Journal of South American Earth Sciences 112 (2021) 103547

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Journal of South American Earth Sciences

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Fish trace fossils from the Lower Cretaceous of Puebla, Mexico

Rubén A. Rodríguez-de la Rosa a, *, María del Rosario Fernández-Barajas b,
Nelson A. Valdes-Vergara c, Erick Prado-Escamilla b
a
Unidad Académica de Ciencias Biológicas, Universidad Autónoma de Zacatecas, Calzada Solidaridad, esq. Paseo de la Bufa S/N, Campus II, C.P. 98060, Zacatecas,
Mexico
b
Laboratorio de Ciencias de la Tierra y Paleontología, Facultad de Estudios Superiores Iztacala, Universidad Nacional Autónoma de Mexico, C.P. 54090, Tlalnepantla de
Baz, Estado de Mexico, Mexico
c
Posgrado en Ciencias del Mar y Limnología, UNAM. Coyoacán, Ciudad Universitaria, Cto. Exterior S/N Coyoacán, 04510, CDMX, Mexico

A R T I C L E I N F O A B S T R A C T

Keywords: A great amount of studies dealing with vertebrate paleoichnology, are based on footprints made by terrestrial
San Juan Raya formation vertebrates; studies on fish paleoichnology, however, are less common, in spite that fish behaviour has the
Osculichnus tepitsin potential to leave different types of bioerosion and/or bioturbation structures. Recent fieldwork on southeastern
Daandavichnus batoideum
Puebla (Central Mexico), has yielded two Lower Cretaceous fish trace fossils, from the sedimentary sequence of
fish paleoichnology
the San Juan Raya Formation (upper Valanginian to lower Hauterivian). Osculichnus tepitsin isp. nov., is repre­
sented by small epichnial traces, lenticular in outline, with two elongate lip-like lobes, separated by a M-shaped
elongate furrow. The lower jaw lobe is thick, and nearly subrectangular; the upper jaw lobe bears a ventral
convex projection of the medial portion; two short, sub-triangular, maxillary barbel impressions are preserved.
Osculichnus tepitsin ichnosp. nov., is a benthic feeder fish trace fossil that represents, to date, the smallest ich­
nospecies of the ichnogenus Osculichnus; it is the first time that this ichnogenus is reported from Mexico, from the
American Continent, and from Lower Cretaceous rocks anywhere in the world. Daandavichnus batoideum ich­
nogen. and ichnosp. nov., is a trace fossil circumscribed within a small, sub-circular to trapezoid, outline; the
anterior area bears a deep impression with a M-shaped external boundary; the posterior area, preserves two
circular depressions, with smooth borders; Daandavichnus batoideum ichnogen. and ichnosp. nov., is interpreted
as the impression of the distal part of the jaw apparatus of a small-sized member of Batoidea, originated during
the symmetrical protrusion, while feeding in the benthos. Together with the ichnogenera Piscichnus and Oscu­
lichnus, Daandavichnus becomes the third ichnogenus that represents a fish feeding trace fossil, in this case a small
member of Batoidea. The close assoaciation of these new trace fossils, together with invertebrate traces such as
Skolithos and Helminthoidichnites; as well as xiphosuran feeding traces (Selenichnites), give insights into the well
stablished trophic relationships of one of the Lower Cretaceous ecosystems, represented within the sedimentary
sequence of the San Juan Raya Formation in southern Puebla, Mexico.

1. Introduction
Most studies dealing with vertebrate paleoichnology, are based on
footprints made by amphibians, reptiles, birds and mammals (e.g.,
Lockley and Hunt, 1995). Studies on fish paleoichnology, however, are
less common; in spite that fish behaviour, has the potential to leave
different types of bioerosion and/or bioturbation structures on a given
substrate (Muñiz et al., 2015).
Through feeding, hunting, walking and/or burrowing, modern fishes
produce several types of epi- or endogenic bioturbation structures in
both fresh and marine waters and at a variety of depths (Muñiz et al.,
2015). There is a great amount of papers dealing with modern fish
ichnology; these reports include the families Acipenseridae, Cen­
trarchidae, Cepolidae, Cichlidae, Gasterosteidae, Gobiidae, Mala­
canthidae, Oxudercinae, Petromyzontidae, Sparidae, Tetraodontidae,
Trachinidae, Protopteridae and members of Batoidea as well (Pearson
et al., 2007; Muñiz et al., 2015 and citations therein; Falkingham and
Horner, 2016).
However, the amazing diversity of extant fish traces is not reflected
in the fossil record (Muñiz et al., 2015). Some of the first reported

* Corresponding author.
E-mail address: rubenrodriguezr@uaz.edu.mx (R.A. Rodríguez-de la Rosa).

https://doi.org/10.1016/j.jsames.2021.103547
Received 14 December 2020; Received in revised form 29 August 2021; Accepted 30 August 2021
Available online 4 September 2021
0895-9811/© 2021 Elsevier Ltd. All rights reserved.
R.A. Rodríguez-de la Rosa et al.
Fig. 1. Geographic location of the tracksite, bearing the fish trace fossils (asterisk), south to Santa Ana Teloxtoc, in southeastern Puebla, central Mexico. 1 to 5, are
different localities bearing vertebrate tracks and traces.
ichnofossils related to a fish origin, were natural casts of burrows from
the Permian of Texas, these were attributed to aestivation activity of
Paleozoic lungfishes, and have been attributed to the genus Gnathorhiza
(Romer and Olson, 1954; Carlson, 1968; Dalquest, 1968; Olson and
Bolles, 1975; Berman, 1976; McAllister, 1988; Dalquest et al., 1989).
Among ichnofossils related to fish feeding, several coprolites, with a
conspicuous morphology, have been related to a fish origin (Hunt et al.,
2015); for example, complex spiral coprolites have been related to
hybodontid sharks, such as Hybodus (Coy, 1995), or Lonchidion and
Pristrisodus (Rakshit et al., 2018). Other amphipolar spiral coprolites
have been related to fishes such as Asipenceriformes and possibly sar­
copterygians (Rummy et al., 2021). Přikryl et al. (2012) reported gut
contents, or cololites, from the Lower Jurassic actinopterygian fish
Pachycormus; these cololites and pre-coprolites, contained numerous
hooklets attributed to the coleoid cephalopod Phragmoteuthis (Přikryl
et al., 2012).
An extraoidrinary find, is that of complex radial structures, that have
been interpreted as evidence of possible courtship behaviour in a
Devonian fish; these structures are, morphologically, similar to the
courtship structures constructed by male Japanese pufferfishes (Zong
and Gong, 2018).
The diversity of trace fossils attributed to fishes is low and only five
ichnogenera have been described to date: Undichna, Parundichna,
Broomichnium, Piscichnus and Osculichnus (Gregory et al., 1979; Feibel,
1987; Gregory, 1991; Simon et al., 2003; Minter and Braddy, 2006;
Benner et al., 2008; Demírcan and Uchman, 2010; Muñiz et al., 2015;
Szrek et al., 2016).
Undichna Anderson, 1976, it is the most commmon trace attributed
to swimming fishes; comprises fossils with a single horizontal wave, or
set of horizontal waves (paired and parallel, or unpaired) of common
wavelength and direction of travel (Minter and Braddy, 2006; Muñiz
et al., 2015).
Parundichna Simon et al., 2003, consists of swimming traces, in
which undulation of scratches is induced by an active gait of paired fins
with protruding fin rays; these traces have been attributed to coelacanth
fishes (Simon et al., 2003).
2
Journal of South American Earth Sciences 112 (2021) 103547
Broomichnium Kuhn, 1958, is a small, bilaterally symmetrical trace,
attributed to small demersal fishes, such as the members of Cottoidei
(Benner et al., 2008).
The two other ichnogenera, Piscichnus and Osculichnus, are attributed
to feeding activity of fishes. In the case of Piscichnus Feibel, 1987, its
emended diagnosis describes it, as a steep-sided, cylindrical or plug-like
to shallow, circular, dish-shaped structure of moderate to large size
oriented concave upward, more or less vertical to bedding (Gregory,
1991). Although originally interpreted as a nesting trace (Feibel, 1987);
it is actually considered as a feeding structure of rays jetting water
(Gregory et al., 1979; Gregory, 1991; Uchman et al., 2018).
The ichnogenus Osculichnus was described by Demírcan and Uchman
(2010) as hypichnial, bilobate mounds, generally elliptical or crescentic
in outline, having a smaller and a larger, lip-like lobe separated by an
undulate furrow (Demírcan and Uchman, 2010). Osculichnus is regarded
as the impression of the anterior part of a fish mouth (Demírcan and
Uchman, 2010; Szrek et al., 2016).
To date, four previous reports of the ichnogenus Osculichnus have
been made, these include the two known ichnospecies. Osculichnus
labialis, from the Eocene-Oligocene of Turkey (Demírcan and Uchman,
2010, 2016); Szrek et al. (2016), reported Osculichnus tarnowskae, from
the Lower Devonian of Poland (Szrek et al., 2016). The ichnogenus, was
also reported from Upper Devonian rocks near Wuhan, China (Fan et al.,
2019).
In recent times, the ichnogenus Osculichnus was found on a sandstone
block from the Lastres Formation, of Late Jurassic (Kimmeridgian) age,
in Spain (Piñuela-Suárez pers. com., 2020).
An ichnofossil, similar to Osculichnus, was reported by Pieńkowski
(1985), from Early Jurassic rocks in Poland; although these were not
refferred to a particular ichnogenera, the author recognized a fish origin
for these impressions (Pieńkowski, 1985).
Recently, Poropat et al. (2021) reported fish feeding traces from the
Upper Cretaceous Winton Formation of Queensland, Australia; these
were attributed to a lungfish and to an unknown actinopterygian
(Poropat et al., 2021).
Recent fieldwork conducted near Santa Ana Teloxtoc, Tehuacan
R.A. Rodríguez-de la Rosa et al. Journal of South American Earth Sciences 112 (2021) 103547

Fig. 2. Trace fossils of Osculichnus tepitsin ichnospecies nov.; A, STA-BAP-2, fine-grained sandstone slab bearing O. tepitsin traces, some indicated with black ar­
rowheads; B, CP.UAZ-015, slab preserving O. tepitsin traces (black arrowheads), together with xiphosuran traces attributed to Selenichnites (Sel); C, CP.UAZ-015, O.
tepitsin epichnial trace, note the typical lenticular outline; D, STA-BAP-3, epichnial trace with silicone cast, note the two lobes typical of the ichnogenus; E, silicone
cast of holotype STA-BAP-3, note the ventral convex projection of the medial portion of the upper jaw lobe (ujl, also seen in d and f), as well as one of the short, sub-
triangular, maxillary barbel impressions (arrow), lower jaw lobe (ljl); F, STA-BAP-3, silicone cast. Coin diameter is 23.0 mm; scale bars equal 50 mm in b, and 10 mm
in c-f.

Municipality, in southern Puebla, Mexico, has revealed Lower Creta­
ceous outcrops, of the San Juan Raya Formation, that preserve the tracks
and trackways of invertebrates, such as xiphosurans; and vertebrates,
such as turtles, crocodilians, pterosaurs and dinosaurs; fish trace fossils
are present as well.
In this paper, we report two fish feeding fossil traces. One of these
represents the third ichnospecies of the ichnogenus Osculichnus. It is the
first time that the ichnogenus Osculichnus is reported from the American
Continent and from Lower Cretaceous age rocks. The second feeding
trace is herein interpreted as a batoid feeding trace. To report these trace
fossils, as well as its paleoecological implications are the main purposes
of this paper.
2. Geological setting
The sediments of the San Juan Raya Formation, oucrop in the
Zapotitlán Basin, southern part of the state of Puebla, central Mexico
(Fig. 1). Its rich, and diverse, fossil fauna has been studied since the XIX
Century; Among the fossils found in this Lower Cretaceous sedimentary
sequence are corals, gastropods, cephalopods, pelecypods, brachiopods,
echinoderms and arthropods (Nyst and Galeotti, 1840; D’Orbigny, 1850;
Desor, 1850; Coquand, 1869; Müllerried, 1933; Alencáster de Cserna,
1956; Reyeros, 1963; Buitrón, 1970; González-Arreola, 1974; Buitrón
and Barceló-Duarte, 1980; Feldmann et al., 1995; Escalante-Ruiz, 2006;
Escalante-Ruiz and Quiroz-Barroso, 2006; Mora-Almazán and
Quiroz-Barroso, 2006; Löser, 2006; Mora-Almazán, 2008;

3
R.A. Rodríguez-de la Rosa et al. Journal of South American Earth Sciences 112 (2021) 103547

Fig. 3. Size range and morphological differences between Osculichnus tepitsin isp. nov. (A), Osculichnus labialis (B, Demícran and Uchman, 2010); Osculichnus cf.
labialis (C, Demícran and Uchman, 2016); Osculichnus tarnowskae (D, Szrek et al., 2016); Osculichnus from China (E, Fan et al., 2019); fish traces reported by
Pieńkowski (1985; F). In the case of A, C, D and E, the gray drawing depicts the maximum size, while the darker drawing represents the minimum size. In the case of
Osculichnus tepitsin isp. nov. (A), note the maxillary barbel impressions (arrows), the ventral convex projection of the medial portion of the upper jaw lobe (ujl) and
the lower jaw lobe (ljl).

Mendoza-Rosales, 2010; González-León et al., 2014, 2015).
The sedimentary sequence of the San Juan Raya Formation, it is
composed of an alternated sequence of sandstones, shales, limestones,
limolites and conglomerates. Sandstones are frequently found with
Ophiomorpha, Thalassinodides, Skolithos, Psilonichnus and Macanopsis
bioturbation structures; dinosaur tracks are knwon as well (Mendo­
za-Rosales, 2010; Rodríguez-de la Rosa et al., 2012; González-León
et al., 2014, 2015; Serrano-Brañas and Centeno-García, 2014).
The San Juan Raya Formation is divided into three members. The
lower member is known as Agua del Burro; it is composed of thin to
massive layers of limestones bearing rudists, pelecipods, corals and bi­
valves; these layers are interbedded with sandstone and shale
(Mendoza-Rosales, 2010). The middle part consists of marls, claystones,
argillaceous limestones, sandstones and limestones; while the upper part
is composed of a rhythmical alternace of marls, argillaceous limestones
and intercalated fossiliferous limestones lenses (Mendoza-Rosales,
2010).
According to Mendoza-Rosales (2010), the lower member is upper
Barremian in age; while the middle and upper parts of the San Juan Raya
Formation, are assigned to a lower Aptian age (Mendoza-Rosales, 2010).
However, it is probable, that part of the formation is older in age;
González-León et al. (2015) and Vega et al. (2019), suggest an age,
rangging from upper Valanginian to lower Hauterivian; this, based on
the presence of calcareous nannoplankton, in a sedimentary sequence

4
R.A. Rodríguez-de la Rosa et al.
with fossils of the lobster Atherfieldastacus magnus and the isopod
Natatolana sp.
Due to predominance of the fossils above mentioned, the San Juan
Raya Formation was thought to be marine in origin for several years
(Feldmann et al., 1995; Mendoza-Rosales, 2010; González-León et al.,
2014, 2015). However, the finding of dinosaur tracks near the towns of
San Martín Atexcal (Rodríguez-de la Rosa et al., 2012; Bravo-Cuevas and
Rodríguez-de la Rosa, 2014) and near San Juan Raya (Bravo-Cuevas and
Rodríguez-de la Rosa, 2014; Jiménez-Hidalgo et al., 2015), permited to
appreciate that, indeed, this sedimentary sequence preserves a great
diversity of paleoenvironmental conditions.
This sedimentary sequence bears a thickness of 800–900 m (Bar­
celó-Duarte, 1978); in spite of the presence of fossil vertebrate tracks
and traces in many outcrops, not the whole sequence represents shallow,
coastal environments; toward the medial and the uppermost portion, it
is possible to find storm deposits and reef limestone lenses, it is evidence
of the relative curve of the sea level (Mendoza-Rosales, 2010;
Serrano-Brañas and Centeno-García, 2014).
The ichnofossils, here described, were found near the small town of
Santa Ana Teloxtoc; at this area, the San Juan Raya Formation consists
of gray/bluish-gray and greenish-gray calcareous mudstone, interca­
lated with gray calcareous fossiliferous siltstone that ranges in thickness
from 10 to 40 cm, and gray calcareous fine to medium-grained sand­
stone, from 10 cm to 2 m in thickness. Sandstone forms both planar and
heterolithic stratification (Serrano-Brañas and Centeno-García, 2014).
The San Juan Raya Formation unconformably underlies the Cipiapa
Formation, at the Santana Teloxtoc area (Serrano-Brañas and Cen­
teno-García, 2014). The great abundance of fossils, as well as their di­
versity, has allowed the San Juan Raya Formation to be recognized
worldwide; however, this sedimentary sequence, still need a thorough
study regarding its tectonic and sedimentary history.
3. Material and method
The sandstone layer, bearing the traces, was entirely collected, due to
the fact that it represented a small outcrop, in risk, because its location
within the creek. Some silicone casts were made, to appreciate the
traces. Slabs, as well as the silicone casts, are housed in the Paleonto­
logical Collection of the Laboratorio de Ciencias de la Tierra y Paleon­
tología, Facultad de Estudios Superiores Iztacala, Universidad Nacional
Autónoma de Mexico, under the catalogue numbers STA-BAP-1 to STA-
BAP-5. Two additional slabs are housed in the Paleontological Collection
of the Unidad Académica de Ciencias Biológicas, Universidad Autónoma
de Zacatecas, under the catalogue numbers CP.UAZ-015, CP.UAZ-016.
Measurements were taken with the use of a digital vernier calliper.
4. Systematic ichnology
Ichnogenus Osculichnus Demírcan and Uchman, 2010.
Type ichnospecies: Osculichnus labialis Demírcan and Uchman
(2010).
Osculichnus tepitsin isp. nov. (Fig. 2).
Derivation of name: From the Nahuatl word, tepitsin, meanning
“tiny”, in reference to the small size of this fish feeding trace.
Diagnosis: Trace fossils lenticular in outline, wth two elongate lip-
like lobes, separated by a M-shaped elongate furrow; these differ from
the known Osculichnus ichnospecies in having an elongate, nearly sub­
rectangular, lower jaw lobe, with fusiform lateral ends; upper jaw lobe
with a ventral convex projection in its medial portion; presence of two
short, sub-triangular, maxillary barbel impressions.
Holotype: STA-BAP-3, slab and associate silicone cast, catalogued
under the same number (Fig. 2e).
Material: Five, medium-to fine-grained greyish sandstone slabs were
collected and numbered STA-BAP-1 to STA-BAP-5; CP.UAZ-015, CP.
UAZ-016.
Description. The trace fossils, here described, are preserved as
5
Journal of South American Earth Sciences 112 (2021) 103547
epichnial traces, in association with those of invertebrates, such as
Skolithos and Helminthoidichnites; as well as xiphosuran feeding traces,
similar to Selenichnites (Fig. 2b). This trace fossil bears a density, over the
substrate, of 8.2 traces per dm2 (5, minimum to 11, maximum).
These are lenticular to oval in outline; 0.6–11.3 mm in width, and
2.2–4.5 mm in height (Fig. 2c). Some traces, preserve a small elongate
ridge in their middle portion (Fig. 2d); in other specimens, this ridge is
poorly preserved or undistinguishable.
The ridge subdivides the trace into two areas, that are better
appreciated at the silicone casts; these areas, or lip-like lobes lobes, are
elongate and narrow, separated by an elongate furrow (Fig. 2d). In
vertical cross-section, most impressions are V-shaped to U-shaped; these
penetrate sediment 2.8–4.2 mm deep.
The elongate furrow, in the middle portion of the epichnial impres­
sions, is M-shaped, in horizontal outline; it is possible to appreciate that
both lobes are symmetric in shape; however, in some cases, one of the
lobes is slightly larger than the other (Fig. 2E, F). The upper jaw lobe,
bears a thicker medial portion and a ventral convex projection (Fig. 2E,
F). Two small sub-triangular structures are preserved laterally to the
medial portion of the upper jaw lobe, these are likely to represent short
maxillary barbel impressions (Fig. 2E). Lower jaw lobe is thick, with
fusiform lateral projections, that connect the lateral projections of the
upper jaw lobe.
The morphological details of these epichnial traces varies from well
preserved to specimens with absent furrow and lobes; in this way, the
impressions are seen as lenticular in outline, with a smooth inner
surface.
Remarks. The trace fossil, here reported, fits the general features of
the ichnogenus Osculichnus, in having a lenticular to oval in outline, with
a smaller and a larger, lip-like lobes, separated by an undulate furrow
(Demírcan and Uchman, 2010, 2016); thus, it is refferred to that ich­
nogenus; however, as mentioned below, shows features that support its
inclusion into a new ichnospecies (Fig. 3).
In 2010, Demírcan and Uchman, reported Osculichnus labialis from
Eocene-Oligocene sediments of the Mezardere Formation in the Thrace
Basin, near Malkara, Turkey. Its mean width and height, are 37 mm and
28.5 mm, respectively (Demírcan and Uchman, 2010).
Posteriorly, Demircan and Uchman (2016), reported Osculichnus
from the prodelta deposits of the Mezardere Formation, outcropping at
the Gökçeada Island, western Turkey (Demircan and Uchman, 2016).
This trace is 7–17 mm in width, and 7–16 mm in height. Although of a
smaller size, the reported traces did share the same morphological fea­
tures with Osculichnus labialis, and has been refferred to as Osculichnus cf.
labialis by the authors (Demircan and Uchman, 2016).
Szrek et al. (2016) reported Osculichnus tarnowskae, from the Lower
Devonian of Poland; its width ranges from 40 to 83 mm, while its height
from 39 to 77 mm; it is to date, the largest Osculichnus ichnospecies
(Szrek et al., 2016, Fig. 3).
Osculichnus was reported from Upper Devonian rocks of Wutong
Formation, near Wuhan, China (Fan et al., 2019). Its width ranges from
23.5 to 59.3 mm, while its height from 17.9 to 69.4 mm.
An ichnofossil, similar to Osculichnus, was reported by Pieńkowski
(1985: pl. 2G); from Early Jurassic rocks of the Holy Cross Mountains, in
Poland; although these were not refferred to a particular ichnogenera,
the author recognized a fish origin for these impressions (Pieńkowski,
1985); the size of these traces are ca. 10 mm in width and 0.75 mm in
height.
As mentioned above, the ichnogenus Osculichnus has been found in
rocks of the Late Jurassic (Kimmeridgian) Lastres Formation in Spain
(Piñuela-Suárez pers. com., 2020). The overall morphology and size of
this finding, recalls those of Osculichnus labialis, from Turkey (Demírcan
and Uchman, 2010).
The homgeneous size of the fossil traces, here described, in addition
to their density, suggest a single, small, tracemaker. As seen before,
other Osculichnus occurrences show a wide range of sizes, suggesting
small to large individuals (Demircan and Uchman, 2016; Fan et al.,
R.A. Rodríguez-de la Rosa et al.
Fig. 4. Daandavichnus batoideum ichnogen. and ichnosp. nov.; A, STA-BAP-2.4, holotype preserved as an epichnial trace, the anterior and posterior areas are shown
above and below the photograph, respectively; B, silicone cast of STA-BAP-2.4; C, STA-BAP-2.7; D-F, morphological variants of Daandavichnus batoideum ichnogen.
and ichnosp. nov. (D, STA-BAP-1.1; E, STA-BAP-2.6; F, STA-BAP-2.5). Note, in A-C, the anterior M-shaped external border, the two posterior circular depressions, as
well as the striations, forming an arch, preserved between these circular depressions. Scale bars equal 20 mm.
2019).
The ichnogenus Osculichnus shows a notable morphological varia­
tion, suggesting different fish trace-makers. Morphology ranges from the
typical lip-like traces of Osculichnus labialis (Demírcan and Uchman,
2010) to the rather “square” mouth of Osculichnus tarnowskae, attributed
to a lungfish (Szrek et al., 2016). Other Osculichnus traces show lip-like
lobes of rather equal size and shape, such as those reported by Demircan
and Uchman (2016) and Fan et al. (2019).
Among the features that supports Osculichnus tepitsin as a new ich­
nospecies, are the thick, nearly subrectangular, lower jaw lobe; the
ventral convex projection of the medial portion of the upper jaw lobe;
and the two short, sub-triangular, maxillary barbel impressions.
The lenticular outline, observed in Osculichnus tepitsin isp. nov., is
quite similar to that observed in the ichnogenus Lockeia, a bibalve
resting trace (Schlirf et al., 2001). However, a notable difference
regarding the diagnostic features of Lockeia, is the presence of a distinct
central median ridge, or crest, on its hypichnial mounds (Radley et al.,
1998; Schlirf et al., 2001; Paranjape et al., 2013). This crest, or ridge, is
unexistant in Osculichnus tepitsin isp. nov.; instead of this, a M-shaped
elongate furrow is present, corresponding to the space between dorsal
and ventral lobes of the fish mouth.
Osculichnus tepitsin isp. nov., is herein interpreted as a benthic feeder
fish trace fossil, that represents the first time that this ichnogenus is
reported from Mexico, from America and from Lower Cretaceous rocks
anywhere in the world.
Daandavichnus ichnogenus nov. Rodríguez-de la Rosa.
Diagnosis: As for the ichnospecies.
Derivation of name: From “da′ an davi”, the self-denomination of the
Mixteco language, the regional native language in the area; plus “ich­
nos”, latinised Greek “ikhnos”, meanning trace. Also, the generic name
honors Daniel and David, Rodríguez-Robles (sons of the first author).
Daandavichnus batoideum ichnospecies nov. Rodríguez-de la Rosa
(Fig. 4).
Derivation of name: In refference to the suborder Batoidea; and
“oideum”, from Latin -oides -oid + -eum, neuter of -eus -eous, “in
6
Journal of South American Earth Sciences 112 (2021) 103547
resemblance to”. Making refference to the general similarity of the mouth
structure of some members of Batoidea, during symmetrical protrusion
while feeding.
Diagnosis: Trace fossil, circumscribed within a sub-circular to trap­
ezoid outline; the anterior area preserves a curved impression with a M-
shaped external boundary; posterior area preserves two circular de­
pressions with smooth borders, occasionally connected by striations
forming an arch.
Holotype: STA-BAP-2.4; it is an epichnial impression on slab STA-
BAP-2; its silicone cast is catalogued under the same number (Fig. 4a
and b).
Material: STA-BAP-1.1 to 1.3, STA-BAP-2.4 to 2.8.
Description. This trace fossil is circumscribed within a faint, sub-
circular to trapezoid, outline, 30 mm in anteroposterior length and 25
mm in lateral width (Fig. 4a–f). Two components are observed, anterior
and posterior, of the same trace fossil. The anterior area preserves a deep
impression, with a M-shaped external boundary (Fig. 4a–d); another
variant, preserves two lateral impressions, lenticular in outline, and
connected by a faint U-shaped to V-shaped groove, resulting in the M-
shape external boundary, as mentioned above (Fig. 4c). This M-shaped
structure is 21.3 mm in mean width, and 11 mm in mean height. The
posterior area of this trace fossil, preserves two circular depressions,
with smooth borders; these depressions are 3.7 mm in mean diameter
(Fig. 4c). These are separated by a mean length of 8.1 mm. In some cases,
some striations are preserved, forming an arch and connecting these
circular depressions (Fig. 4a, c). Some variants of this trace are known
from the same rock layer (Fig. 4d–f). This trace fossil is herein inter­
preted, as the impression of the distal portion of the jaw apparatus of a
benthic feeder organism, during jaw protrusion.
Remarks. Feeding on benthic versus pelagic prey presents different
challenges for predators; in this way, some benthic feeders evolved
buccal specializations, such as jaw protrusion (Hernandez and Staab,
2015).
Jaw protrusion is an adaptation for feeding that evolved within
several actinopterygian lineages (Hernandez and Staab, 2015); also, it is
R.A. Rodríguez-de la Rosa et al.
an adaptation that evolved within members of Batoidea (Gregory, 1904;
Dean and Motta, 2004; Huber et al., 2019). Batoid elasmobranchs
exhibit the greatest degree of jaw kinesis among the cartilaginous fishes,
due to the possession of an euhyostylic jaw suspension; during prey
capture, can protrude the jaws symmetrically and/or asymmetrically
(Gregory, 1904; Dean and Motta, 2004; Huber et al., 2019).
The distal end of palatoquadrates and Meckel’s cartilage, together
with the labial cartilages of batoids, have a morphology capable of
leaving traces during the symmetrical protrusion of the jaw apparatus
(e.g., narcinoid batoids; Dean and Motta, 2004: Fig. 11; Huber et al.,
2019).
Batoids, such as myliobatids, leave feeding structures by jetting
water (Gregory, 1991; Gregory et al., 1979; Howard et al., 1977; Uch­
man et al., 2018). These trace fossils are known as Piscichnus Feibel,
1987; these are characterized by being dish-shaped structures, of mod­
erate (25–30 cm in diameter; Gregory et al., 1979) to large size (135 cm
in diameter; Feibel, 1987). The general features of Piscichnus, clearly
differ from those observed in Daandavichnus batoideum ichnogen. and
ichnosp. nov.
As described by Pearson et al. (2007), the traces left by modern
sturgeons comprise a crescent-shaped impression (snout) and a
plug-shaped excavation (mouth). In the case of Daandavichnus batoideum
ichnogen. and ichnosp. nov., the trace fossils are composed by the im­
pressions of two parts of the same structure, in this case the fish mouth.
Under this point of view, any comparison with the sturgeon traces is
ruled out.
As mentioned above, Daandavichnus batoideum ichnogen. and ich­
nosp. nov., is herein interpreted as the impression of the distal part of the
jaw apparatus of a benthic feeder organism; in this case a small-sized
member of Batoidea; it was originated during the symmetrical protru­
sion of the jaw apparatus, while feeding in the benthos; probably,
looking for preys, such as the Skolithos and Helminthoidichnites trace-
makers.
5. Discussion
As previously mentioned, the studies dealing with fish traces, are less
common; in spite that fish behaviour, have the potential to leave
different types of bioerosion and/or bioturbation structures on a given
substrate (Muñiz et al., 2015). The amazing diversity of extant fish
traces is not reflected in the fossil record, and only five ichnogenera
attributed to fish behavior have been described to date (Gregory et al.,
1979; Feibel, 1987; Gregory, 1991; Simon et al., 2003; Minter and
Braddy, 2006; Benner et al., 2008; Demírcan and Uchman, 2010; Muñiz
et al., 2015 and citations therein; Szrek et al., 2016).
Undichna Anderson, 1976, Parundichna Simon et al., 2003 and
Broomichnium Kuhn, 1958, are traces attributed to swimming fishes
(Simon et al., 2003; Minter and Braddy, 2006; Benner et al., 2008; Muñiz
et al., 2015). The two other ichnogenera, Piscichnus Feibel, 1987 and
Osculichnus Demírcan and Uchman, 2010, are traces attributed to the
feeding activity of fishes (Demírcan and Uchman, 2010; Szrek et al.,
2016).
In the case of Piscichnus Feibel, 1987; this trace has been interpreted
as a feeding structure of rays jetting water (Gregory et al., 1979; Greg­
ory, 1991; Uchman et al., 2018), while Osculichnus Demírcan and Uch­
man, 2010, is regarded as the impression of the anterior part of a fish
mouth, while feeding on invertebrates (Demírcan and Uchman, 2010;
Szrek et al., 2016).
The ichnogenus Osculichnus has been previously reported from the
Eocene-Oligocene of Turkey (Osculichnus labialis Demírcan and Uchman,
2010; Demircan and Uchman, 2016); the Lower Devonian of Poland
(Osculichnus tarnowskae Szrek et al., 2016) and Upper Devonian of China
(Fan et al., 2019); in recent times, Osculichnus was found in the Late
Jurassic of Spain (Piñuela-Suárez pers. com., 2020).
Pieńkowski (1985) reported an ichnofossil, similar to Osculichnus
from Early Jurassic of Poland; although these were not refferred to a
7
Journal of South American Earth Sciences 112 (2021) 103547
particular ichnogenera, the author recognized a fish origin for these
trace fossils (Pieńkowski, 1985).
The size of the ichnogenus Osculichnus ranges from small to large
(Fig. 3). To date, Osculichnus tarnowskae represents the largest ichno­
species (Szrek et al., 2016). Osculichnus tepitsin ichnosp. nov., fit the size
of the smallest specimens of Osculichnus cf. labialis reported by Demircan
and Uchman (2016; Fig. 3); however, the trace fossils, here described,
are morphologically different.
Among the features of Osculichnus tepitsin isp. nov., the presence of a
pair of short maxillary barbels is notable. Some Osculichnus labialis traces
also preserve barbel impressions; however, these differ in being larger
and rounded in shape (Demírcan and Uchman, 2010: figs. 3A and 4G).
Barbels are an essential sensory organ, for the food-seeking ability, of
some actinopterygian fishes (e.g., Siluriformes, Perciformes, Stomii­
formes and/or Cypriniformes); thus, it is possible to find them preserved
in association to these type of ichnofossils; its presence has been
demostrated, as well, by neoichnological experimental work (Demírcan
and Uchman, 2010).
An interesting observation is, that the number and size of barbels
varies among the different groups of fishes, and that barbels are strik­
ingly correlated with fish eye size. For example, large-eyed forms, live in
relatively clear water and have one pair of short barbels; while super­
numerary, long barbels, develop as an adaptation to silty streams in
compensation for reduced vision (Moore, 1950; Davis and Miller, 1967).
Poropat et al. (2021), reported fossil feeding fish traces from the Late
Cretaceous Winton Formation, in Queensland, Australia. An isolated
trace fossil, shows similarities with the feeding traces of extant stur­
geons; while, a second type has been related to a lungfish origin (Pearson
et al., 2007; Poropat et al., 2021).
Modern sturgeon feeding traces preserve curved impressions, that
represent the traces of barbels (Pearson et al., 2007: fig. 2). Although not
mentioned, some impressions, associated to the fish trace fossils from
Winton Formation, seem to represent barbel impressions (Poropat et al.,
2021: fig. 31C, and probably fig. 31B); it is interesting to note that, if
these represent barbels, it is in accordance with a reduced vision field
(Moore, 1950; Davis and Miller, 1967); in this case, caused by the bio­
turbation of turtles, crocodiles and dinosaurus (Poropat et al., 2021).
In the case of Osculichnus tepitsin isp. nov., the evidence of a potential
trace maker is scarce; to date, an isolated and partial skeleton of a small
fish, of an unknown affinity, has been recovered from rocks of the San
Juan Raya Formation; however, no detailed study has been done with
this evidence.
Osculichnus tepitsin ichnosp. nov., is a benthic feeder fish trace, that
represents to date the smallest ichnospecies of the ichnogenus Oscu­
lichnus; also, as previously mentioned, it is the first time that this ich­
nogenus is reported from Mexico, from the American Continent, and
from Lower Cretaceous rocks anywhere in the world.
Daandavichnus batoideum ichnogen. and ichnosp. nov., is herein
interpreted as the impression of the distal part of the mouth, of a small-
sized member of Batoidea; it was originated during the symmetrical
protrusion of the jaw apparatus, while feeding in the benthos; probably,
looking for preys, such as the Skolithos and Helminthoidichnites trace-
makers.
To date, osteological remains of batoids are unexistant in the sedi­
mentary sequence of the San Juan Raya Formation; however, relativelly
recent findings, indicate their presence; in this case, the trace of a large
batoid is known from a nearby site (Valdés-Vergara et al., 2016). It has
been interpreted as the trace left by a batoid, walking with the anterior
lobes of its pelvic fins (Valdés-Vergara et al., 2016). Also, some copro­
lites wtih a spiral morphology, frequently refferred to batoids, have been
found in a stratigraphic level close to that of the trace previously
mentioned.
The finding of the guitarfish Tlalocbatos applegatei (Brito et al., 2019)
in the Early Cretaceous (Upper Albian) sedimentary sequence of the
Tlayúa Formation of Tepexi de Rodríguez, Puebla, is noteworthy by two
different situations, explained below.
R.A. Rodríguez-de la Rosa et al.
Fig. 5. Paleoenvironmental reconstruction of the Osculichnus tepitsin-Daandavichnus batoideum association, at the San Juan Raya Formation in southern Puebla,
Mexico. The scene shows small fishes as the producers of Osculichnus tepitsin ichnosp. nov., a small batoid as the producer of Daandavichnus batoideum ichnogen. and
ichnosp. nov.; both preying on invertebrates, such as the Skolithos and Helminthoidichnites tracemakers. A xiphosurid, making Selenichnites traces, is also present in the
central portion of the image.
Although of a different age, theTlayúa Formation, outcrops close to
the geographic area, where the San Juan Raya Formation outcrops. It is
interesting to note, that Feldmann et al. (1998) described the marine
isopod Archaeoniscus from the sediments of the Upper Albian Tlayúa
Formation; this marine isopod, has been reported as well, from the
Valanginian-Hauterivian San Juan Raya Formation (Vega et al., 2019).
As mentioned by Vega et al. (2019), the presence of Archaeoniscus offers
an opportunity to study its presence in the same region, through time
(Vega et al., 2019).
Tlalocbatos represents a small individual, with a total length esti­
mated in 22.5 cm (Brito et al., 2019). Among small forms, such as Tla­
locbatos, could be the trace-maker of Daandavichnus batoideum ichnogen.
and ichnosp. nov.
Together with Piscichnus Feibel, 1987 and Osculichnus Demírcan and
Uchman, 2010, Daandavichnus becomes the third ichnogenus that rep­
resents a fish feeding trace, in this case a small member of Batoidea.
As mentioned above, the San Juan Raya sedimentary sequence pre­
serves a great diversity of paleoenvironmental conditions (Bar­
celó-Duarte, 1978). In regard to the environment represented in the
Osculichnus-Daandavichnus layer; it seems to represent a shallow marine
facies (Fig. 5). However, the paleoevironments represented, among the
different Osculichnus reports, range from marginal-marine, brackish
marine, estuarine embayment, fluvio-deltaic and prodelta facies
(Valenzuela et al., 1998; Demírcan and Uchman, 2010, 2016; Szrek
et al., 2016; Fan et al., 2019; Piñuela-Suárez pers. com., 2020).
The close assoaciation of Osculichnus tepitsin ichnosp. nov., Daanda­
vichnus batoideum ichnogen. and ichnosp. nov.; invertebrate traces such
as Skolithos and Helminthoidichnites; as well as the xiphosuran feeding
trace Selenichnites, give insights into the well stablished trophic re­
lationships of one of the Lower Cretaceous ecosystems, represented
within the sedimentary sequence of the San Juan Raya Formation in
southern Puebla, Mexico.
Author contributions
Rubén A. Rodríguez-de la Rosa: Investigation, Conceptualization,
Methodology, Writing. María del Rosario Fernández-Barajas: Data
curation, Reviewing and Editing. Nelson A. Valdes-Vergara: Writing –
original draft preparation, Reviewing and Editing. Erick Prado-
Escamilla: Fieldwork support.
8
Journal of South American Earth Sciences 112 (2021) 103547
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgements
Authors thank all persons from the Museo Histórico, Cultural y
Paleontológico (HICUPA), at Santa Ana Teloxtoc, Puebla, for allowing us
to work on their community. Authors want to thank to Alfred Uchman
(Institute of Geological Sciences, Jagiellonian University) and Ricardo
N. Melchor (Universidad Nacional de La Pampa, Argentina), for their
kind review and great improvement to the original manuscript. RARR
thanks to the proyect UAZ-2019-37852.
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