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Tahnaichthys magnuserrata gen. and sp. nov., a double-hump pycnodontid

(Actinopterygii, Pycnodontiformes) fish from the Albian limestones of the Tlayúa
Quarry, Puebla, Mexico.

Stephanie Pacheco-Ordaz, Omar Mejía, Jesús Alvarado-Ortega

PII: S0895-9811(24)00499-1
DOI: https://doi.org/10.1016/j.jsames.2024.105277
Reference: SAMES 105277

To appear in: Journal of South American Earth Sciences

Received Date: 21 September 2024

Revised Date: 22 November 2024
Accepted Date: 23 November 2024

Please cite this article as: Pacheco-Ordaz, S., Mejía, O., Alvarado-Ortega, J., Tahnaichthys
magnuserrata gen. and sp. nov., a double-hump pycnodontid (Actinopterygii, Pycnodontiformes) fish
from the Albian limestones of the Tlayúa Quarry, Puebla, Mexico., Journal of South American Earth
Sciences, https://doi.org/10.1016/j.jsames.2024.105277.

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Tahnaichthys magnuserrata gen. and sp. nov., a double-hump pycnodontid (Actinopterygii,

Pycnodontiformes) fish from the Albian limestones of the Tlayúa Quarry, Puebla, Mexico.

Stephanie Pacheco-Ordaz1*, Omar Mejía2, and Jesús Alvarado-Ortega3

1. Posgrado en Ciencias Quimicobiológicas, Escuela Nacional de Ciencias Biológicas,

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Instituto Politécnico Nacional, Unidad Professional Lázaro Cárdenas. Prolongación de

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Carpio and Plan de Ayala S/N, Colonia Santo Tomás, Alcaldía Miguel Hidalgo, Ciudad de

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México, 11340 México. stephy.pacheco@gmail.com
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2. Departamento de Zoología, Escuela Nacional de Ciencias Biológicas, Instituto

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Politécnico Nacional, Unidad Professional Lázaro Cárdenas. Prolongación de Carpio and

Plan de Ayala S/N, Colonia Santo Tomás, Alcaldía Miguel Hidalgo, Ciudad de México,
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11340 México City, 11340 México. hmejiagpn@ipn.mx

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3. Instituto de Geología, Universidad Nacional Autónoma de México. Circuito de la

Investigación S/N, Ciudad Universitaria, Alcaldía Coyoacán, Ciudad de México, 04510

México. alvarado@geologia.unam.mx

* Corresponding author
Abstract

Tahnaichthys magnuserrata gen. and sp. nov. is described here based on three well-

preserved Albian specimens recovered in the lagoon-coastal deposits of the Tlayúa Quarry,

Puebla, Mexico. This species belongs to the order Pycnodontiformes and shows the

diagnostic features of the family Pycnodontidae, including the presence of a posterior

parietal process distally branched, toothless reniform maxilla, anterior thin wings at the

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base of the haemal and neural spines, and slender dentary with two teeth. In addition, this

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species shows unique features never observed before in other pycnodontids as the presence

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of two unequal predorsal humps or prominences, including a slight hump just behind the
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occiput and a large hump rising in front of the dorsal fin. The predorsal scutes are thick,
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have a longitudinal row of unequal spines, and show thin lateral wings projected ventrally,
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covering a large part of the predorsal trunk region. In the new species, the modified bar-like

scales of the predorsal region have tiny posterior wings ornamented with tubercles scattered
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or arranged in dorsoventral short ridges. In addition, the premaxilla of this new species is
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ventrally broad and has a large single incisiform tooth opposing two small, oval dentary

teeth. This new species exhibits a peculiar mixture of osteological features that, for the

moment, complicate its taxonomic designation at the suprageneric level (subfamily or

tribe). Although this is the second pycnodontiform species from Tlayúa formally described,

the present discovery increases the taxonomic diversity of the pycnodontids in North

America and the Paleogulf of Mexico domain.

Keywords: Albian, Tlayúa, Tahnaichthys, Pycnodontidae, Puebla, Mexico.

 INTRODUCTION

The order Pycnodontiformes is a monophyletic clade of fishes (about 60 genera

organized into eight families) erected by Berg (1937, 1940) to include Pycnodus Agassiz,

1833, and close relatives. During nearly 175 million years, between the Norian (Late

Triassic) and the Middle Eocene (Paleogene), these successful actinopterygian fishes

inhabited shallow tropical and temperate seas around the world (Nursall, 1996a, 1999;

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Martin-Abad and Poyato-Ariza, 2013; Cawley and Kriwet, 2017; Cooper and Martill,

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2020), and even some of them were able to venture into freshwater environments near the

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coasts (Cavin et al., 2020; Cawley and Kriwet, 2024).
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This extinct incertae sedis clade has an amazing Bauplan combining derived and
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primitive osteological features. This fact hinders the recognition of the intra and
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interrelationships of the pycnodontiforms (Nursall, 1996a, b). Pycnodontiforms share a

high, rounded, and laterally compressed body. Their vertebrae are unossified, and the
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notochordal canal separates the ventral and haemal arcocentra along their trunk. Their
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hypertrophied and complex mesethmoid bone occupies most of the ethmoid skull region, is

T-shaped in a longitudinal section, and consists of a large medial posterior thin lamina plus

an anterior transversal lamina expanded laterally to both sides. Moreover, in their

mandibles, the principal dental occlusion is tripartite and includes a battery of massive,

rounded teeth aligned in both branches of the lower jaw and the vomer (Nursall, 1996a, b,

1999; Poyato-Ariza and Wenz, 2002).

In his of the Mexican pycnodontiforms, Applegate et al. (2004) concluded that the

Tlayúa Quarry is a North American hot spot to study this extinct group, not only for its

well-preserved fossils but also because these constitute a predominant component in the
ichthyofauna of this site, with potential new taxa still undescribed. Despite this, the

previous formal studies of Mexican pycnodontiforms have been scarce and, up to now,

limited to the description of Tepexichthys aranguthyorum Applegate, 1992, and the

identification of a vomerine toothed plate possibly corresponding to the genus

Neoproscinetes Silva Santos, 1970 (Applegate, 2001). Given this historical context, this

paper aims to describe and discuss the possible affinities of a new species of

pycnodontiform fish discovered in the Albian limestones of the Tlayúa Quarry near Tepexi

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de Rodríguez, Puebla, central Mexico.

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Formal studies of other pycnodontiform fossils discovered elsewhere in Mexico (in

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paleontological sites within the states of Chiapas, Hidalgo, Oaxaca, Puebla, San Luis
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Potosí, Coahuila, and Nuevo León) are equally scarce. Sapper (1899) reported the presence
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of fossil fish remains preserved in the marly slabs incorporated in different archaeological
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contexts within the Maya city of Palenque, Chiapas. Later, Mülleried (1951) suggested that

these Palenque fossils are Eocene remains of the genus Pycnodus. Alvarado-Ortega et al.
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(2018) recently corroborated such generic taxonomical identification and recovered

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numerous specimens of these fishes from the Danian deposits of the Tenejapa-Lacandón

geological unit. Otherwise, Dunkle and Maldonado-Koerdell (1953) discovered the dental

elements of these fishes in the Jurassic deposits exposed in the Tamán region, San Luis

Potosí, which they conferred Gyrodus macrophthalmus Agassiz, 1843. Alvarado-Ortega et

al. (2014) reported Gyrodus sp. based on similar Kimmeridgian fossils from the Sabinal

Formation recovered near Tlaxiaco, Oaxaca. Coniacian and Turonian fossil remains of

Paranursallia gutturosa (Arambourg, 1954) and Nursallia tethysensis Capasso, Abi Saad,

and Taverne, 2009, have been recovered from different outcrops of the Agua Nueva and the

Austin formations along the states of Nuevo León, San Luis Potosí, and Puebla (Blanco-
Piñón, 2003; Giersch, 2014; Giesrch et al., 2008; Stinnesbeck et al., 2019; Alvarado-Ortega

et al., 2019, Galavíz-Hernández, 2022). Alvarado-Ortega et al. (2020) reported

indeterminate remains of Nursallia Arambourg, 1954, from the Campanian deposits of the

Angostura Formation exploited in the Tzimol Quarry near Comitán, Chiapas.

The Tlayúa Quarry is a Mexican emblematic lagerstätte site with a predominance of

bony fishes (Martill, 1989; Alvarado-Ortega et al., 2007). In addition, the fossil assemblage

of Tlayúa includes ichnites, wood, pollen, plant impressions, algae, foraminifera, ostracods,

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sponges, corals, annelids, gastropods, bivalves, ammonites, belemnites, crustaceans,

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arachnids, insects, echinoids, holothuroids, crinoids, ophiuroids, and reptiles such as turtles,

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crocodiles, lizards, tuataras, and pterodactyls (e.g., Applegate, 1966; Seibertz and Buitrón,
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1987; Feldmann et al., 1998; Alvarado-Ortega, 2005; Reynoso et al., 2000; González-
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Rodríguez et al., 2004; Applegate et al., 2000, 2006; Alvarado-Ortega et al., 2006; Herrera-
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Flores, 2024).

The Tlayúa Quarry is a mined outcrop of the Middle Member of the Tlayúa Formation,
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described by Pantoja-Alor (1992) based on the limestone deposits exposed along the Tlayúa
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Ravin, near to the east of Tepexi de Rodríguez town, Puebla. Kashiyama et al. (2004)

suggested that this member of the Tlayúa Formation represents an Aptian open-ocean inter-

cratonic deposit. Nonetheless, other numerous studies show a compelling alternative

scenario, proposing that this sequence represents an Albian shallow tropical costal pot-

lacunar, with an almost permanent connection to the sea alternated with sporadic periods of

isolation, short episodes of freshwater influence, and occasional subaerial exposure events

(e.g., Applegate et al., 1984; Cantú-Chapa, 1987; Pantoja-Alor, 1992; Espinosa-

Arrubarrena and Applegate, 1996; Benammi et al., 2006; Riquelme et al., 2021; Solé et al.,
2022). A paleoenvironmental proxy for the fossiliferous strata of Tlayúa is the modern

Great Banks of the Bahamas (Juárez-Arriaga et al., 2023).

MATERIAL AND METHODS

Fossil preparation— The mechanical and chemical procedures, expertly described by

Toombs and Rixon (1959), allowed the preparation of the specimens referred to in this

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work. The fossil preparation includes 12 hours of immersion in a 5-10% acetic acid

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aqueous solution saturated with dibasic phosphate and 12 hours of posterior baths in clean,

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running fresh water. Under the stereo microscope, fine brushes helped with the punctual
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application of this solution on the fossil. We repeated this chemical procedure until we
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achieved the total remotion of limestone from the fossil surfaces. We used dental
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excavators and dissecting needles to remove the remaining small limestone patches

mechanically. Both parts of the holotype described here are transferred to a crystal-clear
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polyester casting resin.

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We glued the broken and fractured bones with cyanoacrylate and hardened the fossil

structures with plexigum and ethyl acetate solution. The specimens were analyzed by the

naked eye, under a stereo microscope, and through high-resolution photographs obtained

under different lighting and contrast conditions, including natural, white, and long-wave

UV (254 nm) lights. In addition, we studied high-contrast images of gray tones obtained

with the specimens coated with magnesium and X-ray plate images of the specimens

transferred to resin.

Institutional abbreviations— The following institutional acronyms are mentioned in this

work. IGM, Colección Nacional de Paleontología at the Instituto de Geología (anteriorly

known as Instituto Geológico de México) of the Universidad Nacional Autónoma de

México. IHNFG, Museo de Paleontología "Eliseo Palacios Aguilera”, Tuxtla Gutiérrez,

Chiapas, Mexico. MUPAL, Museo de Palenque of the Instituto Nacional de Antropología e

Historia (INAH), Chiapas, Mexico.

Anatomical nomenclature— The anatomical terminology and the abbreviations

considered here follow those used in similar previous manuscripts (e.g., Nursall, 1996a, b,

1999; Kriwet et al., 1999; Poyato-Ariza and Wenz, 2002; Capasso et al., 2009; Taverne and

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Capasso, 2021).

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Comparative material— This work includes the following fossils with comparative

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purposes. Nursallia sp. Blot, 1987: IGM 11401 from the Tzimol Quarry, an outcrop of the
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Campanian deposits of the Angostura Formation near Comitán, Chiapas, Southeastern
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Mexico (Alvarado-Ortega et al., 2020). Tepexichthys aranguthyorum Applegate, 1992:

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IGM 3286 (holotype) and IGM 3288 to IGM 3300 (paratypes), from the Albian deposits of

the Tlayúa Quarry, Puebla, Mexico. Pycnodus sp. Agassiz, 1833: IGM 4551 and IHNFG-
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4890, from the Danian deposits of the División del Norte Quarry, Chiapas, Mexico; two
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flagstones with fish mass mortalities of the Danian deposits of the Tenejapa-Lacandón

Unit, the MUPAL 2587 with two specimens and MUPAL 2731 with seven specimens, both

recovered in the Archaeological context of the Mayan city of Palenque (Alvarado-Ortega et

al., 2015, 2018).

RESULTS

Systematic paleontology
Subclass Actinopterygii, Cope, 1887.

Division Halecostomi Regan, 1923, sensu Patterson, 1973.

Orden Pycnodontiformes Berg, 1937.

Suborder Pycnodontoidei Nursall, 1996, sensu Poyato-Ariza and Wenz, 2002.

Family Pycnodontidae Agassiz, 1833, sensu Nursall, 1996a.

Tahnaichthys, gen. nov.

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Nomenclatural Act — LCID XXXXXXXXXXX

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Type species— Tahnaichthys magnuserrata sp. nov., described below.
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Diagnosis— As in the type species below.
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Etymology– The name of the genus is composed of the words Tahná and ichthys. "Tahná"
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is the original name that the Popolocas gave to Tepexi el Viejo ancient locality, which gave

rise to the current Tepexi de Rodríguez. This word means hill, small mountain, or hump.
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"Ichthys" in Greek means fish. Thus, this generic name means "the Tahná fish" or "the
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humpback fish."

Tahnaichthys magnuserrata sp. nov.

(Figures 2-10, Table 1)

Nomenclatural act— LCID number provided by ZOOBANK

Holotype— IGM 14024, a complete specimen of 165.2 mm total length, with no additional

field data, preserved in part (IGM 14024a) and counterpart (IGM 14024b), and transferred

to resin (Figures 2 and 3).

Paratypes— IGM 14025, complete specimen with nearly 73.74 mm total length, preserved

in part (IGM 14025a) and counterpart (IGM 14025b), and recovered in the section of the

Tlayúa Quarry IGM-loc 1971 and stratigraphic level Z-8 (Figure 3A). IGM 14026, a nearly

complete specimen with 84.08 mm of total length, from the IGM-loc 2828 quarry section

and stratigraphic level Z-XVII (Figure 3B).

Occurrence— Albian laminar limestones of the Middle Member of Tlayúa Formation.

Tlayúa Quarry, near Tepexi de Rodríguez, Puebla, Mexico.

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Etymology—The name derives from two Latin words, "magnus" or large and "serrata" or

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serrated. The generic and specific names mean "the Tahná fish with large saws or the

humpback fish with large saws."

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Diagnosis— A member of the family Pycnodontidae that exhibits a unique combination of
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features, the presence of two dorsal prominences or unequal humps bordering the predorsal
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region of the trunk, predorsal bordering scutes with broad, uneven lateral wings covering

the laterodorsal flank of the humps extensively, and the modified, bar-like scales of the
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predorsal region have tiny wings ornamented with aligned tubercle or ridges. In addition,
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this fish has a combination of non-exclusive characters that includes the elongated head

occupying about half of the standard length; elongated muzzle, premaxillary bone with a

wide incisiform tooth, dermocranial fenestra, distally branched posterior parietal process or

pinniculus parietalis, slender triangular vomer with three rows of teeth, elongated

prearticular with three tooh rows, dentary with two small, rounded incisive teeth; all fins

are rounded; pelvic fin is small, opposes the posterior predorsal hump, and rises near the

beginning of the posterior third of the body, the unpaired fins oppose to each other and

extend into the last fifth of the body length; there are 28-29 dorsal fin rays; the anal fin has

22-23 rays; the vertebral column with 23 small, rounded precaudal arcocentra, including 13
abdominal arcocentra and 10 caudal arcocentra; adjacent dorsal and haemal arcocentra have

simple lateral contact and not invading the notochordal canal; neural and haemal spines

anterior of caudal-peduncle with anterior triangular wings; small caudal peduncle including

only two haemal and two neural spines; caudal skeleton with six epichordals, eleven

hypochordals and two urodermals; anal and predorsal scutes with 5-7 robust conical spines

of increasing size and placed closely side by side on a single longitudinal row; peltate scale

pattern; bar-like predorsal scales and broad abdominal scales ornamented with tubercle

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scattered or arranged in dorsoventral short ridges; cloacal vestibule bordered by two

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acicular-like scales and with no bifid scale; caudal formula iii+I+20+I+iii.

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Description
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General features and proportions— Table 1 summarizes the measurements and

proportions observed in the specimens studied here and their averages considered to
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describe the present new species. The holotype (IGM 14024) has 135.53 mm of standard
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length (SL, considered in the straight line drawn from the anterior end of the snout to the

middle part of the notochordal canal where the proximal tips of the most anterior caudal

procurrent rays appear) and is the largest specimen. In contrast, the paratype IGM 14025 is

the smallest and only has 60.41 mm of SL. This fish is oblong-bodied, tapered anteriorly,

rounded posteriorly, and shows two unequal predorsal humps protruding upward between

the occiput and the dorsal fin. The anterior predorsal hump is just behind the head and

noticeably less elevated than the posterior hump. The posterior and largest predorsal hump

is as high as the distal margin of the dorsal fin and coincides with the dorsal apex of the
trunk at 70% of the SL on average. On average, the maximum body height represents 68%

of the SL.

In this work, the length of the head is measured in a longitudinal line between the

anterior end of the snout and the posterior margin of the cleithrum. In contrast, the height of

the head is considered in the transversal line that goes from the most dorsal end of the skull

to the ventral tip of the cleithrum. So, the length and height of the elongated triangular head

of this fish represent 51 and 49 % of the LS on average, respectively. The preorbital or

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snout length is elongated, tapers anteriorly, occupies almost two-thirds of the head, and, on

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average, represents 30 % of the SL. In the dorsal half of the skull, the orbit is relatively

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tiny, barely occupies a fifth of the head length, and on average, the orbital diameter is only
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9 % of the SL. Its postocular region occupies the remaining 12 % of the head length.
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Besides, in the lateral view, the skull of this fish is somewhat trapezoidal and strongly tilted
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anteroventrally. On average, the skull length and height are 48 and 35% of the SL,

respectively. Here, the preorbital, orbit, and postorbital skull parts are about three-fifths,
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one-fifth, and one-fifth of the skull length.

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All the fins are rounded. The small pectoral arises far from the ventral edge, at one-third

of the distance between the abdomen and the vertebral column, and probably extends up to

the anterior half of the abdominal region. The other fins rise in the posterior half of the

body. On average, the small pelvic fin at 65% of the SL, close to the abdominal edge and

the anal fin, opposing the posterior predorsal hump. The dorsal fin is relatively long,

extends into the posterior quarter of the body, and represents 18.49% of the SL on average,

rising between 82 and 99% of the SL. The anal fin is shorter and opposes the dorsal fin,

represents 16 % of the SL on average, and extends between 82 and 98 % of the SL. The

short and shallow caudal peduncle is bordered by the last anal and dorsal rays that extend
posteriorly, reaching the caudal fin, and only represents the posterior 0.95 % of the SL on

average. The caudal fin is broad and fan-shaped but less elevated than the trunk.

Neurocranium— The specimens studied here only expose the lateral surface of the

neurocranium. The skull bones are ornamented intensely with tubercles and reticulated

ridges. The endocranial elements are entirely covered. The lateral view of the neurocranium

is somewhat rectangular, tilted anteroventrally, and its preorbital region is elongated. Here,

the small orbit is at the end of the middle third of the neurocranium length and occupies

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one-quarter of the neurocranium height. The dermosupraoccipital, parietals, frontals,

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dermopterotics, autosphenotics, and dermosphenotics attach firmly through sharp and deep

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sinuous sutures (Figure 4). A small oval dermocranial fenestra pierces the neurocranium
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laterally in the dermosupraoccipital, frontal, and parietal tripartite junction. The
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dermosupraoccipital is triangular and forms the upper dorsal corner of the neurocranium,
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posterodorsal the orbit. The frontal is a short, curved bone slightly inclined posterodorsally,

with anterior and posterior ends expanding and bordering the orbit's dorsal half. The
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parietal is an X-shaped bone expanded in the posterior part of the neurocranium, ventral to
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the dermosupraoccipital. The pinniculus parietalis of the parietal is a posterior parietal

process, elongated, slender, and poorly branched distally.

The dermopterotic is a broad trapezoidal bone that occupies most of the neurocranium

postorbital region and extends from the orbit to the nape. Ventrally, the autosphenotic and

dermosphenotic are small bones at the posterodorsal edge of the orbit. The extrascapular

exposes a small triangular lateral surface sutured to the posteroventral edge of the

dermopterotic.

Anteroventral to the orbit, much of the neurocranium consists of the mesethmoid,

parasphenoid, and vomer. The mesethmoid is a complex bone; it includes an expanded

anterior lamina laterally to both sides, forming the anterior edge of the head, plus a

hypertrophied middle lamina that supports other snout bones and occupies almost the entire

anteroventral half of the neurocranium. The mesethmoid has a deep V-shaped opening of

the olfactory cavity in the anteroventral surface of its anterior lamina. Unfortunately, the

parasphenoid is elongated and almost entirely covered. Ventral to the orbit, the

parasphenoid shows an inflection that defines its two regions; its anterior descending

section is laminar and projects anteroventrally, reaching the vomer and the anterior ends of

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the vomer and mesethmoid, while its postorbital part extends posteriorly, forming part of

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the neurocranium base. The prefrontal is relatively thin and long, extends below the frontal

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along and above the mesethmoid, and its anterior end reaches the premaxilla (Figure 4).
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The vomer has a dorsal wing attached to the parasphenoid, plus a triangular massive ventral
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teeth-bearing plate that tapers anteriorly. The vomer bears three longitudinal rows of round-
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oval vomerine teeth that become slightly smaller in posteroanterior order. Concentric

crenulations and tiny central grooves ornament the crushing surface of these teeth (Figure
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6). Among these vomer tooth rows, the middle row has 12 rounded small teeth, and the
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lateral rows have only 11 round-oval teeth (Figures 5).

Orbital bones— Two thin sclerotic bones form a ring that occupies the orbit (Figure 4).

The circumorbital series is open and involves atrophied bones. Supraorbital bones are

absent (Figure 5). The skull incorporates the dermosphenotic. A robust, slightly

ornamented, and triangular infraorbital bone borders the anteroventral part of the orbit;

other infraorbitals are reduced to poorly ossified tubular structures scattered below the orbit

and on the lateral surface of the ethmoid bones (Figure 4).

Upper Jaws— The upper jaw consists of the premaxilla and maxilla. The premaxilla is a

triangular bone with a broad alveolar end and a sharp dorsal process exposed laterally to the
anterior edge of the ethmoid skull region (Figure 4 and 5). This bone occupies the

anteroventral third of the snout. It is three to four times higher than long, slightly curved,

and dorsoposteriorly tilted. A single wide and shallow incisiform tooth occupies most of

the premaxillary alveolar edge. The maxilla is a longitudinal oval, thin, smooth, and

toothless bone with a broad dorsal notch. The maxilla covers the labial surface of the

anterior half of the vomer behind the premaxilla.

Lower jaw— Each branch of the lower jaw includes a dentary, prearticular, angular, and

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articular. These bones form a stout, elongated triangular structure that exhibits a low

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symphysis, a high coronoid process, and a small postarticular process. The lower jaw

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extends half of the snout and joints the quadrate and symplectic bones below and slightly
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ahead of the orbit (Figure 4).
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Although both dentary bones occlude against the premaxillae, the principal jaw
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occlusion in this fish is tripartite and includes the vomer and both prearticular bones. The

dentary is a high acicular curved bone in the anterior part of the lower jaw that bears two
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small conical teeth. The prearticular is triangular and forms most of the lower jaw length
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and the high coronoid process. The prearticular shows a dental battery consisting of three

rows of oblong crushing teeth with surrounding crenulations and middle grooves

ornamenting their occlusal surfaces. The prearticular teeth are small and slightly decreasing

in the posteroanterior direction. Among these, the medial row has 12 teeth about three times

wider than long, while the lateral rows have smaller, oval, and poorly crenulated teeth. The

prearticular sutures the small angular and articular bones in the posteroventral part of the

jaw (Figures 5 and 6).

Suspensorium— The suspensory bones are well preserved and partially exposed (Figure

4). The quadrate is a robust triangular bone, laterally sutured with a slender and elongated
symplectic bone; together, these articulate the articular facet on the postarticular process of

the lower jaw. Most of the dorsal border of the quadrate sutures the broad rectangular

endopterygoid bone. In contrast, the ectopterygoid is a small, smooth, crescent-shaped bone

that attaches to the anterodorsal border of the quadrate. The metapterygoid is a broad bone

mostly covered below the orbit and sutures the ventral region of the parasphenoid

inflection.

The hyomandibula is a robust bone with an elongated and sinuous articular head. The

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opercular process is not present in the posterior edge of this bone. Its ventral part sutures

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and is covered by the preopercle. The exposed and dermalized dorsal part of the

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hyomandibular is marginally smaller than the preopercle; it is exposed laterally and ornated
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with deep pores and ridges like those present in the dermal bones of the skull.
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Opercular bones and hyoid— The opercle is a crescent-shaped bone, small and high, with
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sharp ends, and placed ahead of the dorsal half of the cleithrum. The preopercle is a flat

trapezoidal bone, longer than high, ventrally expanded, with a horizontal and dorsal sinuous
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suture to join the dorsal and dermal parts of the hyomandibula. The ornaments of the
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opercular and preopercular surfaces resemble those of the hyomandibula (Figure 4). This

new taxon, as all Pycnodontidae has no subopercle or infraopercle.

Ventral to the preopercle, some bones of the hyoid arch are recognizable as the

hypohyals, ceratohyals, and branchiostegal rays (Figures 4, 5). The ventral and dorsal

hypohyals are small triangular bones articulating the anterior ceratohyal. The anterior

ceratohyal is a large, robust, rectangular bone with a middle constriction. The posterior

ceratohyal is also rectangular but smaller than the anterior ceratohyal. The two

branchiostegal rays are elongated, thin, and flat bones extending almost horizontally and

articulating the posterior ceratohyal laterally. In IGM 14024, some pharyngeal teeth are
exhibited under UV light and through the cheekbones. These teeth are robust, oval,

dorsoventrally flattened, and show a sharp, anteriorly bent apex (Figure 6).

Vertebral column— The vertebral column has an incomplete ossified centra and the

notochordal canal is exposed. Instead, the trunk axis includes 23 pairs of neural arcocentra

(Table 1), including 13 abdominals and ten caudals (opposed by the respective haemal

arcocentra). An indeterminate number of these arcocentra constitute the axis of the caudal

skeleton, which in the same way are recognized by the ossified epichordal and hypochordal

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elements. The anterior seven arcocentra seem fused, forming a synarcual, and their

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respective neural spines are autogenous. The posterior neural and haemal spines fuse the

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arcocentra (Figures 2 and 9). Ten long and curved rod-like ribs enclose most of the
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abdomen, with posterior laminar or wing projection in their proximal half.
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Haemal and neural arcocentra are rounded, massive, and smooth. These do not surround
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the notochordal canal and are joined laterally with the adjacent ones through a simple

contact that involves intertwining an anterior and a posterior zygapophysis for each
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arcocentra. Each zygapophyses shows a short, triangular extension or flange.

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The haemal and neural spines located anteriorly to the caudal peduncle have anterior

sagittal flanges. These thin wings are triangular and rise from the anterior flange of the

respective arcocentra. These wings are inconspicuous in the first three or four neural spines;

however, further back and anterior to the caudal peduncle, the height of haemal and neural

spines and their respective wings decreases harmoniously according to the curvature of the

posterior edges of the trunk. The postcoelomic is an acicular and smooth bone. The last

postcloacal scutes cover most of the ventral expansion of this bone. Its dorsal end is nearly

straight, longer than the anterior most pterygiophores of the anal fin, and ends far from the

vertebral column (Figures 2 and 9).

 The caudal peduncle is short and involves two neural and two haemal arcocentra. The

spines of these arcocentra have significantly reduced anterior wings (Figure 9).

Remarkably, the peduncular neural spines are shorter than the adjoining neural and

epichordal spines, and their length is half of the opposite haemal spines. Besides, the

lengths of the peduncular haemal spines are intermediate between the shorter anterior

adjacent haemal spines and the elongated hypochordal that supports the caudal rays.

Pectoral fin and girdle— The pectoral fin comprises 19 to 22 distally segmented and

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branched rays. This fan-shaped fin has a narrow base, and its rounded and broad posterior

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part covers the anterior half of the abdominal length. The proximal ends of the pectoral rays

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joined the 6 or 8 elongated pectoral radials with broadened ends, ordered as a bouquet with
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a rounded base (Figure 7).
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The cleithrum is the most prominent bone of the pectoral girdle. This bone extends two-
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thirds of the posterior height of the head, from the abdominal margin to the articulation of

the skull and vertebral column (Figures 4 and 8). The cleithrum is crescent-shaped, concave
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anteriorly, acute dorsally, and expanded ventrally in lateral view. The pectoral fin arises in
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the middle of the trunk, between the ventral edge and the vertebral column. At the posterior

end of its ventral third, the cleithrum shows a shallow posterior pectoral notch, in which the

base of the pectoral fin is nested. The dorsal and ventral ends of this notch do not show

conspicuous processes. The medial bones of the pectoral girdle are mostly covered.

The supracleithrum and posttemporal are oblong bones dorsal to the cleithrum, slightly

inclined anterodorsally, and barely in contact. The supracleitrum is nearly a quarter of the

height of the cleithrum, lies behind the head, overlaps the dorsal tip of the cleithrum, and

covers the most anterior elements of the vertebral column. The posttemporal extends

behind the parietal and extends up to the pinniculus parietalis. Like in the skull, small
tubercles and reticular ridges ornament the lateral surfaces of the supracleithrum,

posttemporal, and ventral half of the cleithrum.

Pelvic fin and girdle— The pelvic fin is small and fan-shaped with a rounded posterior

border and narrow base. This fin probably has five distally segmented and branched rays

that barely reach the cloacal cavity (Figures 8 and 10). An indeterminate number of tiny

radials articulate the proximal ends of these rays. The broad and imbricated abdominal

scales cover most of the pelvic bones, which are triangular and taper anteriorly. The pelvic

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girdle is close to the ventral trunk edge, slightly ahead of the cloacal cavity, and opposes

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the posterior predorsal hump.

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Dorsal fin— Both unpaired fins are in the posterior fifth of the SL. The dorsal fin is
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rounded and a little longer and larger than the anal fin. This involves 29 dorsal rays and
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rises behind the posterior predorsal hump and dorsal apex of the trunk (Figure 2, Table 1).
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The dorsal fin expansion is barely smaller than the dorsal part of the trunk caudal region,

and posteriorly, it overhangs the level of the most anterior caudal fin rays. All the dorsal fin
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rays are long, segmented, and branched except for the first, which is notably short. The rays
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4-6 are the more extended in the fin, while the anterior and posterior ones are progressively

smaller. The dorsal fin support includes a series of 29 proximal pterygiophores that relate

1-to-1 with the dorsal fin rays. Such long, straight, and flat pterygiophores have small

articular heads and penetrate a little bit into the interneural spaces. The pterygiophores of

the dorsal and anal fins are symmetrically opposed.

Anal fin— The anal fin is also rounded and entirely opposes the dorsal fin (Figure 2). The

fin consists of 22 to 23 anal rays. The first is short, unbranched, and unsegmented, while

the others are elongated, distally segmented, and branched.

 The anal fin support consists of a series of 22 to 23 proximal pterygiophores that relate

1-to-1 with the anal rays. These long anal pterygiophores are straight, evenly broad, have

small articular heads, and penetrate deep into the trunk, barely exceeding the distal end of

the haemal spines.

Caudal fin— Although poorly preserved in all specimens, a large fan-shaped caudal fin

rises behind a short, shallow peduncle. This fin has no recognizable separate lobes, is

slightly higher than long, equals the half maximum body height, and shows a harmoniously

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convex distal edge (Figure 9). IGM 14026 only preserves a small ventral part of this fin

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(Figure 3); however, this is dorsal and ventrally contracted in IGM 14025, causing it to

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appear to be shallower than the caudal fin of IGM 14024.
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The caudal fin has 28 rays ordered in the formula iii+I+20+I+iii (Figure 9, Table 1). The
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posterior haemal spine supports the lower procurrent rays of this fin. Here, the procurrent
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rays are unsegmented and unbranched, both principal rays are segmented, and all the inner

caudal rays are segmented and intensely branched distally.

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The caudal skeleton comprises two small oval urodermal bones, eleven hypochordals,
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and six epichordals (Figure 9). Both hypochordals and epichordals rise from broad and

rounded arcocentra that show decreasing sizes in an anteroposterior order, have no

projections invading the notochordal canal, and laterally, these barely have contact with

each other. Although caudal-fin rays cover the distal ends of epichordals, these elements

are shaped similarly but are longer and progressively thinner than the adjacent neural

spines. Besides, the hypochordals are similarly long but slightly broader than the last

haemal spines. The hypochordals 1-7 occupy the ventral and posterior part of the caudal

skeleton, whereas the hypochordals 8-11 form its dorsal part. The hypochordals 1-5 are

spatuliform and have a similar anterodorsal inclination to the posterior most haemal spines.
The hypochordals 6-8 are triangular and become more horizontal, and their size tends to be

smaller. Finally, the hypochordals 9-11 become spatuliform, smaller, and tilted

anteroventrally.

Scales— The scales cover the trunk following the peltate pattern previously defined by

Nursall (1996a. ch. 39, fig. 19). The caudal trunk region is entirely nude. On the contrary,

scales of two kinds cover the anterior part of the trunk. Above the vertebral column,

between the occiput and the dorsal fin base, 11 rows with no more than five rod-like

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modified scales each poorly cover the predorsal trunk region, while below, 15 rows of

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expanded scales cover the entire abdominal trunk region (Figure 10).

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The bar-like predorsal scales are high, slightly curved, tilted anterodorsally, and have
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ends tightly sutured, forming continuous rows. These modified scales show tiny posterior
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wings with aligned lateral apicobasal tubercles that are sometimes fused, forming sinuous
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ridges. On the contrary, 15 rows of broad, thick scales cover most of the abdomen. These

broad scales are high, oval, and posteroanteriorly imbricated. Among these abdominal scale
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rows, the anterior one has a single scale, the posterior has two scales, and the middle rows
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have up to five scales. In the precloacal region of the trunk, most ventral scales are

rhomboidal and almost as high as long, while in the postcloacal region, these scales become

triangular and at least four times higher. The scales in one row become higher from ventral

to dorsal. Small tubercles randomly scattered or forming dorsoventral rows ornament the

lateral surfaces of most of these abdominal scales; however, in the larger scales, these

ornaments fused, forming short sinuous ridges.

The cloacal vestibule is deep and penetrates about one-fifth into the abdomen. Two high

acicular cloacal scales border this cavity, have acute ends, overlap dorsally, and are

ornamented poorly with tiny, scattered tubercles (Figures 8 and 10). The posterior cloacal
scale is nearly twice as high as the anterior one. Posterior of the pelvic fin, the anterior

cloacal scale joins the posterior precloacal scute and the posterior cloacal scale contacts the

anterior postcloacal scute.

Predorsal scutes— The predorsal region of the trunk, between the occiput and the dorsal

fin base, is bordered entirely by a continuous series of 11 predorsal scutes (dorsal ridge

scales in Poyato-Ariza and Wenz, 2002). These scutes are broad triangular-trapezoidal

shaped, with ventral wings evenly sized covering the upper third of the flanks of the trunk.

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The tips of these scutes are in contact. Those scutes located in the apex of both predorsal

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humps have wings that are larger than those of adjacent scutes (Figure 10). The largest

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scutes cover the apex of the posterior hump and are about four times higher and two and a
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half times longer than the smaller scutes placed in the predorsal central depression between
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the humps. All the predorsal scutes have a longitudinal medial row of 2-5 broad spines,
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which increase in size posteriorly. These spines are conical, sharp-tipped, broad-based,

curved backward, and placed side by side without leaving any gaps in between.
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The predorsal scutes 1 and 2 are trapezoidal, high, and subequal in size. The anterior one
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sutures the posterodorsal part of the skull, while the second is slightly higher. Both scutes

form the apex of the anterior predorsal hump and ventrally reach the pinniculus parietalis.

The scutes 3-5 are triangular, a little longer than high, project ventrally, and have

slightly concave dorsal edges; moreover, these are progressively larger in anteroposterior

order and border the central predorsal depression. Besides, the scutes 6-8 are high

rectangular, border the apex of the posterior predorsal hump, and have the largest ventral

wings in the predorsal scute series. Finally, the scutes 8-11 are triangular, higher than long,

become smaller in anterior to posterior order, and the last one reaches the anterior end of

the dorsal fin base.

 The ventral wings of the predorsal scutes show a dorsoventral thickening bar and

scattered small tubercles irregularly distributed or dorsoventrally aligned. A series of 2-5

spines form a longitudinal medial row of these scutes. These spines are conical, broad-

based, sharp-tipped, and slightly curved posteriorly; their sizes increase in the

anteroposterior order, and their bases are side by side with no empty spaces in between.

Although most scutes have three or four spines, their arrangement varies throughout the

series, and among the specimens studied, these spines occupy part of the posterior half of

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scutes 1-6, while in the apex of the posterior predorsal hump, the spines occupy the anterior

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half of the scutes 7-8; and further back, these tend to occupy the middle part of the most

posterior scutes.
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In the small specimen, IGM 14025, the predorsal humps are closer to each other, scute 6
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has a slightly convex anterior edge, and the spines of scutes 8-11 are nearly straight. On the
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other hand, in the large specimens, IGM 14024 and IGM 14026, the humps are farther

away, the scute 6 has a concave dorsal edge, and scutes 8-11 have spines curved anteriorly.
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Preanal scutes— Between the isthmus and the origin of the anal fin, the abdomen is
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entirely bordered with a series of 13-16 preanal scutes (ventral keel scutes in Poyato-Ariza

and Wenz, 2002), including 11-14 precloacals and two postcloacals (Table 1, Figures 2, 3,

8, and 10). These preanal scutes are triangular-trapezoidal shaped, barely expanded

dorsally, and have anterior and posterior acute ends. These scutes have contact with each

other through their anterior and posterior ends, forming a continuous series only interrupted

by the deep cloacal cavity.

Among the precloacal scutes, those in the middle of the abdomen are slightly larger and

have a longitudinal series of 5 ventral bordering spines. In contrast, those scutes placed

anterior, or posterior become smaller and have fewer spines, never less than two. The
precloacal scute ventral to the pelvic fin base is small and has only two spines. The spines

of these scutes are conical, sharp, broad-based, curved posteriorly, form a continuous series

without empty spaces, and extend along the ventral edge of the scute. Here, the spines show

increasing sizes posteriorly. Small and scattered tubercles ornament the laterodorsal wings

of the preanal scutes. There are two postcloacal scutes. The anterior one is slightly longer

than the posterior one, none is ventrally notched (the postclacal notch is not present), and

both show 4-5 spines like those of the anterior preanal scutes but occupy only the posterior

of
half of these scutes (Figures 8 and 10).

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DISCUSSION
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Tahnaichthys magnuserrata gen. and sp. nov. is a scarce species. The Tlayúa Quarry
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highlights for its fossil richness and abundance of pycnodontiform specimens. However,

after nearly 50 years of the activity of the Tlayúa Project in the UNAM, so far, only the
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three double-hump pycnodont specimens described here have been recovered. The
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specimens of the type series here referred were fruitlessly analyzed at the beginning of the

century by Dr. Shelton Pleasant Applegate of the UNAM (Applegate, 2001) and by Dr.

Lucio Paulo Crivano Machado of the UERJ (Universidade do Estado do Rio de Janeiro);

however, only until now as part of the Doctoral Research by the first author (SPO), it was

possible to integrate a complete description of this species.

As explained in this section, the finding of Tahnaichthys magnuserrata reveals that the

taxonomic and morphological diversities of the Pycnodontidae fishes are still partially

unknown. Today, the eclectic and vast quantity of data and fossil preservation in fishes of

this group deserves an accurate and in-depth analysis before its inclusion in a
comprehensive phylogenetic analysis to allow the generation of a robust hypothesis of the

relationships between these fishes and their natural classification. The achievement of such

a task is beyond the possibilities and goals of this study; therefore, here, we will limit

ourselves to demonstrating the uniqueness of T. magnuserrata and discussing its possible

inclusion within some of the nominal taxonomical hierarchies of this extraordinary fish

group.

Undoubtedly, Tahnaichthys magnuserrata has adequate characteristics to support its

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inclusion within the order Pycnodontiformes, the suborder Pycnodontoidei, superfamily

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Pycnodontoidea, and family Pycnodontidae, as they were defined previously by different

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authors (e.g., Nursall, 1996 a, b, 1999; Poyato-Ariza, 2005, 2020). First, a glance at the
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fossils of T. magnuserrata reveals the most outstanding features of the Pycnodontifomes
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bauplan; this is a rounded and high-bodied fish, strongly compressed laterally, with empty
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notochordal canal, a noticeable modification, and reduction of the opercular bones; and a

peculiar tripartite jaw occlusion between the vomer and the prearticular bones. In addition,
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this species shares the following diagnostic Pycnodontiformes features listed by Poyato-
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Ariza and Wenz (2002): The antorbital and ethmoid skull regions are hyperthrophied and

include a huge mesethmoid composed by two laminar parts, one medial and other

anterolateral, which are T-shaped in a coronal section; the frontals are curved and short; the

parasphenoid is toothless and its orbital-ethmoid section inflects downward; the frontal is

short and curved; the dermal supraoccipital is unpaired (dermo supraoccipital in this work);

the dorsal process of the premaxilla is long and superficial; the dentary is reduced to a

slender bone; there is no supramaxilla, subopercle, nor infraopercle; the opercular process

in the hyomandibula is absent or strongly reduced; the opercle is reduced; teeth on vomer

and prearticular arranged in longitudinal rows; the main vomerine tooth row has 10 or more
teeth; the dentary has four or less teeth; the maxilla is toothless and loosely attached; the

cleithrum is large and has a ventral palaform; ribs are alate (winged); and the predorsal and

preanal edge of the trunk have a continuous series of thick scutes.

Here, we conclude that Tahnaichthys magnuserrata belongs to Pycnodontoidei sensu

Poyato-Ariza and Wenz (2002) because it possesses the following diagnostic features of

this suborder. The opercular process of hyomandibula is absent; the gular region is

unossified; there are only two thin and separated branchiostegal rays; the vomerine and

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prearticular teeth are absent or reduced; the scales do not cover the whole body; the

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posterior part of the nude between the unpaired fins is nude [this is the peltate or clathrate

scale pattern described by Nursall (1996b)].

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Tahnaichthys magnuserrata gen. and sp. nov. is also identified as a member of the
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Pycnodontoidea, as defined by Payato-Ariza and Wenz (2002). This new Mexican species
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shares the following synapomorphies of this superfamily. The hyomandibula has a small to

enlarged dermalized region ornamented superficially and ventrally suturing the preopercle:
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there is no opercular process in the hyomandibula; anterior wing or sagittal flanges in the
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base of the neural and haemal spines. Additionally, this species shows other non-esclusive

Pycnodontoidea features, including the dentary has three or fewer teeth; the adjacent neural

and hemal arches are in contact; there are two or fewer urodermals; there are no nuchal

plates; the predorsal scutes are less than 18; the spines of dorsal scutes becomes larger in

anteroposterior order; the preanal scutes are less than seven; cloacal vestibule with anterior

and posterior modified scales not forming a mosaic. In this case, T. magnuserrata

contravenes this diagnosis of Pycnodontoidea; according to the previous records in this

superfamily, the premaxillary and dentary teeth are incisiform, and the predorsal trunk

region has no dorsal prominence or hump. On the contrary, the Mexican species has two
small tubercular teeth (Figure 6) on the dentary, and the double hump in the predorsal

region is its most outstanding feature.

Moreover, Tahnaichthys magnuserrata also belongs to the Pycnodontidae, as defined by

Poyato-Ariza and Wenz (2002). Visibly, this species exhibits the panniculus parietalis, the

only synapomorphy defining the family. Additionally, T. magnuserrata also shows most of

the features non-exclusive of Pycnodontidae, as the presence of an enlarged and

ornamented infraorbital behind the orbit, the cheek is naked because of the lack of

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suborbitals; the maxilla is toothless and oval-reniform; dentary has two-three teeth; the

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anterior ten or less neural spines are autogenous; bar-like scales cover the predorsal part of

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the abdominal regions of the trunk; and the preanal scutes with spines located side-to-side
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with no free space in between.
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After the phylogeny of Pycnodontiformes made by hand by Nursall (1996a, b), which
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exposes the monophyly of the family Pycnodontidae, the interrelations of this clade have

different configurations, including deep contradictions, in best-performed phylogenetic

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hypotheses later published (Figure 11). These differences are not a result of the natural,
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historical accumulation of data about the anatomy and taxonomical diversity of

pycnodontid fossils but of the arbitrary selection (and not always adequately explained) of

the species and characters that are used or excluded from these studies, as well as to the

different computer programs and search protocols implemented by the various authors. Not

counting Tahnaichthys magnuserrata, the review of the taxonomical diversity of

pycnodontid shows that this family contains 90 valid nominal species representing 42

genera (see Appendix.); curiously, almost half of these genera have not been included

formally in any of the hypotheses mentioned above, probably because many of these taxa

are represented only by fragmentary and isolated remains of the body, jaws, and teeth.
Despite this problematic scenario, here we explore the possible affinities of T.

magnuserrata by comparing the osteological attributes of this species with nine selected

characters included in the diagnosis of the three subfamilies named and phylogenetically

defined in the hypotheses mentioned above (Table 2).

Poyato-Ariza and Wenz (2002) supported the monophyly of Nursallinae, and recently,

without any conflict, Flagellipinna rhomboides Cawley and Kriwet, 2019, was added to

this subfamily (Taverne and Capasso, 2020; Poyato-Ariza 2020). Tahnaichthys

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magnuserrata and members of this subfamily share only three non-exclusive features

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(Table 2, Figure 11): the well ossification of the lateral part of the skull, the lack of free

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anterior dorsal pterygiophores; and the absence of ventral and dorsal process enclosing the
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pectoral notch of the cleithrum. Tahnaichthys magnuserrata is excluded from this
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subfamily because it does not have any of the Nursallinae synapomorphies (the posterior
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hypocordals are paddle-like and hypertrophied; and the arcocentra are expanded

surrounding the notochordal canal and have a complex to hypercomplex between adjacent
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arcocentra).
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Tahnaichthys magnuserrata and members of the subfamily Proscinetinae sensu Ebert

(2020) share seven features, including one putative synapomorphy (Table 2, Figure 11).

These features include: the scale pattern peltate [this is a putative synapomorphy of

Proscinetinae because in Thiollierepycnodus wagneri (Thiollière, 1852), the scale pattern is

more like to the pattern peltate-amplified, described above, in which, in addition to

covering the predorsal region, the bar-like scales extend further back, into the caudal region

and between the unpaired fins (see Ebert, 2020, figs. 2, 3, 6)]; the well ossification of the

lateral part of the skull; the pectoral notch in the cleithrum has no dorsal nor ventral

processes; there cloacal vestibule has no bifid scales; the posterior hypochordals are not
hypertrophied, the notochordal canal is not surrounded by the arcocentra; and the adjacent

arcocentra have simple contact [within Prosicnetinae, Neoproscinetes penalvai Silva-

Santos, 1970, show different states in the last two features because its posterior arcocentra

are expanded, surrounding the notochordal canal and having a slightly complex contact

between adjacent arcocentra (Figueiredo and Silva Santos, 1987; Wenz, 1991)]. Here, T.

magnuserrata is excluded from the subfamily Proscinetinae because it does not share the

most notable of its synapomorphies (the free anterior dorsal pterygiophores).

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Furthermore, the hypotheses proposed by Ebert (2016, 2020) rejected the naturalness of

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the subfamily Pycnodontinae; however, recently, Poyato-Ariza (2020) reaffirmed that this

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is a valid clade. Tahnaichthys magnuserrata and members of this subfamily share the
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following features (Table 2, Figure 11): The presence of the dermocranial fenestra; the lack
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of free anterior dorsal pterygiophores; the posterior hypochordals are not hypertrophied the
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notochordal not surrounded by the arcocentra, and the adjacent arcocentra have simple

contact. The last two features are shared only partially because, in Pycnodontinae, the
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arcocentra are variable; in some cases, these are somewhat expanded and surround part of
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the notochordal canal and develop slightly complex adjacent contacts. Tahnaichthys

magnuserrata is excluded from Pycnodontinae because it does not have either of its two

synapomorphies (the poorly ossification of the lateral part of the skull and the presence of

conspicuous processes in the ventral and dorsal ends of the pectoral botch of the cleithrum).

Although in the summary of the comparison between T. magnuserrata and the three

subfamilies Pycnodontidae (Table 2), this species and Pycnodontinae share the presence of

the dermocranial fenestra; this is not a real synapomorphy because such feature is present in

other pycnodontids [e.g., Tepexichthys aranguthyorum (Figure 12), Costapycnodus costae

(Heckel, 1856), and Akromystax tilmachiton Poyato-Ariza and Wenz, 2005 (also see
Applegate, 1992; Taverne et al., 2019)], and at least one pyncnodontine, Polazzodus

coronatus Poyato-Ariza, 2010, do not have this cranial fenestra.

The reduction in the number of premaxillary teeth to only one large one represents a

derived and highly specialized character that merits the review and comparison of the

pycnodontiformes that possess it. Up to now and in order of appearance in the fossil record,

these fish are Tahnaichthys magnuserrata from the Albian of Tlayúa, Mexico, described

here; Rhinopycnodus gabriellae Taverne and Capasso, 2013, from the Cenomanian deposits

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of Haqel, Lebanon; Polazzodus coronatus from the Turonian-early Senonian deposits of

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"Calcari di Aurisina, in Polazzo, Gorizia, Northeastern Italy; and Tergestinia sorbinii

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Capasso, 2000, from the Late Campanian-Early Paleocene of Lubornica Formation, Trieste,
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northern Italy (also see Taverne et al., 2019; Poyato-Ariza, 2020). Since the phylogenetic
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analyses show that these genera are not closely related, the single premaxillary tooth is
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undoubtedly a convergent character, acquired by these fish independently at least twice

(Figure 11). Poyato-Ariza (2020) recognized the last two of these species as part of the
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subfamily Pycnodontinae, and Ebert (2016) concluded that Rhinopycnodus is the sister taxa
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of the other pycnodontids. The dentary is unknown in Tergestinia, but in different genera,

the dentary has two teeth, which are incisiform and markedly unequal in Polazzodus (the

anterior one is three or four times longer than the posterior), incisiform and equally tiny in

Rhinopycnodus, and similarly small and tubercular in Tahnaichthys. There are five

vomerine tooth rows in all these fishes except Tahnaichthys, which has only three. The

snout is remarkably elongated in all fish except in Tergestinia, which only has a moderately

elongated snout. In Tahnaichthys, all the fins are rounded; on the contrary, the unpaired fins

are strip-like (uniformly low) in Tergestinia and falcate (low with a slight anterior high

lobe) in Rhinopycnodus and Polazzodus trip-like. In contrast, the caudal fin is stalked (with
two forked lobes) in Polazzodus and double emarginated in Tergestinia and

Rhinopycnodus. Finally, the caudal peduncle is relatively long and involves three neural

and haemal spines Polazzodus; however, this peduncle is short and involves only two of

such spines in Tahnaichthys and Tergestinia, while in Rhinopycnodus peduncle is non-

existent because there are no hemal and neural spines separating the odd fins from the

caudal fins.

The comparison between Tahnaichthys magnuserrata, its closest coeval neighbor, and

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the only previously described species of Tlayúa, Tepexichthys aranguthyorum (Figure 12),

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is mandatory. Tepexichthys is a diamond-shaped body fish without predorsal humps and the

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dorsal apex coinciding with the origin of the dorsal fin (Applegate, 1992); on the contrary,
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Tahnaichthys is an oblong-shaped and double-hump fish in which the posterior predorsal
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hump also represents the dorsal apex of the trunk and is in front of the dorsal fin and also.
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The snout of Tepexichthys is contrastingly short compared to the long snout of

Tahnaichthys. Besides, Tahnaichthys has rounded fins, whereas Tepexichthys has falcated
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unpaired fins and a stalked caudal fin. Tepexichthys has a supraorbital bone, but
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Tahnaichthys lacks this. Both have a scale pattern peltate, but in Tepexichthys, the scales

are smooth, whereas, in Tahnaichthys, all the scales are ornamented intensively with

tubercles scattered or aligned, forming short ventrodorsal ridges.

Regarding the preceding paragraphs, the nomination and the recognition of

Tahnaichthys magnuserrata as a new member of Pycnodontidae is fully justified. As noted,

this species cannot be placed in any of the subfamilies or genera of this clade; it also differs

from all pycnodontids because it is the only double-hump with strongly ornamented scales.

Besides, this is one of the four species with a single premaxillary tooth.
 CONCLUSIONS

Tahnaichthys magnuserrata is the second nominal species known from the Albian

deposits of the Tlayúa Quarry. Here, we reveal that this species exhibits adequate

osteological attributes to support its nomination and place it as a new member of

Pycnodontiformes, Pycnodontoide, and Pycnodontidae. The contrasting morphology of

Tahnaichthys shows that the taxonomic and morphological diversity of pycnodontiformes

of
in North America is vast and includes unexpected features, such as the double-hump

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condition, and common convergent features with species from other regions and

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temporalities, such as the reduction of the premaxillary teeth. The development of a
re
desirable comprehensive phylogenetic analysis of the family Pycnodontidae is a giant
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pending task that goes beyond the scope of this manuscript, which also requires the in-
na

depth review of all the taxa attributed to this family and the conciliation of the data

previously considered in the different hypotheses already proposed (Figure 11). The present
ur

manuscript is the first result of a PhD research project of the first author (SPO), which deals
Jo

with reviewing, describing, and revising the phylogenetic relationships of pycnodontiforms

of the Tlayúa Quarry.

Acknowledgments

We thank the Aranguthy family and their Tlayúa Quarry workers for their genuine interest

in paleontology, friendship, and daily effort. UNAM provided financial support for this

research through the project DGAPA-PAPIIT IN115223. IPN offered additional support

through Project 20240772. This work results from the Doctoral research of the first author
(SPO), who also received a doctoral grant from CONACYHT, Mexico. We are grateful for

the valuable contributions of Dr. Martin Ebert, three additional anonymous reviewers, and

Dr. Francisco Javier Vega Vera (JSAES editor), which significantly improved the scope

and content of this manuscript.

References.

Agassiz L., 1833. Recherches sur les poissons fossiles. Vol 2, part 1, Neuchâtel et Soleure,

of
Petitpierre.

ro
Agassiz L., 1843. Recherches sur les poissons fossiles. Vol 2, part 2, Neuchâtel et Soleure,

Petitpierre.
-p
re
Alvarado-Ortega J., 2005. Sistemática de los peces Ichthyodectiformes de la Cantera
lP

Tlayúa, Puebla, México. PhD Thesis, Universidad Nacional Autónoma de México,

na

Distrito Federal, México.

Alvarado-Ortega J., Barrientos-Lara J.I., Espinosa-Arrubarrena L., Melgarejo-Damián

ur

M.P., 2014. Late Jurassic marine vertebrates from Tlaxiaco, Oaxaca State, southern
Jo

Mexico. Palaeontologia Electronica, 17, 1, 24A. https://doi.org/10.26879/454.

Alvarado-Ortega J., Cantalice K.M., Martínez-Melo A., García-Barrera P., Than-Marchese

B.A., Díaz-Cruz J.A., Barrientos-Lara J.I., 2020. Tzimol, a Campanian marine

paleontological site of the Angostura Formation near Comitan, Chiapas, southeastern

Mexico. Cretaceous Research, 107, 104279.

https://doi.org/10.1016/j.cretres.2019.104279.

Alvarado-Ortega J., Cantalice Severiano K.M., Barrientos-Lara J.I., Díaz-Cruz J.A., Than-

Marchese B.A., 2019. The Huehuetla quarry, a Turonian deposit of marine vertebrates
 in the Sierra Norte of Puebla, central Mexico. Palaeontologia Electronica, 22, 1, 13A.

https://doi.org/10.26879/921.

Alvarado-Ortega J., Cuevas-García M., Cantalice K., 2018. The fossil fishes of the

archaeological site of Palenque, Chiapas, southeastern Mexico. Journal of

Archaeological Science: Reports, 17, 462-476.

https://doi.org/10.1016/j.jasrep.2017.11.029.

Alvarado-Ortega J., Cuevas-García M., Melgarejo-Damián M.P., Cantalice K.M., Alaniz-

of
Galvan A., Solano-Templos G., Than-Marchese B.A., 2015. Paleocene fishes from

ro
Palenque, Chiapas, southeastern Mexico. Palaeontologia Electronica, 18, 2, 39A.

https://doi.org/10.26879/536.
-p
re
Alvarado-Ortega J., Espinosa-Arrubarrena L., Blanco A., Vega F.J., Benammi M., Briggs
lP

D.E.G., 2007. Exceptional preservation of soft tissues in cretaceous fishes from Tlayúa
na

quarry, central Mexico. Palaios, 22 (6), 682-685.

https://doi.org/10.2110/palo.2006.p06-059r.
ur

Alvarado-Ortega J., K.A. González-Rodríguez, A. Blaco-Piñón, L. Espinosa-Arrubarrena,

Jo

E. Ovalles-Damián., Mesozoic Osteichthyans of Mexico, In Vega, F.J., Nyborg, T.G.,

Perrilliat, M.C., Montellano-Ballesteros, M., Cevallos-Ferriz, S.R.S., Quiroz-Barroso,

S.A. (Eds.), Studies on Mexican Paleontology. Topics on Geobiology 24. Springer,

The Netherlands. https://doi.org/10.1007/1-4020-3985-9_9. 2006. p. 169-207.

Applegate S.P., 1992. A new genus and species of pycnodont from the Cretaceous (Albian)

of central Mexico, Tepexi de Rodríguez, Puebla. Revista Mexicana de Ciencias

Geológicas, 10(2), 164-178.

Applegate S.P., 2001. The pycnodont fauna from the Lower Cretaceous of Tepexi de

Rodríguez (Puebla, Mexico), p. 3. In Tintori, A. (Ed.), Abstract Book, III International

 Meeting on Mesozoic Fishes. Systematics, Paleoenvironment and Biodiversity.

Serpiano, Monte San Giorgo, Italy.

Applegate S.P., Alvarado-Ortega J., García-Barrera P., Ovalles-Damián E., Espinosa-

Arrubarrena L., 2004. Análisis de la diversidad de peces Pycnodontiformes de México.

p.17. In Libro de resúmenes. IX Congreso Nacional de Paleontología. Sociedad

Mexicana de Paleontología, Tuxtla Gutiérrez.

Applegate S.P., Espinosa-Arrubarrena L., Alvarado-Ortega J., and Benammi M., Revision

of
of recent investigations in the Tlayúa Quarry, In Vega, F.J., Nyborg, T.G., Perrilliat,

ro
M.C., Montellano-Ballesteros, M., Cevallos-Ferriz, S.R.S., Quiroz-Barroso, S.A.

-p
(eds.), Studies on Mexican Paleontology. Topics on Geobiology 24. Springer, The
re
Netherlands. 2006. p. 275-304.
lP

Applegate S.P., Espinosa-Arrubarrena L., López-Neri P., 1984. Las Calizas litográficas de
na

la cantera Tlayua en Tepexi de Rodríguez, Puebla. Libreto-guía de la excursion a

Tepexi de Rodríguez, Estado de Puebla. 7a Convención Geológica Nacional.

ur

Applegate S.P., Gonzalez-Rodríguez K.A., and Alvarado-Ortega J., Fish fauna of the
Jo

Tlayua quarries. In Guide book of the Field Trips of the 60th Annual Meeting Society

of Vertebrate Paleontology Publicación especial. Universidad Autónoma del Estado de

Hidalgo, Avances en Investigación. 2000. p. 97–105

Arambourg C., 1954. Les poissons crétacés du Jebel Tselfat: Maroc: Service Géologique du

Maroc, Notes et Mémoires, 118, 1-188.

Benammi M., Alvarado-Ortega J., Urrutia-Fucugauchi J., 2006. Magnetostratigraphy of the

Lower Cretaceous strata in Tlayúa Quarry, Tepexi de Rodríguez, state of Puebla,

Mexico. Earth, Planets and Space, 58, 1295-1302.

Berg L.S., 1937. A classification of fish-like vertebrates. Izvestiya Akademii Nauk USSR,

Seriya Biologischeskaya, 4, 1277–1280.

Berg L.S., 1940. Classification of fishes, both recent and fossil. Труды Зоологического

института, 5, 2, 87-517 [Works of the Zoological Institute of the Academy of Sciences

of the University of U.R.S.S, or, Traveaux de l'Institut Zoologique de l'Académie des

Sciences de l'U.R.S.S]

Blanco-Piñón A., 2003. Peces fósiles de la Formación Agua Nueva (Turoniano) en el

of
Municipio de Vallecillo, Nuevo León, NE-México. PhD Thesis, Universidad

ro
Autónoma de Nuevo León, Linares, Nuevo León, México.

-p
Blot J., 1987. L’ordre des Pycnodontiformes. Studi e Ricerche sui Giacimenti Terziari di
re
Bolca, 5, 1–211.
lP

Cantú-Chapa A., 1987. Las amonitas del Albiano superior de Tepexi de Rodríguez, Puebla.
na

Revista de la Sociedad Mexicana de Paleontología, 1, 159-160.

https://doi.org/10.1186/BF03352624.
ur

Capasso L., 2000. Tergestinia sorbinii gen. nov., sp. nov. del Paleocene inferiore di
Jo

Trebiciano, Trieste (Pisces, Pycnodontiformes). Atti del Museo Civico di Storia Naturale

di Trieste, 48, 261-289.

Capasso L., Abi Saad P., Taverne L., 2009. Nursallia tethysensis sp. nov., a new pycnodont

fish (Neopterygii: † Halecostomi) from the Cenomanian of Lebanon. Bulletin de

l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre, 79, 11, 117-

136.

Cavin L., Garcia G., Valentin X., 2020. A minute freshwater pycnodont fish from the Late

Cretaceous of southern France: Palaeoecological implications. Cretaceous

Research, 106, 104242. https://doi.org/10.1016/j.cretres.2019.104242.

Cawley J.J., Kriwet J., 2018. A new pycnodont fish, Scalacurvichthys naishi gen. et sp.

nov., from the Late Cretaceous of Israel. Journal of Systematic Palaeontology, 16(8),

659-673.

Cawley J.J., Kriwet, J., 2019. A new genus and species of pycnodontid fish Flagellipinna

rhomboides gen. et sp. nov. (Neopterygii, Pycnodontiformes), from the Upper

Cretaceous (Cenomanian) of Lebanon, with notes on juvenile form and ecology:

Journal of Vertebrate Paleontology, 39, 2, e1614012.

of
https://doi.org/10.1080/02724634.2019.1614012.

ro
Cawley J.J., Kriwet J., 2024. The Fossil Record and Diversity of Pycnodontiform Fishes in

-p
Non-Marine Environments. Diversity, 16, 4, 225. https://doi.org/10.3390/d16040225
re
Cooper S.L., Martill D.M., 2020. A diverse assemblage of pycnodont fishes
lP

(Actinopterygii, Pycnodontiformes) from the mid-Cretaceous, continental Kem Kem

na

Group of south-east Morocco. Cretaceous Research, 112, 104456.

https://doi.org/10.1016/j.cretres.2020.104456.
ur

Cope E. D., 1887. Geology and palaeontology. Am. Nat. 1887:1014-1019.

Jo

Dunkle D.H., Maldonado-Koerdell M., 1953. Notes on some Mesozoic fossil fish remains

from Mexico. Journal of the Washington Academy of Sciences, 43(10), 311-317.

Ebert M., 2016. The Pycnodontidae (Actinopterygii) in the late Jurassic: 2) Turboscinetes

gen. nov. in the Solnhofen Archipelago (Germany) and Cerin (France). Archaeopteryx,

33, 12-53.

Ebert M., 2020. A new genus of Pycnodontidae (Actinopterygii) from the Upper Jurassic of

France and Germany, included in a phylogeny of Pycnodontiformes. Zoological

Journal of the Linnean Society, 188(2), 434-454.

https://doi.org/10.1093/zoolinnean/zlz087.
Espinosa-Arrubarrena L., Applegate, S.P., A possible model for the paleoecology of the

vertebrate bearing beds in the Tlayúa quarries, near Tepexi de Rodríguez, Puebla,

México. In: Arratia G, Viohl G (eds.), Mesozoic Fishes -Systematics and

Paleoecology. Verlag Dr. Friedrich Pfeil. München. 1996. p. 539-550.

Feldmann R.M., Vega F.J., Applegate S.P., Bishop G.A., 1998. Early Cretaceous

arthropods from the Tlayúa Formation at Tepexi de Rodríguez, Puebla,

Mexico. Journal of Paleontology, 72, 1, 79-90.

of
https://doi.org/10.1017/S0022336000024033.

ro
Figueiredo F.J., Silva S.R., 1987. Considerações taxinômicas dos picnodontidos da

-p
Formaçaõ Gramame (camada de Fostato), Pe. Anais X congresso brasileiro de
re
Paleontologia, Rio de Janeiro. p. 25-31.
lP

Galavíz-Hernández A.I., 2022. Primer reporte formal para la fauna fósil de peces
na

(Actinopterygii) de la Formación Agua Nueva (Cenomaniano-Turoniano) en la

localidad de Xilitla San Luis Potosí. Thesis, Universidad Autónoma de San Luis
ur

Potosí, México.
Jo

Giersch S., Frey E., Stinnesbeck W., Gonzalez Gonzalez A.H., Fossil fish assemblages of

northeastern Mexico: New evidence of Mid Cretaceous actinopterygian radiationIn:

Krempská, Z. (Ed.), 6th Meeting of the European Association of Vertebrate

Palaeontologists, Spiŝská Nová Ves, Slovak Republic, Volume of abstracts. 2008. p.

43–45.

Giersch S., 2014. Die Knochenfische der Oberkreidezeit in Nordostmexiko: Beschreibung,

Systematik, Vergesellschaftung, Paläoökologie und Paläobiogeographie. PhD thesis.

Ruprecht-Karl-University Heidelberg, Deutschland.

González-Rodríguez K., Applegate S.P., Espinosa-Arrubarrena L., 2004. A New World

macrosemiid (Pisces: Neopterygii—Halecostomi) from the Albian of Mexico. Journal

of Vertebrate Paleontology, 24, 2, 281-289. https://doi.org/10.1671/1862.

Heckel J., 1856. Beiträge zur Kenntnis der fossilien Fishe Österreichs. Denkschriften des

Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche

Klasse, 11, 187-274.

Herrera-Flores J.A., The Tlayúa Quarry: An Overview of a Notable Early Cretaceous

of
Fossil-Lagerstätte from Mexico. In: Guerrero-Arenas R., Jiménez-Hidalgo, E. (Eds.),

ro
Past Environments of Mexico. Springer, Switzerland. 2024. p. 389-404.

-p
Juárez-Arriaga E., Barragán R., Núñez-Useche F., Moreno-Bedmar J.A., 2023.
re
Sedimentary environments in the prelude to the lagerstätten conditions of the Tlayúa
lP

Formation (Albian) in central Mexico: A microfacies approach. Journal of South

na

American Earth Sciences, 131, 104650. https://doi.org/10.1016/j.jsames.2023.104650.

Kashiyama Y., Fastovsky D. E., Rutherford S., King J., & Montellano M., 2004. Genesis of
ur

a locality of exceptional fossil preservation: paleoenvironments of Tepexi de

Jo

Rodríguez (mid-Cretaceous, Puebla, Mexico). Cretaceous Research, 25(1), 153-177.

https://doi.org/10.1016/S0195-6671(03)00052-1.

Kriwet J., Poyato-Ariza F.J., Wenz S., 1999. A revision of the pycnodont fish Coelodus

subdiscus Wenz, 1989, from the Early Cretaceous of Montsec (Lleida, Spain). Treballs

del Museu de Geologia de Barcelona 8: 33-65.

Martill D.M., 1989. A new 'Solenhofen' in Mexico. Geology today, 5, 1, 25-28.

Martin-Abad H., Poyato-Ariza, F.J., 2013. Historical patterns of distribution in

Pycnodontiform and Amiiform fishes in the context of moving plates: Geologica

Belgica, 16, 4, 217-226. https://doi.org/10.1111/j.1365-2451.1989.tb00607.x.

Mülleried F.K., 1951. Algunos fósiles marinos del Terciario Inferior y Medio de Palenque,

Chiapas. Revista de la Sociedad Mexicana de Historia Natural, 12, 209-227.

https://doi.org/10.26879/536.

Nursall J.R., Distribution and ecology of pycnodont fishes. In: Arratia G., Viohl G. (Eds.),

Mesozoic Fishes – Systematics and Paleoecology. Verlag Dr. F. Pfeil, München.

1996b, p. 115-124.

Nursall J.R., The phylogeny of pycnodont fishes. In: Arratia, G., Viohl, G. (Eds), Mesozoic

of
fishes -systematics and paleoecology. Verlag Dr. F. Pfeil, München. 1996a. p. 125–152.

ro
Nursall J.R., The pycnodont bauplan: the morphology of a successful taxon. In: Arratia G,

-p
Schultze HP (Eds.), Mesozoic Fishes 2: Systematics and Fossil Record. Verlag Dr.
re
Friedrich Pfeil, München. 1999 p. 189-214
lP

Pantoja-Alor J., 1992. Geología y paleoambiente de la cantera Tlayúa, Tepexi de

na

Rodríguez, Estado de Puebla. Revista Mexicana de Ciencias Geológicas, 9, 2, 156-169.

Patterson C., 1973. Interrelationships of holosteans. In P. H. Greenwood, R. S. Miles, and

ur

C. Patter- son (eds.), Interrelationships of fishes, pp. 233-305. Academic Press,

Jo

London.

Poyato-Ariza F.J., Wenz, S., 2002. A new insight into pycnodontiform fishes.

Geodiversitas, 24, 1, 139-248.

Poyato-Ariza F.J., 2005. Pycnodont fishes: morphologic variation, ecomorphologic

plasticity, and a new interpretation of their evolutionary history. Bulletin of the

Kitakyushu Museum of Natural History and Human History, Series A (Natural

History), 3, 169-184. https://doi.org/10.34522/kmnh.3.0_169.

Poyato-Ariza F. J., 2010. Polazzodus, gen. nov., a new pycnodont fish from the Late

Cretaceous of northeastern Italy. Journal of Vertebrate Paleontology, 30(3), 650-664.

https://doi.org/10.1080/02724631003762955.

Poyato-Ariza F.J., 2020. Studies on pycnodont fishes (II): revision of the subfamily

Pycnodontinae, with special reference to Italian forms. Rivista italiana di Paleontologia

e Stratigrafia, 126, 2, 447-473. https://doi.org/10.13130/2039-4942/13514.

Poyato-Ariza F.J., Wenz S., 2005. Akromystax tilmachiton gen. et sp. nov., a new

of
pycnodontid fi sh from the Lebanese Late Cretaceous of Haqel and En Nammoura.

ro
Journal of Vertebrate Paleontology 25 (1): 27-45. https://doi.org/10.1671/0272-

4634(2005)025[0027:ATGESN]2.0.CO;2.
-p
re
Regan C. T., 1923. The skeleton of Lepidosteus, with remarks on the origin and evolution
lP

of the lower neopterygian fishes. Proceedings of the Zoological Society of London

na

1923:445–461.

Riquelme F., Ramos-Arias M., Aguilar-Franco M., Alvarado-Ortega J., Ruvalcaba-Sil J.L.,
ur

2021. Zircon U-Pb age of the Cretaceous Tlayúa Fossil-Lagerstätte in Central Mexico.
Jo

Neues Jahrbuch für Geologie und Paläontologie, 301, 2, 157-169.

https://doi.org/10.1127/njgpa/2021/1004.

Sapper K., 1899. Ueber gebirgsbau und Boden des noerdlichen Mittelamerika. Petermanns

Mitteilungen, Ergänzungsband, 127.

Seibertz E., Buitrón BE., 1987. Paleontología y estratigrafía de los Neohibolites del

Albiano de Tepexi de Rodríguez, Estado de Puebla (Cretácico Medio, México).

Sociedad Mexicana de Paleontología, 1, 285-299.

Silva Santos R.,1970. A paleoictiofauna da Formação Santana–Holostei: Família

Girodontidae. Anais da Academia Brasileira de Ciências, 42, 3, 445-452.

Solé J., Pi-Puig T., Bermúdez-Chávez C., Garduño-Martínez D., Alvarado-Ortega J., 2022.

Mineralogy, geochemistry, and KAr dating of feldspars and clays from an exceptional

Cretaceous fossil locality (Tlayúa, Puebla, Mexico): Insights into the depositional and

diagenetic ages and processes. Chemical Geology, 612, 121134.

https://doi.org/10.1016/j.chemgeo.2022.121134.

Stinnesbeck E.S., Rust J., Herder F., 2019. Paleobiology and taphonomy of the pycnodont

fish Nursallia gutturosum, based on material from the Latest-Cenomanian-middle

of
Turonian Vallecillo platy limestone, Mexico. PalZ, 93(4), 659-668.

ro
Taverne L., Capasso L., Del Re M., 2019. The pycnodont fishes from the Lower

-p
Cretaceous of the Capo d'Orlando, near Castellammare di Stabia (Naples, Campania,
re
southern Italy), with the description of the new genus Costapycnodus. Geo-Eco-Trop,
lP

43, 1: 53-74.
na

Taverne L., Capasso L., 2013. Osteology and relationships of Rhinopycnodus gabriellae

gen. et sp. nov. (Pycnodontiformes) from the marine Late Cretaceous of

ur

Lebanon. European Journal of Taxonomy, 67, 1-14.

Jo

http://dx.doi.org/10.5852/ejt.2013.67.

Taverne L., Capasso L., 2020. New data on the osteology and relationships of

Flagellipinna rhomboides, a pycnodont fish (Pycnodontidae) from the Cenomanian

(Upper Cretaceous) of Lebanon. Geo-Eco-Trop, 44(1), 175-184.

https://doi.org/10.1080/02724634.2019.1614012.

Taverne L., Capasso L., 2021. Osteology and phylogenetic relationships of Agoultpycnodus

aldrovandii gen. and sp. nov., Geo-Eco-Trop, 45(3), 487-495.

Thiollière V., 1852. Troisième notice sur les gisements à poisons fossiles situés dans le Jura

du département de l’Ain. Annales des Sciences Physiques et Naturelles, Lyon 2 : 354–

446.

Toombs H., Rixon A.E., 1959. The use of acids in the preparation of vertebrate fossils.

Curator, 2 (4), 304-312. https://doi.org/10.1111/j.2151-6952.1959.tb00514.x.

Reynoso V.H., Cabral-Perdomo M.A., Clark J.L., Espinosa-Arrubarrena M., Montellano-

Ballesteros, The reptiles of the Tlayúa Formation. In Guidebook of the Field Trips of

of
the 60th Annu. Meet. Society of Vertebrate Paleontology Publicación especial.

ro
Universidad Autónoma del Estado de Hidalgo, Avances en Investigación. 2000. p.

106-113.
-p
re
Wenz S., 1989. Iemanja palma 1989. In Maisey, J. (Ed.), 1991. Santana Fossils: an illustrated
lP

Atlas. TFH Publications, Inc., Neptune City, New Jersey p. 130-140.

na
ur
Jo
 FIGURE CAPTIONS

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Figure 1. Map of the Tlayúa Quarry.

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— Planned to one-column width or 90 mm —
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Figure 2. IGM 14024, holotype of Tahnaichthys magnuserrata gen. and sp. nov. from the
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Albian deposits of the Tlayúa Quarry, Puebla, Mexico. A) IGM 14024a, part of the
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specimen transferred to resin and observed under UV light. B) Idealized line drawing based

on A.

—Planned to one-page width or 190 mm —

Figure 3. Paratypes of Tahnaichthys magnuserrata gen. and sp. nov. from the Albian

deposits of the Tlayúa Quarry, Puebla, Mexico. A) Part of IGM 14025 under UV light. B)

IGM 14026 under UV light.

 —Planned to one-page width or 190 mm —

Figure 4. Head of Tahnaichthys magnuserrata gen and sp. nov. from the Albian deposits of

Tlayúa Quarry, Puebla, Mexico, based on Figure 2A. Abbreviations:

ang, angular; art, articulate; asph, autoesphenotic; br, branchiostegals; ach, anterior

of
ceratohyal; cl, cleithrum; dpt, dermopterotic; dsoc, dermosupraoccipital; dsph,

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dermoesphenotic; ecpt, ectopterygoid; enpt, entopterygoid; ext, extrascapular; fr, frontal; ft,

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temporal fenestra; hy, hyomandibular; dhyom, dermohyomandibula; io, infraorbitals; d,
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dentary; pch, posterior ceratohyal; pmx, premaxila; met, mesethmoid; mpt, metapterygoid;
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mx, maxilla; op, opercle; pa, parietal; pop, preopercle; pp, peniculus parietalis; pra,
na

prearticular; psp, parasphenoid; prfr, prefrontal; pt, posttemporal; q, quadrate; s, sclerotic;

scl, supracleithrum; sym, sympletic; vo, vomer.

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—Planned to one-page width or 190 mm —

Figure 5. Vomer bones of Tahnaichthys magnuserrata gen and sp. nov. paratype IGM

14025 from the Albian deposits of Tlayúa Quarry, Puebla, Mexico. Abbreviations: lpra, left

prearticular; rpra, right prearticular; s, sclerotic; vo, vomer.

— Planned to one-page width or 190 mm —

Figure 6. Teeth of Tahnaichthys magnuserrata gen. and sp. nov., from the Albian deposits

of Tlayúa Quarry, Puebla, Mexico. A) IGM 14026, close up of the jaw bones coated with

magnesium. B and C) IGM 14024a, close up prearticular teeth (B) and pharyngeal teeth (C)

coated with magnesium. Abbreviations: dt, dentary tooth; mx, maxilla; pmxt, premaxillary

of
tooth; prat, prearticular tooth; vt, vomer teeth.

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— Planned to one-column width or 90 mm —
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Figure 7. Pectoral girdle and fin of the IGM 14025, paratype of Tahnaichthys magnuserrata

gen. and sp. Abbreviations: cl, cleithrum; co, coracoid; pcfr, pectoral fin rays; pcr, pectoral
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fin radians; sc, scapula.

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— Planned to one-page width or 190 mm —

Figure 8. Pelvic find and cloacal scales of Tahnaichthys magnuserrata gen. and sp. nov.

from the Albian deposits of the Tlayúa Quarry, Puebla, Mexico. A) Close-up of the

paratype IGM 14025 under UV light. B) Close-up of the paratype IGM 14026 under UV

light. C) Idealized line draw based on A. Abbreviations: an, anal notch; afr, anal fin ray; cv,
vestibule cloacal; ps, preanal shields; pta, anal pterygophore; pvfr, pectoral fin rays; pclvs,

postcloacal ventral scutes; usc, unmodified scales.

— Planned to one-page width or 190 mm —

Figure 9. Caudal fin line draw based of Tahnaichthys magnuserrata gen. and sp. nov. from

the Albian lagersttäte of Tlayúa Quarry, Puebla, Mexico. Abbreviations: afr, caudal fin

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radius; apt, anal pterygiophore; dfr, dorsal fin ray; dpt, dorsal pterygiophore; e, epiaxial; h,

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hypoaxial; prr, procurrent caudal fin ray; ud, urodermal; black arrows show the upper and

lower principal caudal fin rays.

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Figure 10. Scale and scute pattern of Tahnaichthys magnuserrata gen. and sp. nov. based
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on the holotype IGM 14024. A) Details of the scale. B) Predorsal scutes. C) Preanal scutes.

D) Post-cloacal shields. Abbreviations: bls, bar-like scale; pclvs, postcloacal ventral scutes;

pds, predorsal shields; ps, preanal shields; pvfr, pectoral fin rays; usc, unmodified scales.

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Figure 11. Phylogenetic hypotheses of the subfamilies of Pycnodontidae.

 — Planned to one-page width or 190 mm —

Figure 12. IGM 3286 under UV light, holotype of Tepexichthys aranguthyorum

Applegate, 1992, from the Albian Tlayúa lagerstätte, Tepexi de Rodríguez Municipality,

of
Puebla, central Mexico.

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Table 1. Measurements and meristic data of the type specimens of Tahnaichthys

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magnuserrata gen. and sp. nov. erected here. Measurements are expressed in millimeters;

the proportions are % of the standard length (SL); ≈, +, imprecise data (not considered in
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the averages and modes).

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Table 2. Comparison of Tahnaichthys magnuserrata gen. and sp. nov. and the three

nominal subfamilies of the family Pycnodontidae.

Table 1. Measurements and meristic data of the type specimens of Tahnaichthys

magnuserrata gen. and sp. nov. erected here. Measurements are expressed in millimeters;

the proportions are % of the standard length (SL); ≈, +, imprecise data (not considered in

the averages and modes).

IGM 10003 IGM 20001 IGM 30002 Averages

Total length (TL) 165.2 ≈73.74 ≈84.08 –

Standard length (SL) 135.53 60.41 69.96 –
Head Length (HL) 68.79 31.2 35.02 45
HL as % of SL 50% 52% 50% 51%

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Head height (HH) 66 26.99 33.08 42.02
HH as % of SL 49% 45% 47% 49%

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Skull length (SKL) 64.31 ≈29.79 33.27 48.79
SKL as % of SL 47% ≈49% 48% 48%

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Skull height (SKH) 49.86 ≈17.1 23.57 36.71
SKH as % of SL 37% ≈28% 34% 35%
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Preorbital length (POL) 40.12 16.43 22.45 26.33
POL as % of SL 30% 27% 32% 30%
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Orbit diameter (OD) 13.6 5.33 6.18 8.37

OD as % of SL 10% 9% 9% 9%
Maximum body height (MBH) 98.81 42.66 45.67 62.38
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MBH as % of SL 66% 71% 65% 68%

Predorsal apex length (PDAL) 94.35 41.65 49.76 61.88
PDAL as % of SL 70% 69% 71% 70%
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Predorsal length (PDL) 111.33 48.62 54.94 72.29

PDL as % of SL 82% 80% 79% 80%
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Preanal length (PAL) 112.64 49.29 57.71 73.21

PAL as % of SL 83% 82% 82% 82%
Prepelvic length (PVL) 88.02 38.54 47.22 57.92
PVL as % of SL 65% 64% 67% 65%
Dorsal fin length (DFL) 23.44 11.02 13.96 16.14
DFL as % of SL 17% 18% 20% 18%
Anal fin length (AFL) 21.08 10.68 10.95 14.23
AFL as % of SL 16% 18% 16% 16%
Caudal peduncle length (CPL) 1.4 .47 0.8 0.89
CPL as % of SL 1 0.77 1.1 0.95
Modes
Total archocentra 23 23 23 23
Abdominal archocentra 13 13 13 13
Caudal archocentra 10 10 10 10
Ribs (pairs) 10 10 10 10
Epiaxials/Hypoaxials 8/11 8/11 5+/9+ 8/11
Pectoral fin rays 22+ 20+ 19+ –
Pelvic fin rays 5+ 5+ 4+ –
Dorsal fin rays 29 29 28 29

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Anal fin rays 23 22 22 22
Caudal formula iii+I+20+I+iii iii+I+20+I+iii ? iii+I+20+I+iii

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 Table 2. Comparison of Tahnaichthys magnuserrata gen. and sp. nov and the three

nominal subfamilies of the family Pycnodontidae.

Nursallinae Proscinetinae Tahnaichthys Pycnodontinae

sensu sensu magnuserrata Sensu
Poyato-Ariza and Ebert Poyato-Ariza
Wenz (2002) (2020) This work (2020)
1. Dermocranial fenestra No No Yes Yes
2. Scale pattern Imbricate, Peltate* Peltate Calthrate
Peltate-amplified
3. The lateral part of the Yes Yes Yes No, partially *

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skull is well-ossified ossified,
supraoccipital

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exposed
4. Free anterior dorsal No Yes * No No

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pterygiophores
5. Notochordal canal Slightly-totally * No No No, or
surrounded slightly
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6. Lateral contact Complex- * Simple Simple Simple or slightly
between archocentra hipercomplex complex
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7. Posterior hypocordals Yes, * No No No

hypertrophied paddle-like
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8. Processes in the No No No Yes, *

pectoral notch of the very large
cleithrum
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9. Cloacal bifid scale Yes No No Yes

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 HIGHLIGTHS

• Tlayúa Quarry is a Mexican marine Albian lagerstätte with a diverse fish

assemblage.

• The Tlayuan assemblage of pycnodontiform fish is the most diverse in North

America.

• Tahnaichthys magnuserrata is a peculiar doble-hump pycnodontiform from Tlayúa.

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• Tahnaichthys magnuserrata belongs to the family Pycnodontidae.

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• This fish exhibits a single large incisiform premaxillary tooth.

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Declaration of interests

☒ 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.

☐ The authors declare the following financial interests/personal relationships which may be considered
as potential competing interests:

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