Journal of South American Earth Sciences 88 (2018) 132–143

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Journal of South American Earth Sciences

journal homepage: www.elsevier.com/locate/jsames

Quasimodichthys gen. nov. (Neopterygii: Semionotiformes): A morphological T

and ontogenetic study
Hanna Carolina Lins de Paiva∗, Valéria Gallo∗∗
Laboratório de Sistemática e Biogeografia da UERJ, Universidade do Estado do Rio de Janeiro, Rua São Francisco Xavier, 524. Maracanã, 20550-013, Rio de Janeiro, RJ,
Brazil

A R T I C LE I N FO A B S T R A C T

Keywords: Many Lepidotes species were recorded worldwide during most parts of the twentieth century, even in the
Systematics Brazilian sedimentary basins. Among the Brazilian sedimentary basins, the Late Jurassic “L.” piauhyensis is one
Ontogeny the best described, especially because the relatively great amount of well-preserved and articulated specimens.
Morphology As suggested in the most recent phylogenetic analysis, Lepidotes should be restrict to the Lower Jurassic of
Ginlgymodi
Central Europe and all the Lepidotes species recorded beyond this interval belong to other genera. Herein “L.”
Mesozoic
piauhyensis was revised and presents new morphological and ontogenetic information, which allow us to propose
a new genus, Quasimodichthys. This study reveals additional characters that Quasimodichthys piauhyensis shares
with Hoyasotes tanyrhis, Lepidohyas microrhis and Neosemionotus puntanus. Additionally, four distinct ontogenetic
stages were observed in Quasimodichthys piauhyensis, juvenile (1 and 2), sub-adult, and adult.

1. Introduction
Many Lepidotes species were worldwide recorded during most parts of
the twentieth century, even in the Brazilian sedimentary basins (Gallo and
Brito, 2004; Gallo, 2005; Paiva et al., 2013, 2014; Cavin et al., 2013).
Among the Brazilian “Lepidotes”, “L.” piauhyensis (Roxo and Löfgren, 1936)
is the best described one, especially because of the relatively great amount
of well-preserved and articulated specimens (Paiva, 2017).
“Lepidotes” piauhyensis is recorded in the Upper Jurassic of the
Parnaíba Basin, Northeastern of Brazil. Despite the brief description
furnished by Roxo and Löfgren (1936), Silva Santos (1945) and
Schaeffer (1947) made extensive morphological studies and stated that
“L.” piauhyensis is a Jurassic taxon, related to “L.” congolensis Hussakof,
1917 (Upper Jurassic of the Congo Basin, Africa), due to the strong
similarity in scale morphology.
This relationship was also noted by Gallo-da-Silva (1998) and Gallo
(2005) that redescribed “L.” piauhyensis and established comparisons
with older studies, beyond a discussion about the relationships among
the others Brazilian and African Lepidotes species. Additionally, two
autapomorphies were established to “L.” piauhyensis: presence of bony
lamina on the ventral portion of anterior ceratohyal, and dorsal and
ventral hypohyals.
Although “L.” piauhyensis had been considered a well-known
Brazilian “Lepidotes” representative, López-Arbarello (2012) argued
that Lepidotes genus should be restrict to the Lower Jurassic of Central
Europe. Additionally, López-Arbarello (2012) noted that all the Lepi-
dotes species recorded beyond this interval belong to other genera or
need to be revised. Therefore, herein we revise “L.” piauhyensis, adding
new morphological and ontogenetic data. These new information
complement the Gallo (2005) analysis and provide a new comprehen-
sion about Brazilian paleoichthyofauna and change the “L.” piauhyensis
nomenclature, named here with a new generic epithet, Quasimodichtys.
2. Material and methods
The studied holotype (DGM-297-P) and paratype (DGM-295-P) are
housed at the Museu de Ciências da Terra – Serviço Geológico do Brasil
(CPRM), Rio de Janeiro, Brazil. These are almost complete specimens,
with details of skull and fins. Additional material, analyzed by Gallo-da-
Silva (1998) and Gallo (2005), is housed at American Museum of
Natural History and paleozoological collection of the da Universidade
do Estado do Rio de Janeiro. Except for Callipurbeckia tendaguruensis,
Camerichthys lunae and Pliodetes nigeriensis, all comparative material
was directly examined by the authors.
2.1. Institutional abbreviations
AMNH: American Museum of Natural History, New York, United

∗
Corresponding author.
∗∗
Corresponding author.
E-mail addresses: hanna.clp@gmail.com (H.C.L.d. Paiva), gallo@uerj.br (V. Gallo).

https://doi.org/10.1016/j.jsames.2018.08.010
Received 4 May 2018; Received in revised form 13 August 2018; Accepted 13 August 2018
Available online 15 August 2018
0895-9811/ © 2018 Elsevier Ltd. All rights reserved.
H.C.L.d. Paiva, V. Gallo
States; BMNH/NHMUK PV: Natural History Museum (Paleontology
vertebrates), London, UK; DGM/MCT: Museu de Ciências da Terra –
Serviço Geológico do Brasil, Rio de Janeiro, Brazil; FPH: Fundação
Paleontológica Phoenix, Aracaju, Brazil; MGM: Museo GeoMinero,
Instituto Geológico y Minero de España, Madrid, Spain; MN: Museu
Nacional, Rio de Janeiro, Brazil; MNHN: Muséum national d’Histoire
naturelle, Paris, France; Pz.UERJ: Coleção Paleozoológica da
Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil.
2.2. Comparative material
Araripelepidotes temnurus (Agassiz, 1841): DGM 1104-P (holo-
type), AMNH 11809, AMNH 11810, AMNH 11811, AMNH 11812,
AMNH 11813, AMNH 11814, AMNH 11815, AMNH 11831, AMNH
11832, AMNH 11833, AMNH 11834, AMNH 11835, AMNH 12645,
AMNH 12716, AMNH 13116, Pz.UERJ 039, Pz.UERJ 040, Pz.UERJ 041,
Pz.UERJ 042, Pz.UERJ 043, Pz.UERJ 047, Pz.UERJ 102; Callipurbeckia
minor (Agassiz, 1833): BGS. GSM 27975 (lectotype), NHMUK PV P
1118, NHMUK PV P 2006, NHMUK PV P 4989, NHMUK PV P 5591,
NHMUK PV P 29393, NHMUK PV P 36080, NHMUK PV P 36081,
NHMUK PV P 41157; Callipurbeckia tendaguruensis (Arratia and
Schultze, 1999): MB. f7040a-b (holotype), MB. f7041a-b, MB. f7042,
MB. f7043, MB. f7044, MB. f7045, MB. f7046, MB. f7047; Camer-
ichthys lunae Bérmudez-Rochas and Poyato-Ariza, 2015: MGM-108C
(holotype); “Lepidotes” alagoensis Gallo, 2000: FPH 0120-V (holo-
type), Pz.UERJ 446, Pz. UERJ 466, Pz.UERJ 467, Pz.UERJ 468; “Le-
pidotes” dixseptiensis Silva Santos, 1963: DGM 675-P (holotype), DGM
425-P, DGM 1130-P, MN 4377-V, MN 4378/1-V, MN 4378/2-V, MN
4378/3-V; Lepidotes elvensis (Blainville, 1818): MNHN JRE-545 (ho-
lotype), AMNH 7537, BMNH P 32421, MNHN 10527, MNHN 10528;
“Lepidotes” mawsoni Woodward, 1888: BMNH P 5534 a, b (holotype),
BMNH P 410, BMNH P 412, BMNH P 2280, BMNH P 7114, BMNH P
7712, BMNH P 7713, BMNH P 7718, BMNH P 7340, BMNH P 7341,
BMNH P 7342, BMNH P 7343, BMNH P 7344, DGM 426-P, MN 4361/1-
V, MN 4361/2-V, MN 4361/3-V, MN 4361/4-V, MN 4362-V, MN 4363/
1-V, MN 4363/2, MN 4363/3-V, MN 4363/4-V, MN 4364-V, MN 4365-
V, MN 4366-V, MN 4367-V, MN 4368-V, MN 4369/1-V, MN 4369/2-V,
MN 4369/3-V, MN 4369/4-V, MN 4370/1-V, MN 4370/2-V, MN 4370/
3-V, MN 4371/1-V, MN 4371/2-V, MN 4371/3-V, MN 4371/4-V, MN
4371/5-V, MN 4372-V, MN 4373-V, MN 4374-V, MN 4375/1-V, MN
4375/2-V, MN 4375/3-V, MN 4375/4-V, MN 4376-V, MN 4377-V, MN
4379/1-V, MN 4379/2-V, MN 4379/3-V, MN 4380/1-V, MN 4380/2-V,
MN 4380/3-V, MN 4380/4-V, MN 4381/1-V, MN 4381/2, MN 4381/3-
V, MN 4381/4-V, MN 4381/5, MN 4381/6-V, MN 4381/7-V, MN 4381/
8-V, MN 4381/9-V, MN 4381/10-V, MN 4386/1-V, MN 4386/2-V, MN
4387-V, MN 4388-V; “Lepidotes”oliveirai Silva Santos, 1969: DGM
952-P (sintypes), MCT 1450; “Lepidotes” roxoi Silva Santos, 1953:
DGM 423-P (holotype), DGM 422-P, DGM 429-P, DGM 430-P, DGM
431-P, MN 4383-V, MN 4384-V, MN 4385-V, MN 4386/1-V, MN 4386/
2-V, MN 4387-V, MN 4388-V; Lepidotes semiserratus Agassiz, 1833:
NHMUK PV P 3528; “Lepidotes” souzai Woodward, 1908: BMNH P
10603 (holotype), BMNH P 10566, BMNH P 10601, DGM 424-P, MN
4382/1-V, MN 4382/2-V, MN 4391/1-V, MN 4391/2-V; “Lepidotes”
wenzae Brito and Gallo, 2003: MNHN-BCE 387 (holotype), MN 4791-V
(paratype); Paralepidotus ornatus Agassiz (1833): AMNH 1029; Plio-
detes nigeriensis Wenz, 1999: MNHN.F.GDF 127 (holotype); Scheenstia
laevis (Agassiz, 1833): MNHN.F.JRE 370; Scheenstia mantelli (Agassiz,
1833): NHMUK PV P 6943, NHMUK PV P 6933; Scheenstia maximus
(Wagner, 1863): AMNH 13097, MNHN 1885-19, MNHN 1878-116B,
MNHN 1878-7, MNHN 1868-15, MNHN.F.JRE 369, MNHN.F.JRE 375.
2.3. Anatomical nomenclature
The anatomic description present in this work was compared with
Wenz (1967, 1999), Thies (1989), Gallo-da-Silva (1998), Gallo (2005),
López-Arbarello (2012), Bérmudez-Rochas and Poyato-Ariza (2015)
133
Journal of South American Earth Sciences 88 (2018) 132–143
and López-Arbarello and Wencker (2016).
Ontogenetic study followed Cloutier (2010), beyond a comparison
with Paralepidotus ornatus ontogenetic sequence (Tintori, 1996; Gallo
et al., 2002).
2.4. Anatomical abbreviations
ACh, anterior ceratohyal; Ang, angular; Ao, antorbital; Apal, au-
topalatine; Art, articular; asc.proc.Psph, ascending process of the
parasphenoid; Br, branchiostegal ray; Cl, cleithrum; Dpal, dermopa-
latine; Dpt, dermopterotic; Dsph, dermosphenotic; D, dentary; Ecpt,
ectopterygoid; Enpt, endopterygoid; Exc, extrascapula; fr, fin ray; Fr,
frontal; fr.f., fringing fulcra; Hm, hyomandibula; Hy, hypohyal; Io,
infraorbital; io. s.c., infraorbital sensory canal; Iop, interopercle;
md.s.c., mandibular sensory canal; Mpt, metapterygoid; Na, nasal; Op,
opercle; Pa, parietal; Part, prearticular; Pcl, postcleithrum; PCh, pos-
terior ceratohyal; Pmx, premaxilla; Pop, preopercle; pop.s.c., pre-
opercular sensory canal; Ps, parasphenoid; Pt, posttemporal; Qu,
quadrate; Sang, surangular; Scl, supracleithrum; So, supraorbital;
so.s.c., supraorbital sensory canal; Sob, suborbital; Sop, subopercle;
Vo, vomer.
We also analyzed the phylogenetic relationships of the Brazilian
“Lepidotes” among Ginglymodi. The matrix was built on Mesquite 3.10
(Maddison and Maddison, 2016) and analyzed on Tree Analysis using
New Technology (TNT) 1.5 (Goloboff and Catalano, 2016). The data
were submitted to parsimony algorithm Traditional Search, with ran-
domic adition of 2000 replicates, 10 obtained trees on each step and
TBR (Tree Bisection Reconnection).
The characters used on this study were the same of López-Arbarello
(2012), updated on López-Arbarello and Wencker (2016). The char-
acters list and the matrix are on the appendices (Appendix 1). The inner
group was formed by 33 genera: (Adrianaichthys Meunier et al., 2016;
Araripelepidotes Silva Santos, 1990; Atractosteus Rafinesque, 1820; Cal-
lipurbeckia López-Arbarello, 2012; Camerichtys Bérmudez-Rochas and
Poyato-Ariza, 2015; Dentilepisosteus Grande, 2010; Herreraichthys
Alvarado-Ortega et al., 2016; Hoyasotes Poyato-Ariza and Martın-Abad,
2016; Isanichthys Cavin and Suteethorn, 2006; Kyphosichthys Xu and
Wu, 2012; Lepidotes Agassiz, 1832; Lepidohyas Poyato-Ariza and Martın-
Abad, 2016; Lepisosteus Lacépède, 1803; Lophionotus Gibson, 2013;
Luoxionichthys Wen et al., 2012; Macrosemiocotzus González-Rodíguez
et al., 2004; Macrosemius Agassiz, 1833; Macrosemimimus Schröder
et al., 2012; Masillosteus Micklich and Klappert, 2001; Neosemionotus
Bocchino, 1973; Notagogus Agassiz, 1833; Obaichthys Wenz and Brito,
1992; Occitanichthys López-Arbarello and Wencker, 2016; Oniichthys
Cavin and Brito, 2001; Paralepidotus Stolley, 1920; Pliodetes Wenz,
1999; Propterus Agassiz, 1833; Sangiorgioichthys Tintori and Lombardo,
2007; Scheenstia López-Arbarello and Sferco, 2011; Semiolepis
Lombardo and Tintori, 2008; Semionotus Agassiz, 1832; Thaiichthys
Cavin et al., 2013; Tlayuamichin López-Arbarello and Alvarado-Ortega,
2011). The chosen outgroup was Perleidus sinensis. “L.” dixseptiensis, “L.”
mawsoni e “L.” oliveirai were excluded of phylogenetic analyzes, be-
cause they presented more than 50% of missing data on the firstly
catalogued bibliographic data, according Wilkinson and Benton (1996)
methodology.
3. Geological setting
All the analyzed specimens of this study were collected in Fazenda
Muzinho, Pastos Bons Formation (Piauí State, Brazil). The strata of
Pastos Bons Formation (Fig. 1) are composed of green sandstones and
grey to green siltstones (Vaz et al., 2007).
These sediments are fluvial-lacustrine, deposited in an arid and
seasonal paleoenvironment during the Late Jurassic (Oxfordian) (Vaz
et al., 2007; Petra and Gallo, 2012). Additionally, Petra (2006) and
Petra and Gallo (2012) argued that the Fazenda Muzinho could be re-
lated to a saline lake, in which the water temperature could vary from
H.C.L.d. Paiva, V. Gallo
Fig. 1. Location map of Pastos Bons Formation, Floriano city, State of Piauí, Brazil.
20 °C to 30 °C, interleaved with dry periods, characterizing a cata-
strophic assemblage.
According to Brito and Gallo (2002) and Petra (2006), Fazenda
Muzinho locality contains ostracods, conchostraca, palynomorphs,
plant cuticles and fishes, as well as Mawsoniidae, Macrosemiidae,
Semionotidae and Pleuropholidae, some of them in distinct ontogenetic
stages.
4. Systematic paleontology
NEOPTERYGII Regan, 1923.
GINGLYMODI Cope, 1872 (sensu López-Arbarello, 2012)
SEMIONOTIFORMES Woodward, 1890 (sensu Gallo, 2005)
Quasimodichthys gen. nov.
Type species. Quasimodichthys piauhyensis (Roxo and Löfgren,
1936), by original description (Fig. 2).
v. 1936 Lepidotus piauhyensis Roxo and Löfgren, p. 7, figs. 1, 2.
v. 1945 Lepidotus piauhyensis Silva Santos, p. 1, pls. 1-4.
v. 1947 Lepidotes piauhyensis Schaeffer, p. 12, pl. 1.
v. 2005 Lepidotes piauhyensis Gallo, p. 1, figs. 1-8.
v. 2017 “Lepidotes” piauhyensis Paiva, p. 42-64, figs. 4-13.
Etymology. ‘Quasimodo’ in honor the ‘Hunchback of Notre Dame’
personage, because the hump-backed body format, and ‘ichthys’ fish in
Greek.
Holotype. DGM-297-P (Fig. 2), comprising almost complete spe-
cimen. Paratype. DGM-295-P.
Referred material. AMNH FF 10013; DGM 291-P; DGM 292-P;
DGM 293-P; DGM 294-P; DGM-296-P; DGM 298-P; Pz.UERJ 408;
Pz.UERJ 409; Pz.UERJ 410; Pz.UERJ 411; Pz.UERJ 412; Pz.UERJ 413;
Pz.UERJ 414; Pz.UERJ 415; Pz.UERJ416; Pz.UERJ 513; Pz.UERJ 514;
Pz.UERJ 515.
Type Locality. Fazenda Muzinho, 16 Km northestern from Floriano
city, Floriano-Amarante Road, State of Piauí, Brazil.
Stratigraphical distribution. Pastos Bons Formation, Late Jurassic
134
Journal of South American Earth Sciences 88 (2018) 132–143
Fig. 2. Quasimodichthys piauhyensis, holotype DGM-297-P, scale bar equals
5 cm.
(Oxfordian), Parnaíba Basin (Vaz et al., 2007).
Emended diagnosis. Semionotiform fish with 480 mm of max-
imum standard length (SL); fusiform juveniles, hump-backed sub-adults
and adults; triangular head occupying about ¼ total body length; par-
ietals with ⅓ of frontal length; paired asymmetric extrascapulae;
complete and closed circumorbital ring; most anterior infraorbital
deeper than others; most ventral infraorbital bone longer than the
others; a single row of suborbitals; a crescent-shaped preopercle; rec-
tangular opercle; smooth dermal bones in juvenile 1, however the or-
namentation increases during growth; co-ossified vomers; two series of
pores on dentalosplenial sensory canal; quadratojugal present; deep
endopterygoid; presence of dorsal and ventral hypohyals (*); presence
of bony lamina on ventral portion of anterior ceratohyal (*); dorsal fin
posteriorly located from pre-dorsal elevation, between pelvic and anal
fins; presence of fulcra on all fins; 33–40 diagonal rows of scales from
lateral line; ganoid scales with peg-and-socket articulation and dorsal
and ventral anterior processes; smooth posterior border in the juvenile
1 scales; ornamentation on posterior border of the juvenile 2 anterior
scales, sub-adults and adults with almost all scales ornamented on
posterior border; rectangular shape in anterior scales; quadrate in
medial scales and rhomboid in caudal scales.
(*) autapomorphies.
H.C.L.d. Paiva, V. Gallo
Fig. 3. Quasimodichthys piauhyensis, holotype DGM-297-P. A. Draw of head; B.
Detail of head. Scale bar equals 3 cm.
Description. Quasimodichthys piauhyensis is a medium-sized fish,
known by specimens ranging between 75 and 480 mm of SL. The
smaller individuals have a fusiform body while the larger ones are
hump-backed (sensu Tintori, 1996). The head is triangular and occupies
¼ from total length of body. The skull is ornamented by dense tu-
bercles, with varying pattern among specimens. The dorsal fin is im-
mediately posterior to the predorsal elevation, between the pelvic and
anal fins. All fins contain fringing fulcra. The ganoid scales possess a
reduced peg-and-socket articulation and well-marked anterior pro-
cesses. The posterior border of the scales is ornamented with denticu-
late projections. Additionally, the shape of the scales varies according
to position: the anterior are rectangular, the medial are quadrate and
the caudal are rhomboid.
Snout. The antorbitals (Fig. 3) are reniform, ventrally placed to the
nasals, and bear the pores of the ethmoid comissure. The premaxilla
(Fig. 4) possesses an elongated nasal process, narrowing in the edge and
penetrates the nasal. In Pz.UERJ 515 six narrow and styliform teeth
were preserved, all of them observed only in the right side. The nasals
are also reniform, narrower than the antorbitals and separated by the
premaxilla nasal process.
Skull Roof. The frontals are three times longer than wide, narrower
in the orbit region. The supraorbital sensory canal runs medially on the
frontals and the parietals.
135
Journal of South American Earth Sciences 88 (2018) 132–143
Fig. 4. Quasimodichthys piauhyensis, specimen Pz.UERJ 515. A. Draw of head; B.
Detail of head. Scale bar equals 3 cm.
The parietals are approximately rectangular (Figs. 3–5), with a
sinuous suture. There is a deep groove that extends to the dermop-
terotic and forms the middle pit line.
The dermopterotics are also approximately rectangular, articulated
lateroposteriorly with the frontal. The number of extrascapulae vary
among the specimens; while the holotype, DGM-297-P (Fig. 3), seems to
possesses three representatives, there is a paired and symmetric one in
Pz.UERJ 514 (Fig. 5).
Neurocranium. The parasphenoid is elongated and thin, forming a
relatively wide lamina in the palate, gradually narrowing backwards,
and forking in two relatively elongated and narrow lateral projections.
The posterior margin is also forked, in two short and triangular por-
tions. They are triangular in format and seem similar to the posterior
part of an arrow. Due the preservation mode in samples, it was not
possible to observe foramens, or additional structures.
The unique vomer preserved in Q. piauhyensis was observed in
Pz.UERJ 513 (Fig. 6). It is a co-ossified bone, anterior to the para-
sphenoid and possesses three preserved viliform teeth.
Circumorbital bones. The circumorbital bones form a relatively
small and closed orbit, approximately five times smaller than the head.
This structure is best preserved only in DGM-297-P, in the left side
(Fig. 7).
There are seven infraorbitals in the circumorbital ring. They are
approximately rectangular and the pores of the infraorbital sensory
H.C.L.d. Paiva, V. Gallo Journal of South American Earth Sciences 88 (2018) 132–143

Fig. 5. Quasimodichthys piauhyensis, specimen Pz.UERJ 514. A. Draw of head; B.

Detail of head. Scale bar equals 3 cm.

Fig. 7. Quasimodichthys piauhyensis, holotype DGM-297-P, in left side. A. Draw

of head; B. Detail of head. Scale bar equals 4 cm. Adapted from Gallo (2005).

canal are present in their middle portion. Additionally, there is an

elongated infraorbital bone that occupies the most ventral portion in
the circumorbital ring. One anterior infraorbital was also preserved, on
the left side of DGM-297-P, rectangular in format and narrower than the
others.
A rounded to rectangular dermosphenotic is located between the
last infraorbital and the most posterior supraorbital. In some specimens
(i.e., DGM-295-P, DGM-297-P and Pz.UERJ 414) the dermosphenotic
possesses a uniform ornamentation, but it is smooth in Pz.UERJ
513–515. Three to four big and ornamented supraorbitals are present.
These are rectangular and the first supraorbital reaches the infraorbital
series, closing the orbit (Fig. 7).
The suborbitals are organized in a single row, between the pre-
opercle and the infraorbital series. They are approximately rectangular
and occupy the ventral margin of the infraorbital series.
Opercular series. The preopercle is an elongated and narrow bone,
crescent-shaped and preserved some pores of the preopercular sensory
Fig. 6. Quasimodichthys piauhyensis, specimen Pz.UERJ 513, scale bar equals
canal in its horizontal arm.
1 cm.
The opercle is the biggest bone of the opercular series. It is

136
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rectangular in almost all the specimens, however in Pz. UERJ 514 it

possesses a rounded dorsal margin. In addition, the opercle is orna-
mented in the anterior region. The subopercle is shorter and wider than
the opercle, with a wide ascendant process that reaches ¼ of the opercle
height. The interopercle is relatively big, triangular and smooth; it is
located near to the preopercle ventral portion.
Upper Jaw. The maxilla is a short bone anterior to the orbit. The
posterior portion is highest, tapering forwards and reaching the pre-
maxilla, although the presence of an anteromedial articular process was
not observed. The posterior border of the maxilla is almost straight and
contacts the posterolateral region of the dentalosplenial. The supra-
maxilla was observed only on the left side of the upper jaw in DGM-
297-P (Fig. 7). It is unique, narrow and short, placed dorsoposteriorly to
the maxilla.
Lower Jaw. The dentalosplenial is elongated and narrow in the
anterior portion, extending on the most posterior portion, in which
begins the coronoid process. The pores of the mandibular sensory canal
are disposed in a unique row and reach the angular. In DGM-297-P and
Pz.UERJ 412, the left dentalosplenial possesses additional pores
(probably oral canal), which extends to the middle of the row of the
mandibular sensory canal. Additionally, in DGM-297-P and Pz.UERJ
515 the dentalosplenial has three to 11 preserved marginal and styli-
form teeth, in a unique row, beyond seven rounded and shorter cor-
onoid teeth, observed in DGM-297-P (Fig. 3). The angular is deep and
contacts the surangular through a curvilinear suture.
The surangular is small and narrow, dorsally placed to the denta-
losplenial and angular. It forms an angle posterior to the coronoid
process.
Only Pz.UERJ 408 preserved the articular and prearticular. The
Fig. 8. Quasimodichthys piauhyensis, specimen Pz.UERJ 408. A. Draw of head; B.
articular is triangular and possesses a widest posterior region, beyond a Detail of head. Scale bar equals 3 cm.
delicate curvature in the upper portion, which is longer than the lower
portion. The prearticular is elongated and narrow, but shorter than the
articular. Overall, the cleithrum is crescent-shaped, narrow and elongated but
Suspensorium and hyoid arch. The hyomandibula is irregular, it shows a variation in width and length is some specimens (in DGM-
with a fan-shaped articular facet, a wide vertical branch, a pronounced 295-P, DGM-297-P and Pz.UERJ 515 it is wider than in Pz.UERJ 514; in
preopercular process and concave anterior portion. DGM-295-P and DGM-297-P, it is shorter than in Pz.UERJ 514).
As observed in Pz.UERJ 514, the quadrate is fan-shaped, with a As observed in DGM-297-P, Quasimodichthys piauhyensis possesses
wider dorsal portion and most internally positioned, near to the lower three postcleithra. The most dorsal postcleithrum is the biggest and
suborbital. It has a rounded condyle in the most ventral portion, which longest, while the other two most ventral bones are rectangular.
articulates with the lower jaw. The quadratojugal is narrow and elon- The pectoral fins have 11–12 fine and elongated rays; the first one is
gated, stick-shaped and free from quadrate. covered by one basal fulcra and five to six fringing fulcra.
The ectopterygoid is elongated, narrow and edentulous. The en- Unpaired fins. The dorsal fin is composed by 19–20 rays, a single
dopterygoid is also edentulous, wider and concave ventrally, orna- basal and five to six fringing fulcra (Fig. 9). It is posterior to the pre-
mented with fine grooves on the suture regions. The metapterygoid is dorsal elevation, between the pelvic and anal fins. The anal fin is
irregular and wider than the endopterygoid and ectopterygoid. As the composed by nine rays, a pair of basal and nine fringing fulcra.
first ones, it is edentulous, but in Pz. UERJ 408 it is divided in two parts. Pelvic girdle and fins. The pelvic fins are composed by, at least,
The hypohyals are preserved in Pz.UERJ 408 (Figs. 4 and 8). The five short and robust rays. Only a pair of basal fulcra was preserved.
dorsal hypohyal is rounded, while the ventral one is bigger and oval. Caudal fin. The caudal fin was preserved only in DGM-295-P and
Both of them are anteriorly located in relation to the anterior cer- DGM-297-P. It is abbreviated heterocercal type (Fig. 10), possessing 23
atohyal. rays, being the first and the last covered by fulcral scales, with two pairs
The anterior ceratohyal is large and flattened, with wider ex- of basal and five to six fringing fulcra.
tremities and fine grooves in its central portion. The posterior cer- Scales. The body is covered by ganoid scales, organized on 40
atohyal is smaller than the anterior; it is fan-shaped and also orna- transversal rows in DGM-297-P (holotype) and 33 in DGM-295-P
mented with fine grooves on its anterior portion. (paratype), counted from the superior edge of the supracleithrum to the
A narrow and elongated autopalatine was observed only in Pz.UERJ superior lobe of the caudal fin. The fin/scales formula was designed to
408, which is anterior to the endopterygoid and ectopterygoid. The the holotype as:
dermopalatine bears, at least, two viliform teeth. D21
The branchiostegal rays were preserved in most of specimens, 40
P10 Pe19 A31
generally in number of seven, and ventrally located to the opercular
region. According to this formula, the dorsal fin begins in the twenty-first
Pectoral girdle and fins. The posttemporal is wide, triangular, and transverse row, the pectoral fins in tenth, the pelvic fins in nineteenth
ornamented by tubercles, crossed by pores of lateral line. In the most and the anal fin in thirty first. The ganoine layer is smooth, continuous,
posterior portion it possesses a notch that articulates with the most fine and without ornamentations or grooves. However, in most of
dorsal portion of the supracleithrum. scales, except for dorsal and ventral, the posterior border presents
The supracleithrum is elongated and narrow, and shows orna- denticulate ornamentation. All scales are strongly imbricated and pos-
mentation on the most anterior portion. sess two types of articulation: anterior processes and peg-and-socket. In

137
H.C.L.d. Paiva, V. Gallo Journal of South American Earth Sciences 88 (2018) 132–143

Fig. 9. Dorsal fin of Quasimodichthys piauhyensis, holotype DGM-297-P. Scale

Fig. 11. Juvenile 1 Quasimodichthys piauhyensis, specimen Pz.UERJ 513. Scale
bar equals 1 cm.
bar equals 1 cm.

rounded on the caudal fin base, with an elongated posterior portion,

forming a fringing-like structure.
Beyond the modified dorsal and fulcral scales, the others scales
change their shape according to the position on the body. The anterior
region is composed for big and rectangular scales, the medial region has
quadrate scales and rhomboid on posterior scales, smaller and near to
the caudal fin.

4.1. Ontogenetic series of Quasimodichthys piauhyensis

Three distinct ontogenetic stages were identified and distributed in

four distinct groups (Table 1).

4.1.1. Juvenile 1 of Quasimodichthys piauhyensis

Fig. 10. Caudal fin of Quasimodichthys piauhyensis, holotype DGM-297-P. Scale
bar equals 1 cm.
the most anterior and medial regions, the scales present a dorsal
anterior process more developed than the ventral one. Additionally, the
dorsal anterior process is longitudinally directed, with a sharp edge.
The peg-and-socket articulation is reduced, but present in the most
anterior and medial regions.
The dorsal scales bear extensions posteriorly forming a crest, best
seen before to the dorsal fin. The fulcral scales are narrower and
Material. Pz.UERJ 513 (Fig. 11).
Pz.UERJ 513 is the smallest specimen (75 mm SL), with a fusiform
and smooth body. Additionally, all scales have a fine and continuous
ganoine layer, with a smooth and straight posterior border. Some teeth
were preserved, probably from premaxilla. There is an unpaired vomer,
with small and poorly preserved viliform teeth. Also, the parasphenoid
is delicate and narrow. It is somewhat fragmented in anterior portion,
but possess a similar condition observed in Quasimodichthys piauhyensis
general description.
4.1.2. Juvenile 2 of Quasimodichthys piauhyensis
Material. Pz.UERJ 409; Pz.UERJ 514 (Fig. 12).
Pz.UERJ 514 is an articulated specimen (289 mm SL), bigger than
Pz.UERJ 513 and smaller than the others specimens. In addition, the

Table 1
Measurements (in mm) of Quasimodichthys piauhyensis specimens.
Measurements CPRM AMNH UERJ

DGM 295-P DGM 297-P AMNH FF 10012 AMNH FF 10013 Pz.UERJ 513 Pz.UERJ 514 Pz.UERJ 515

Standard Lenght 415 480 320 325 75 289 365

Head Lenght 113.62 136.42 122 114 22 85 115
Preorbital Lenght - 54.14 43.5 48.6 10 25 45
Orbit Lenght - 25.91 23.5 22.7 - 23 -
Maximum body depth 240 200 145 107.5 27 93 145
Distance from posterior border of opercle to origin of dorsal fin 137.11 165 116 202.5 30 110 120
Distance from posterior border of opercle to origin of anal fin 205 260 131.4 130 - 140 -
Distance from posterior border of opercle to origin of pelvic fin 143.16 146.01 - 95 - 95 -
Distance from origin of pectoral and pelvic fin 126.01 120.81 - 67.5 - 83 -
Height of caudal peduncle 57.02 76.84 42.8 47.5 - - -

138
H.C.L.d. Paiva, V. Gallo
Fig. 12. Juvenile 2 Quasimodichthys piauhyensis, specimen Pz.UERJ 514. Scale
bar equals 3 cm.
preopercle is narrow and its vertical arm is elongated, but no more than
observed in adults. Pz. UERJ 514 is slender, fusiform and possesses
denticulate ornamentation on posterior border of the anterior scales;
the others scales are smooth. In both Pz.UERJ 409 and Pz.UERJ 514,
dermal bones are ornamented with delicate ganoine tubercles, most
concentrated in the supraorbital bones and in the posteriormost portion
of the frontal.
4.1.3. Sub-adults of Quasimodichthys piauhyensis
Material. AMNH FF 10012; AMNH FF 10013; Pz.UERJ 408;
Pz.UERJ 410; Pz.UERJ 413; Pz.UERJ 515 (Fig. 13).
AMNH FF 10012, AMNH FF 10013 and Pz.UERJ 515 are medium-
sized specimens, reaching between 320 mm and 365 mm SL, with a
visible but delicate predorsal elevation. Pz.UERJ 410 and Pz.UERJ 413
are represented by isolated skulls and anterior scales. In AMNH FF
10012, AMNH FF 10013, Pz.UERJ 408, Pz.UERJ 410 and Pz.UERJ 515
the dermal bones are ornamented with most numerous and robust ga-
noine tubercles than those observed in Pz. UERJ 409 and Pz.UERJ 514.
The preopercle is narrow and its vertical arm is elongated, similar
condition observed in Pz.UERJ 514. The parasphenoid is more robust
than in the observed in Pz.UERJ 513, similar to adults. As observed in
Juvenile 2, the anterior scales possess denticulate ornamentation on
posterior border, condition also observed in the middle region.
4.1.4. Adults of Quasimodichthys piauhyensis
Material. DGM 297-P (holotype); DGM 295-P (paratype) (Fig. 14).
Almost complete specimens, robust and hump-backed, with more
than 400 mm SL. In both specimens, the preopercle is wider than the
other specimens. DGM 297-P has a crescent-shaped cleithrum, with a
wider posteriormost portion. All dermal bones present most numerous,
robust and dense ganoine tubercles. Additionally, in both specimens,
Fig. 13. Sub-adult Quasimodichthys piauhyensis, specimen Pz. UERJ 515. Scale
bar equals 4 cm.
139
Journal of South American Earth Sciences 88 (2018) 132–143
Fig. 14. Adult Quasimodichthys piauhyensis, paratype DGM-295-P. Scale bar
equals 5 cm.
except for the dorsal, all scales present denticulate ornamentation on
the posterior border, and the preopercle and the cleithrum are crescent-
shaped.
5. Comparisons and discussion
5.1. Morphological analysis of Quasimodichthys piauhyensis
Quasimodichthys piauhyensis is known to bear tubercles on dermal
bones, especially on the infraorbital and opercular series, which re-
semble those of Camerichthys lunae, Callipurbeckia minor, C. tendagur-
uensis, “L.” souzai, and Scheenstia mantelli. Nevertheless, in
Quasimodichthys piauhyensis these tubercles are more dense and robust
than in C. lunae, however they are delicate if compared with “L.” souzai.
Additionally, Q. piauhyensis shares more than two extrascapular bones
with Araripelepidotes temnurus, “L.” souzai and Paralepidotus ornatus.
According to Gallo (2005), a crescent-shaped preopercle should be a
typical shape for Lepidotes. However, a crescent-shaped preopercle was
also observed in C. lunae, Paralepidotus ornatus, Scheenstia laevis, S.
mantelli and S. maximus. On the other hand, Araripelepidotes temnurus,
Callipurbeckia minor, “L.” alagoensis, “L.” roxoi and Pliodetes nigeriensis
possess a ‘L’-shaped preopercle.
López-Arbarello (2012) affirmed that one of the diagnostic char-
acters of Semionotidae could be a closed circum-orbital ring. However,
this feature was also observed in all Brazilian articulated “Lepidotes”, in
addition to Araripelepidotes temnurus, Camerichthys lunae, Scheenstia
laevis, S. mantelli, S. maximus and Pliodetes nigeriensis. Generally, neop-
terygians have a circumorbital ring composed of supraorbitals, der-
mosphenotic and infraorbitals. Semionotiforms, macrosemiids and gars
could possess a varied number of anterior infraorbitals, cited on lit-
erature as lachrymal or preorbital (Olsen and McCune, 1991; Wenz,
2003; López-Arbarello, 2012).
In addition, according to the current Lepidotes diagnosis,
Quasimodichthys piauhyensis (firstly described as “L.” piauhyensis), only
closes to the genus due the presence of the middle pit line contained in
a groove excavated in dermopterotic and parietal and reduced peg-and-
socket articulation in scales. However, many ginglymodian re-
presentatives possess these characteristics.
5.2. Scales of Quasimodichthys piauhyensis
Regarding scales, the ganoids ones are the most frequent in sedi-
mentary basins, considering the age and sediments of the outcrops
where they were found (Gayet and Meunier, 1986).
Quasimodichthys piauhyensis possesses scales morphologically si-
milar to those of Araripelepidotes temnurus, “L.” alagoensis, “L.” oliveirai,
“L.” dixseptiensis and “L.” wenzae, mainly because of the continuous and
fine ganoine layer. However, the fine ganoine layer of “L.” alagoensis
could be result of the fossilization process (Gallo, 2000). Additionally,
Q. piauhyensis has a distinct posterior border of the scales, with denti-
culate margin. This characteristic differs from other analyzed
H.C.L.d. Paiva, V. Gallo
specimens, mainly A. temurus (a unique and strong projection in the
medial scales and the most anterior scales with grooves in the posterior
margin) and C. lunae (more regular forward projections, forming a
serrated margin) (Silva Santos, 1990; Gallo and Brito, 2004; López-
Arbarello, 2012; Bérmudez-Rochas and Poyato-Ariza, 2015).
Araripelepidotes temnurus and Brazilian “Lepidotes” share ganoid
scales and the peg-and-socket articulation that provided slow move-
ment capacity, indicating a life mode on shallow seas or calm waters
environment (Gallo-da-Silva, 1998).
Additionally, López-Arbarello (2012) and Murray et al. (2015)
pointed out that Lepidotes could be characterized by ganoid scales with
strongly longitudinal articulation, through ventral and dorsal anterior
processes, and reduced peg-and-socket; all of them also observed in Q.
piauhyensis.
5.3. Quasimodichthys piauhyensis dentition
Jain (1983) classified the Ginglymodi dentition on three functional
groups: strongly tritoral (Scheestia maximus and S. laevis); moderately
tritoral (Callipurbeckia minor) and non-tritoral (Lepidotes elvensis). Ad-
ditionally, the presence and quantity of styliform marginal teeth could
be indicative of a less tritoral dentition (Jain, 1983). However,
Bérmudez-Rochas and Poyato-Ariza (2015) stated that “styliform” does
not reflect the morphology of the marginal teeth and “tritoral” is a more
functional than morphological term. Beyond, they suggested a mor-
phological description of Camerichthys lunae dentition, not arguing the
functionality of heterodonty. Additionally, López-Arbarello and
Wencker (2016) divided the Ginglymodi in groups, according to the
marginal teeth morphology: conic, pencil like and graceful, high, with
globular to cylindrical crowns, and molariform. Therefore, according to
the authors, Ginglymodi teeth would be separated uniquely by mor-
phology.
Overall, many Lepidotes occurrences are based on isolated bones and
scales, being rarer teeth records (Gallo-da-Silva and Azevedo, 1998).
However, isolated teeth can be useful for proposals of the paleoenvir-
onment (Thies and Mudroch, 1996).
Applying the morphological classification in Quasimodichthys piau-
hyensis, it possesses pencil like marginal dentition and molariform
medial dentition.
5.4. Ontogenetic series of Quasimodichthys piauhyensis
The analyzed Quasimodichthys piauhyensis specimens revealed some
ontogenetic characteristics that were distributed on four distinct stages:
juvenile 1; juvenile 2; sub-adult and adult.
According to Cloutier (2010), the juvenile stage is difficult to define,
because it is a transitional phase, however the individuals of this stage
are characterized by incomplete squamation and corporal proportions
similar to adults. Among the specimens, Pz.UERJ 409, Pz.UERJ 513 and
Pz.UERJ 514 were considered the youngest Q. piauhyensis individuals.
Pz.UERJ 513 have no ornamentation, fine ganoine layer and the
minor standard length, while in both of Pz.UERJ 409 and Pz.UERJ 514
there are evidences of soft ornamentation on the posterior portion of
the frontal and on the posterior border of scales, observed only in the
anterior region. Although these last evidences had been observed on the
others Quasimodichthys piauhyensis specimens, the shape of the para-
sphenoid, preopercle and cleithrum, in addition to the absence or the
smoothest ornamentation on dermal bones and scales ornamentation
restricted to the anterior portion of the body could be also related to a
juvenile stage. However, among the observed juveniles (Pz.UERJ 513,
Pz.UERJ 409 and Pz.UERJ 514), the absence of ornamentation in
Pz.UERJ 513 allowed to suggest that it could be considered younger
than Pz.UERJ 409 and Pz.UERJ 514. Therefore, in this paper, the ju-
venile stage was divided in 1 (Pz.UERJ 513) and 2 (Pz.UERJ 409 and
Pz.UERJ 514).
As observed on juvenile 2, AMNH 10012, AMNH 10013, Pz.UERJ
140
Journal of South American Earth Sciences 88 (2018) 132–143
410, Pz.UERJ 413 and Pz.UERJ 515 possess evidences of ornamentation
on dermal bones and posterior border of the scales. However, unlike in
juvenile 2, in these specimens, herein considered sub-adults, the orna-
mentation reaches the anterior portion of the frontal and the su-
praorbital bones. Additionally, the anterior scales and some of the
medial region also present ornamentation on the posterior border. Also,
while in juveniles 1 and 2 the specimens present a fusiform body, in
sub-adults it was noted a soft pre-dorsal elevation. Although these
specimens possess some differences related to juveniles, they cannot be
considered adults yet, because these characteristics indicates not totally
differentiated individuals, and they would be transitional between ju-
venile 2 and adult, named herein as sub-adults.
Regarding the holotype and paratype, the largest specimens, they
are distinct of all others, because of the most robust and hump-backed
body, a most dense and robust ornamentation on dermal bones and
recover the entire circumorbital ring and the frontal, beyond the scales;
all of them, except the dorsal and caudal ones, possess ornamentation
on the posterior border. Regarding others specimens, the holotype and
the paratype of Q. piauhyensis seem to be the oldest individuals, con-
sidered herein the adult stage for the species.
These ontogenetic stages resemble the observed on Paralepidotus
ornatus from the Neotriassic (Norian) of Italy. Tintori (1996) and Gallo
et al. (2002) observed ontogenetic variations for this taxon and argued
that the variations in ornamentation of the dermal bones and scales are
useful on identification of ontogenetic stages. Additionally, as P. ornatus
as Q. piauhyensis show differences on dentition morphology, which re-
semble an ontogenetic transition. According to Tintori (1996) and
Gallo-da-Silva (1998), the dentition transition reflects also an en-
vironment transition along the development. Gallo-da-Silva (1998) ar-
gued that fusiform and slender body in juveniles Q. piauhyensis could be
related to a fast swim and the specimens could occupy shallower en-
vironments, while the oldest specimens, most robust and slowest, would
occupy deeper and calm waters, likely feeding on benthic mollusks.
Ontogenetic sequences are rare in fossils, except in cases of cata-
strophic events, as large floods, tornadoes, prolonged droughts, which
could reach a population and cause a non-selective death of its in-
dividuals, associated to great preservation and diagenetic conditions
(Leal, 2006). Leal (2006) suggested that catastrophic assemblages could
allow the record of distinct ontogenetic stages and are especially useful
on the studies of morphological patterns. Additionally, Petra and Gallo
(2012) argued that the Pastos Bons Formation was a saline lake en-
vironment, with seasonal drying events that caused alterations in the
water temperature, increase on salinity levels, and decrease of oxygen
rates, which could cause mass death of the Q. piauhyensis population,
explaining the record of ontogenetic stages for the taxon at the locality.
Finally, based on the observed characteristics, Quasimodichthys
piauhyensis possesses some distinct ontogenetic stages: juvenile 1; ju-
venile 2; sub-adult; adult.
5.5. Phylogenetic relationships of Quasimodichthys piauhyensis among
Ginglymodi
The parsimony analysis generated 10 most parsimonious trees,
length of 399 steps, consistency index of 0.316 and retention index of
0.695 (Fig. 15).
Thies (1989) stated that Lepidotes could be a para or polyphyletic
taxon, due the great amount and variety of described species of the
genus. However, Lepidotes is a poorly defined genus, considered a
“waste basket” taxon, because the majority of the records are based in
fragmented and isolated elements, with contradictory characters, thus
considered nomina dubia (Lombardo and Tintori, 2008; Cavin, 2010).
Besides the constant debate about the Lepidotes relationships, the
general purpose included it in Semionotiformes, most precisely in
Semionotidae family. However, López-Arbarello (2012) analyzed the
Ginglymodi group and proposed a diagnosis for Lepidotes, in which it
could be a monophyletic group, most related with Lepisosteiformes and
H.C.L.d. Paiva, V. Gallo Journal of South American Earth Sciences 88 (2018) 132–143

Fig. 15. Strict consensus of 10 most parsimonious trees, length = 399 steps; consistency index = 0.316; retention index = 0.695. Quasimodichthys piauhyensis ap-
pears in a basal position to the green clade. Bootstrap values above (if higher than 50%) for 1000 replicates and Bremer support values (below) pointed out in clades.
Italic numbers (blue point) indicated the cited nodes. (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of
this article.)

restrict to Lower Jurassic of Central Europe. Additionally, all the others
“Lepidotes” species out of the estimated range need a review, as ante-
riorly proposed in Cavin (2010), which was tested in some studies (e.g.
Cavin et al., 2013; Deesri et al., 2014; Cavin et al., 2018). In addition,
López-Arbarello and Wencker (2016) and López-Arbarello and Sferco
(2018) improved their studies and affirmed that true Lepidotes remains
in Lepisosteiformes, included in Lepidotidae, a family previously pro-
posed in Owen (1860).
However, neither López-Arbarello (2012) nor López-Arbarello and
Wencker (2016) and López-Arbarello and Sferco (2018) used Brazilian
“Lepidotes” species, some of which are housed on foreign paleontolo-
gical collections.
When Brazilian “Lepidotes” taxa were included in the López-
Arbarello analysis, they showed a non-monophyletic relationship re-
presented by homoplastic characters, incertae sedis among
Semionotiformes.
In Brazil, Lepidotes records are mainly based on isolate and frag-
mentary elements (scales and isolated bones), often assigned to
“Lepidotes” sp. Gallo and Brito (2004) stated that describing species
from isolated scales is a hazardous practice, as there may be consider-
able diversity in scale morphology (including ornamentation) even in a
single specimen. Among the Brazilian “Lepidotes” representatives, few
are articulated, but the relationships between these taxa and the type
species of the genus are debatable.
Furthermore, the phylogenetic analysis herein applied agreed with
the morphological observations about “Lepidotes” piauhyensis and

141
H.C.L.d. Paiva, V. Gallo
Fig. 16. Quasimodichthys piauhyensis reconstitution. Draw by the paleoartist
Maurilio Oliveira.
confirmed it as a distinct taxon (i.e., Quasimodichthys), which does not
belong to the true Lepidotes representatives. Therefore, it is the first
occurrence of Quasimodichthys in Jurassic of Brazil and the others
Brazilian “Lepidotes” need more studies (Fig. 16).
On the other hand, we observed a relationship between
Quasimodichthys piauhyensis (Upper Jurassic of Brazil), N. puntanus
(Lower Cretaceous of Argentina), Lepidohyas microrhis (Lower
Cretaceous of Spain) and Hoyasotes tanyrhis (Lower Cretaceous of
Spain), in the node 90 (highlighted in green, with blue point), sustained
by two homoplasies: absence of denticles in branchial and lateral
margins of the cleithrum [71 (0)] and conspicuous dorsal ridge scales
with lower spine [78 (1)].
In addition, we observed a sister-group relationship, formed by
Lepidohyas microrhis and Hoyasotes tanyrhis species, in the node 89
(highlighted in blue point), based on closed circumorbital ring, pre-
sence of dentalosplenial teeth disposed in unique row, presence of
fringing fulcra in pectoral fins, supraorbital sensorial canal running the
parietal almost in the edge. However, these characteristics are weak to
support the genus, because they were observed in other
Semionotiformes and Lepisosteiformes species.
6. Conclusions
Three distinct ontogenetic stages were observed in Quasimodichthys
piauhyensis, juvenile (1 and 2), sub-adult and adult. These stages are
similar to those observed in Paralepidotus ornatus. Although fossils re-
presenting ontogenetic sequences of fishes are rare; those preserved in
mass mortalities are specially useful on studies about the evolutionary
morphological transformations. The fossil assemblage recovered in
Pastos Bons Formation offer an unparalleled opportunity to study the
morphological transformation through out the ontogeny of Q. piau-
hyensis. Quasimodichthys piauhyensis presents some morphological
characteristics related to L. elvensis, the type-species. Although López-
Arbarello (2012) affirmed that Lepidotes was restrict to Lower Jurassic
of Central Europe and formed a monophyletic group, the inclusion of
Brazilian taxa showed that not only Lepidotes, but Ginglymodi needs
more studies for a better comprehension about its taxonomic relation-
ships.
Quasimodichthys piauhyensis appears in a basal position to the clade
formed by (N. puntanus (Lepidohyas microrhis, Hoyasotes tanyrhis)), on
the basis of absence of denticles in branchial and lateral margins of the
cleithrum and conspicuous dorsal ridge scales with lower spine.
Acknowledgments
We would like to thank MSc. Rodrigo da Rocha Machado (CPRM),
Dr. Hugo Ricardo Secioso Santos (UERJ), Deise Dias Rêgo Henriques
(Museu Nacional/UFRJ), John Maisey (AMNH), Emma Bernard
(NMUK), and Gaël Clement (MNHN) for the access to paleontological
collections, the photographers Regiana Salgado (PROATEC-UERJ),
Kevin Webb (NMUK) and Philippe Loubry (MNHN), the paleoartist
142
Journal of South American Earth Sciences 88 (2018) 132–143
Maurilio Oliveira (Museu Nacional/UFRJ), and to Rodrigo Tinoco
Figueroa (UERJ) for the revision of the manuscript. HCLP thanks
Programa de Pós-graduação em Ecologia e Evolução (UERJ) for her
MSc. studies. HCLP was supported by Capes-DS. VG is supported by
CNPq and have an UERJ/FAPERJ Prociência grant.
Appendix A. Supplementary data
Supplementary data related to this article can be found at https://
doi.org/10.1016/j.jsames.2018.08.010.
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