines. Each of the
subsequent 8 pterygiophores (3 through 10 in the series) is
located in the interneural spaces. Other pterygiophores (11-
24 in the series) are shorter and in some cases form couples
that t in a single interneural space.
The rst pterygiophore is stout and hook-shaped; it
is three times higher than long and its vertical and horizon-
tal limbs form an anterior angle of 50–53 degrees. In this
pterygiophore the vertical limb consists of a stout central
column anked by two laminar sections located forward
and backward, whereas its horizontal limb is a stout spine
projecting forward. The second pterygiophore is the highest
in the series (being slightly higher than the rst), and as in
all the others has stout columnar and laminar sections. All
subsequent pterygiophores gradually decrease in height
and their laminar sections tend to be smaller; the height
of the last one is about half of the rst one. The laminar
sections of the pterygiophores meet each other, forming
interdigitating or sigmoid junctions, except for the eight
Figure 6. Caudal skeleton of Zoqueichthys carolinae gen. and sp. nov.; a) close-up of specimen IHNFG-2224 with contrast heightened by smoke of mag-
nesium; b) line drawing based on a). Abbreviations: ep, epural; h, hypural; hspu, hemal spine of preural vertebra; nspu, neural spine of preural vertebra;
pc, pseudurostylar complex; ph, parhypural; pu; preural vertebra; sc, scale; st, stegural (= ural 1 expanded); u, ural; un, uroneural; black arrows, enclose
the principal n rays; white arrow, shows the caudal fenestra.
Alvarado-Ortega and Than-Marchese
744
most posterior ones where these sections are so small that
they do not meet each other. The vertical limbs of the rst
and second pterygiophores are in the space between the rst
neural spines of vertebrae 1 and 2 (Figure 4).
The dorsal n spines are smooth structures projecting
backward. The height of the rst and smallest spine is one
half and one fth the height of the second and third spines
respectively. The rst and second spines are stout whereas
the tip of the third spine is clearly thinner. The rst three
anterior soft rays are higher than the posterior spine and form
the anterior acuminate end of the dorsal n; here the rst
ray is about two times higher than this spine. The height of
the remaining dorsal rays (2 through 24 in the series) tends
to decrease, and in the most posterior one this is about one
fourth the height in the rst ray.
Anal n. This n is strip-like with a small acuminate ante-
rior border. It is relatively short, opposes the posterior half
of the dorsal n, and is placed between 70 and 96% of the
SL (Figure 2). This n involves 17 branched and segmented
anal rays and three stout anterior spines supported by 18
pterygiophores, the rst of these represents the hemaxanal
complex described by Gayet (1980a). The rst two spines
join the hemal complex and the last spine and all of the soft
rays are articulated one to one with the remaining pterygi-
ophores (Figure 5).
The rst anal pterygiophore or hemaxanal complex is
sickle-shaped, two times higher than long, with two limbs
forming an angle of about 80 degrees and an anterior round-
ed concave edge. The vertical limb of this complex projects
into the space between the rst hemal spine (attached on
preural centra 14) and the last couple of ribs (which are
attached to the parapophysis of vertebra 11). It consists
of a thick column running along its entire height and two
laminar sections placed forward and backward and reach-
ing only half the height. In contrast, its horizontal limb has
a thick ventral edge that joins the anterior laminar section
where the rst two anal spines are strongly attached. The
second pterygiophore is about the height of the vertical limb
of the hemaxanal complex. The posterior pterygiophores
gradually decrease in height; the second one slightly more
than half the height of the rst pterygiophore, and the last
one slightly less than half. These also show a thick middle
column anked by two laminar sections of equal size that
extend anteriorly and posteriorly to meet the laminar sec-
tions of adjacent pterygiophores forming almost straight
joins. In the last ve pterygiophores these laminar sections
tend to be smaller and fail to join each another.
The spines are stout structures projecting backward
with the lateral surfaces ornamented by ne longitudinal
striations (Figure 5). The rst and smallest spine is tri-
angular, two times longer than it is high, and attached to
the anterior pterygiophore by simple contact. The second
is a better developed spine, almost as long as the rst but
almost three times higher; it joints the rst petrygiophore
through a small process developed at its base, and with the
rst and third spines through rounded anterior and posterior
processes located near the base. The third and longest spine
is almost two and a half times higher than the second one;
this joins the second pterygiophore and the rst soft ray
through a pair of small processes present in its base and
rear respectively; it also joins the second spine through a
small facet formed in the anterior edge and near the base,
where the posterior process of the second anal spine sits. At
least the rst two anal n rays are slightly longer than the
anal spines and form the acuminate anterior end of this n.
Although the height of the remaining anal rays (3 through
17 in the series) progressively decreases, they are regular
in size and just slightly smaller than the third anal spine.
All of the anal rays are segmented and branched from the
middle to the tip.
Caudal n. The caudal n is homocercal, deeply forked,
and has the upper and lower lobes showing the same shape
and size. There are 19 principal n rays, of which 17 are
branched, with nine in the upper and eight in the lower lobes
respectively (Figure 6). Ten and eight procurrent rays are
Figure 7. Previous hypotheses of interrelationships of aipichthyoids.
Characters: 1, Predorsal formula equals 0+0+0/x/; 2, lost of antorbital; 3,
presence of a plate-like anterior process in the hyomandibular; 4, pelvic
n in thoracic position; 5, distal ceratohyal without fenestra; 6, seven
pelvic n rays (reduction); 7, complex supraoccipital crest (formed with
the participation of the supraoccipital, frontal, and mesethmoid bones);
8, two epurals (reduction from three); 9, Five hypurals (reduction from
six); 10, Fusion of hypurals 1 and 2 with the pseudurostylar complex; 11,
six pelvic n rays (reduction from seven, see character 6); 12; rst pelvic
n ray unbranched.
Zoqueichthys carolinae gen. and sp. nov. 745
present in the anterior end of the upper and lower caudal
lobes respectively. A small caudal fenestra lies between the
rst principal and last procurrent rays of the dorsal caudal
lobe, similar to the fenestra that allows the insertion of the
upper bundle of interradialis muscle (Winterbottom, 1974)
or of the exa dorsalis (Otero and Gayet, 1996).
Specimen IHNFG-2224 has three long epurals over
the bifurcated spiny ends of the neural arch on the second
preural vertebra (this is the 25th of the entire vertebral se-
ries). The rst preural centrum and ural 1 are fused, forming
a pseudurostylar complex. This specimen has six autogenous
hypurals separated by a diastema between hypurals 2 and 3,
and two long uroneurals. The rst uroneural bears a wide
anterior membranous outgrowth or stegural that covers the
lateral dorsal surfaces the pseudurostylar complex. The par-
hypural and hypurals 1 and 2 are autogenous bones. There
is no evidence for the presence of urodermals (Figure 6).
Scales and scutes. Scales are cycloid and ovoid, slightly
higher than long, and cover the entire body, part of the
cheek and opercle. Seven or eight smooth ventral scutes of
regular size are aligned and occupy the abdominal ventral
border between the rst anal n spine and the pelvic n
base (Figure 5).
DISCUSSION
The unranked Aipichthyides, a monophyletic group
delineated by Otero and Gayet (1995) later recognized as
superfamily Aipichthyoidea, was distinguished based on
the following six presumed synapomorphies: 1) predorsal
formula equals 0+0+0/x/; 2) loss of antorbital; 3) presence of
a plate-like anterior process in the hyomandibular; 4) pelvic
n in thoracic position; 5) distal ceratohyal without fenes-
tra; and 6) seven pelvic n rays (Figure 7). Subsequently,
when Otero and Gayet (1996) formally named superfamily
Aipichthyoidea, they did so based only on characteristics
1 through 3; and in the later phylogenetic analysis of this
group, Otero (1997) resorted solely to synapomorphies 2
through 5. The inclusion of Zoqueichthys carolinae as a
new member of this group is strongly supported because it
shows characteristics 1 through 4 (Figures 2, 4-5).
Regarding the other two characteristics listed in the
original diagnosis, 5 and 6, do not agree with the present
material assigned to Zoqueichthys. However, there is scarce
documentation available on the distal ceratohyal among
Aipichthyoidea and this bone is obscured in Zoqueichthys
carolinae, so it is impossible to provide a useful assessment
of characteristic 5. On the other hand, data now available
shows that pelvic n rays are between 8 and 6 in these shes
(Table 2) and Z. carolinae is unique in having eight rays, a
number comparable to that found among some Beryciformes
(Fahay, 1997: 842).
Aipichthyoides and probably Aspesaipichthys
(Taverne, 2004) have a long and complex supraoccipital
crest involving the supraoccipital, frontal, and mesethmoid
bones, and extending the entire length of the skull roof
(character 7 in Figure 7). Zoqueichthys show a peculiar
long crest that is comparatively simpler in its composition,
involving only the supraoccipital and frontal bones (Figure
4). In contrast, all other aipichthyoid genera have a short,
simple crest, which rises above the postocular and part of
the ocular skull areas and involves only a dorsal laminar
projection from the supraoccipital bone. In the caudal skel-
eton of all Aipichthyoidea, including Zoqueichthys, there
are three epurals and six hypurals, except in Aipichthyoides
that has only two epurals and ve hypurals (characters
8 and 9 in Figure 7). Additionally, hypurals 1 and 2 are
autogenous in Zoqueichthys and other Aipichthyoidea,
except for Aipichthyoides and Aspesaipichthys, in which
these hypurals are fused to the pseudurostylar complex
(character10 in Figure 7).
Fins Vertebrae
Dorsal Anal Pectoral Pelvic Caudal
Aipichthyoides galeatus II+30/(31) III+23/(I+23) 10 76+I+9-8+I+6 11+15(4)
Aipichthyoides formosus II+25/(26) IV+18/(I+18) 10 78+I+9-8+I+7 11+15(4)
Aipichthys petriosus III+31/(?) III+19/(?) 12?-14? ? ? 12+14(?)
Aipichthys velifer III+22-24/(25-26) IV+15-17/(I+17-18) 11 67+I+9-8+I+6 11+16(4)
Aipichthys minor II+27-29/(29-30) IV-V+18-19/(I+18-19) 11 67+I+9-8+I+6 11+15(4)
Aipichthys oblongus II+26/? IV+16/(I+17) ? ? ? 10+16(4)
Aipichthys nuchalis ? ? ? 1+? ?10+6?
Aspesaipichthys cavaensis IV+29/(31) III-V+16-19/(I+19) ? ? 7+I+9-8+I+6 8+16(4)
Freigichthys elleipsis III+24/(26) IV+?/(I+16) ? 7? 7+I+8-?+?+? 11+16(4)
Paraipichthys lusitanicus IV+10/? V+7/? 10 67+I+9-8+I+6 10+?(4)
Zoqueichthys carolinae III+24/(26) III+17/(I+17) 12 8 10+I+9-8+I+8 11+16(4)
Table 2. Comparative sketch of components of paired and unpaired ns and vertebrae of all the species in superfamily Aipichthyoidea, including Zoqueichthys
carolinae gen. and sp. nov. Accounts of vertebrae involve abdominal (abd) and caudal (ca) vertebrae, as well as vertebrae with hemal and neural spines
supporting the caudal n skeleton (between parenthesis and as part of caudal vertebrae). Accounts for n elements include spines (in Roman numbers),
soft rays (in Arabic numbers), plus pterygiophores (enclosed into parenthesis). Anal perygiophores are divided in the rst pretygiophore modied in the
hemaxanal complex (I) and normal pterygiophores (Roman numbers).
Alvarado-Ortega and Than-Marchese
746
Interrelationships of superfamily Aipichthyoidea
Results of the phylogenetic analysis annexed to this manu-
script (see electronic complementary data) show that
Aipichthys, Freigichthys, Paraipichthys, Aipichthyoides,
Aspesaipichthys, and Zoqueichthys form the monophyletic
superfamily Aipichthyoidea (node 2, Figure 8), which is
supported by two synapomorphies: the plate-like anterior
process of the hyomandibular [15(1)] and the occurrence of
three predorsal bones in front of the rst neural spine [17(1)].
Additionally, this clade, Amblyopsidae, and Aphredoderidae
share a homoplasic characteristic, absence of the antorbital
bone [14(1)].
Results of the phylogenetic analysis show two mono-
phyletic groups outlined within superfamily Aipichthyoidea,
these are the families Aipichthyoidae and Aipichthyioididae.
The composition and supporting characteristics of these
groups depict some differences with previous hypotheses
(compare Figures 7 and 8). Remarkable in this new cla-
distic array are the inclusion of Paraipichthys in family
Aipichthyidae together with Freigichthys and Aipichthys, and
the position of Zoqueichthys within family Aipichthyoididae
together with Aipichthyoides and Aspesaipichthys.
Monophyly of Aipichthyoididae is supported by two
characters (node 3, Figure 8). Although in Aspesaipichthys
the supraoccipital crest is unknown, in this family the
crest is partially formed by the frontal [18(1)], and it is a
long laminar projection from the nasal area to the occiput
[21(1)]. Among aipichthyoidids, Aspesaipichthys and
Aipichthyoides share one synapomorphy, inclusion of the
mesethmoid bone in the supraoccipital crest [19(1)], but
the crest is unknown in the rst taxon. Additionally these
latter genera also share two homoplasic characteristics, they
have only ve hypurals [24(1)] as in Omosomopsidae, and
hypurals 1 and 2 are fused with the pseudurostylar complex
[25(1)] as in Amblyopsidae and Apheredoderidae. In this
analysis Zoqueichthys is unique among Aipichthyoidea in
having eight branchistegal rays [4(1)] and eight pelvic n
rays [10(0)].
The natural status of family Aipichthyidae is weakly
supported on the combination of two homoplasic charac-
teristics (node 5, Figure 8), the occurrence of six pelvic
fin rays [10(2)] as in Omosomopsidae, and the occur-
rence of one supramaxilla [28(1)] as in Omosomopsidae
and Sphenocephalidae. In this family the monophyletic
group Aipichthys + Freigichthys is supported by two syn-
apomorphies (node 6, Figures 8), the anterior edge of the
supraoccipital is thickened [22(1)] and the interfrontal is
at [27(1)] as previously noted by Otero and Gayet (1996).
These species also share one homoplasic condition, the half
boomerang shape of the supraoccipital crest [20(1)] also
found in crown acanthomorph clades (i.e. Amblyopsidae,
Percopsidae, among others). In this phylogenetic hy-
pothesis Aipichthys is presented as a paraphyletic group
with A. velifer apart from other species of Aipichthys. In
contrast, Freigichthys, A. minor, and A. oblongus consti-
tute a monophyletic group weakly supported on a single
homoplasic characteristic, opening of the frontal branch
of the sensory canal through pores [3(0)]. Finally, in this
group Freigichthys is unique in having seven pelvic n rays
[10(1)], which represents a homoplasy.
Although Taverne (2004) pointed out that the single
specimen thus far known of Aspesaipichthys does not
preserve the diagnostic features (or synapomorphies) of
Aipichthyoidea (Otero and Gayet, 1996), he supported its
inclusion in this superfamily because it shows a caudal
n with 19 principal rays, true epipleural bones, and three
predorsal bones located in front of the rst neural spine.
Nevertheless the rst two plesiomorphic features are also
present in other primitive acanthomorphs (i.e., Johnson and
Patterson, 1993; Patterson, 1993); whereas the predorsal
arrangement is doubtful according to the illustrations of
Aspesaipichthys published by Taverne (2004, gs. 3, 4),
in which these bones appear disarticulated and are located
behind the rst two neural spines. Tavern (2004) also justi-
ed inclusion of Aspesaipichthys in family Aipichthyoididae
because it shares with Aipichthyoides some common
characteristics, which include a reduced neural arch on
preural 2, the lack of fenestra in the caudal n, and rst
two hypurals fused to the pseudurostylar complex (ural 1 +
preural 1). These arguments appear weak because the rst
condition is a plesiomorphic one present in other primi-
tive acanthomorphs, and Aipichthyoides has a real caudal
n fenestra (Gayet 1980a, gs. 4 and 15). Fusion of the
rst two hypurals to the pseurodtylar complex is a feature
also present in derived acanthomorphs, i.e., Amblyopsidae
(Otero and Gayet, 1996).
Figure 8. Interrelationships of superfamily Aipichthyoidea suggested in
the present study. A complete description of this hypothesis is present in
the complementary electronic data.
Zoqueichthys carolinae gen. and sp. nov. 747
CONCLUSIONS
The phylogenetic hypothesis presented in this paper
attempts to depict the relationships of Zoqueichthys caroli-
nae within Aipichthyioidea. Recognition of the phylogenetic
position of this superfamily within Acanthomorpha requires
a comprehensive study, which in the future should include
all possible forms of basal recent and fossil acanthomorphs.
The mixture of primitive and derived characteristics of
Zoqueichthys allows its recognition as a new genus and
species in the family Aipichthyoididae, and validates this
Mexican sh as the rst Aipichthyoidea found in America.
Diagnostic characteristics of the superfamily
Aipichthyoidea include some of those described by other
authors (Otero and Gayet, 1995, 1996; Otero, 1997).
Both families Aipichthyidae and Aipichthyodidae are
shown here to be organized differently than in previous
hypotheses (compare Figures 7 and 8). Today the former
includes Paraipichthys, Aipichthys, and Freigichthys
whereas the latter includes Zoqueichthys, Aipichthyoides,
and Aspesaipichthys. In this new classication the family
Aipichthyidae is well characterized by a supraoccipital
crest that is half-boomerang shaped, relatively short, proj-
ects over the posterior half of the orbit and the postocular
skull areas, and has a thickened anterior edge. Otero and
Gayet (1995, 1996) discussed generic characteristics that
distinguish Aipichthys, and later Otero (1997) did the same
for Freigichthys. It is noticeable that the six pelvic n rays
shared by Aipichthys and Paraipichthys, and once recog-
nized as a synapomorphy, is here considered a homoplasic
condition.
On the other hand, family Aipichthyoididae is diag-
nosed by a high and extremely long supraoccipital crest.
The crest includes participation of the supraoccipital and
frontal bones, and extends over the whole skull roof, from
occiput to the nasal area. Zoqueichthys represents the sister
group of other genera in this family.
The discovery of Zoqueichthys complements our
understanding of the taxonomic diversity of basal acantho-
morphs. Efforts to describe the processes involved in their
evolution, origin, and early diversication must now include
an expanded biogeographical framework, and involve new
fossil evidence from the Cretaceous deposits of America.
ACKNOWLEDGEMENTS
We thank M.A. Coutiño and G.F. Carbot for the fa-
cilities to perform the present study. M. Gayet, O. Otero,
B. Khallou, and D. Mayrinck provided us with a valuable
bibliography. L.E. Gómez, F. Riquelme, and A. Alaniz
helped us during the eldwork. C. Ross kindly helped us
improve the English. A. López-Arbarello an O. Rauhut
gave us the facilities to review fossils of the Bayerische
Staatssammling für Paläontologie und Geologie. L.P.C.
Machado took the photographs shown in this manuscript.
We are indebted to S. Cevallos and his students, who col-
lected the fossils described in this manuscript. Our friend
L. Espinosa-Arrubarrena provided critical technical sup-
port for the development of this research. Valuable critical
suggestions by O.Otero, L.Taverne, and T. Lehman sig-
nicantly improved the content of this article. Universidad
Nacional Autónoma supports J.A.O. through the project
PAPIIT IN106011. B.A.T.M. acknowledges the support
of the project: Prospección y Resguardo del Patrimonio
Paleontologico de Chiapas.
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Manuscript received: April 17, 2012
Corrected manuscript received: July 22, 2012
Manuscript accepted: July 24, 2012.
