See

discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/229730630

New Albian (Early Cretaceous) ophiuroids from

the Tlayúa Quarry, Puebla, Mexico. Palaeontol

Article in Palaeontology · December 2008

DOI: 10.1111/j.1475-4983.2008.00836.x

CITATIONS READS

4 35

3 authors, including:

Ben Thuy Pedro García-Barrera

Musée national d'histoire naturelle de Luxem… Universidad Nacional Autónoma de México
60 PUBLICATIONS 255 CITATIONS 33 PUBLICATIONS 187 CITATIONS

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Ophiuroid evolution and phylogeny View project

PaleoDeep: Paleobiology of the deep-sea through time and space View project

All content following this page was uploaded by Ben Thuy on 05 December 2014.

The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document
and are linked to publications on ResearchGate, letting you access and read them immediately.
[Palaeontology, Vol. 52, Part 1, 2009, pp. 83–94]

NEW ALBIAN (EARLY CRETACEOUS) OPHIUROIDS

FROM THE TLAYÚA QUARRY, PUEBLA, MEXICO
by LEONORA MARTIN-MEDRANO*, BEN THUY  and
PEDRO GARCÍA-BARRERA*
*Museo de Paleontologı́a, Departamento de Biologı́a Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México, Ciudad Universitaria, c.p. 04510,
D.F. México, México; e-mail: [email protected]
 Institut für Geowissenschaften, Universität Tübingen, Sigwartstraße 10, D-72076 Tübingen, Germany

Typescript received 20 February 2007; accepted in revised form 7 April 2008

Abstract: On the basis of six articulated individuals from morphologically. The rarity of ophiuroids, their complete
the middle to late Albian lithographic limestone at Tlayúa preservation as well as the unnatural contortions of their
near Tepexi de Rodrı́guez (Puebla, Mexico), a new species arms suggest these specimens to be allochthonous, having
of ophiuroid, Ophiactis applegatei, is described. The mate- been transported into a hostile environment where burial
rial adds to the rather poor record of Early Cretaceous was rapid.
brittlestars and represents the oldest known member of the
family Ophiactidae, being based on well-preserved and Key words: Albian, Tlayúa Formation, Mexico, Ophiuroi-
articulated specimens and thus reasonably well defined dea, taxonomy, palaeoecology.

Often referred to as the ‘Mexican Solnhofen’, the Tlayúa terms of individuals. The present paper, in which a new
Quarry near Tepexi de Rodrı́guez (State of Puebla, Mexico) species, Ophiactis applegatei, is erected, constitutes the
ranks amongst the more important fossil localities in the first formal description of fossil ophiuroids from Mexico.
New World. Strata exposed there have yielded a vast array Similar to other biota from Tlayúa, these brittlestars
of biota, which are remarkable in view of their diversity are well preserved and reveal hardly any sign of disinte-
and exceptional preservation, inclusive soft tissues in fish gration. As such, they offer a unique insight into skeletal
(Alvarado-Ortega et al. 2007). Over the past few decades, structure and the species’ palaeobiology. This is of special
the highly fossiliferous lithographic limestone at Tlayúa has note, because little is known about Early Cretaceous brit-
been the focus of studies which aim to characterize the tlestars in comparison to the well-documented Late Juras-
biota and reconstruct their palaeoecology. To date, only 5 sic and Late Cretaceous ophiuroid diversity (Hess 1975;
per cent of all biota represented at Tlayúa have been Kutscher and Jagt 2000). Here, we assess the present
described, which means that a majority (>6500 specimens) faunule taxonomically and briefly discuss its taphonomy
remain largely unassessed taxonomically. The presence of and palaeoecology.
brittlestar remains has been noted previously by Applegate
and Espinosa-Arrubarrena (1982), Applegate (1987, 1996),
Buitrón and Malpica-Cruz (1987), Feldmann et al. (1998), GEOLOGICAL AND
Applegate et al. (2000), Martin-Medrano (2003, 2006), PALAEONTOLOGICAL CONTEXT
Alvarado-Ortega (2005), Applegate et al. (2006) and
Martin-Medrano and Garcı́a-Barrera (2006). The Albian Tlayúa Formation, introduced by Pantoja-Alor
In Mexico, fossil ophiuroids are rare in comparison to (1992), comprises a succession of carbonates in the Tlayúa
other echinoderms and echinoids in particular (Nieto- Ravine area, near the city of Tepexi de Rodrı́guez (Text-
López and Garcı́a-Barrera 2006). To date, brittlestar fig. 1), which is subdivided into three members. About
remains have been recorded from strata of Late Carbonif- 50 m of massive white limestone constitutes the Lower
erous (Pennsylvanian) to Pliocene age (Buitrón et al. Member which yields miliolid foraminifera and small-sized
1994; Quiroz-Barroso and Sour-Tovar 1995), at just a molluscs; the occurrence of the rudist Toucasia polygyra
handful (seven in all) of localities. Amongst these, the Alencáster, 1973 and the bivalve genus Chondrodonta Stan-
Cretaceous accounts for three sites, namely the Tlayúa ton, 1901 dates this member as early Albian (Pantoja-Alor,
Quarry (Puebla; the present record), Redoma in the State 1992). The Middle Member (thickness c. 35 m; Text-fig. 2)
of Coahuila and Altares in Chihuahua (Martin-Medrano comprises mainly micritic, yellowish brown lithographic
2006), with the first-named being the richest locality in limestone with intercalated, hematite-rich clayey layers,

ª The Palaeontological Association doi: 10.1111/j.1475-4983.2008.00836.x 83

84 PALAEONTOLOGY, VOLUME 52

T E X T - F I G . 1 . Location of the Tlayúa

Quarry, near Tepexi de Rodrı́guez,
Puebla.

parallel to bedding, which outcrop over a stretch of some levels. Similar to all macrofossils from the Middle Member,
700 m (Pantoja-Alor 1992; Applegate et al. 2006). these brittlestars are preserved within thin clayey deposits
Magnetostratigraphic data strongly suggest a middle to late on bedding planes. We used water and soft brushes to
Albian age for these highly fossiliferous limestones remove the adhering clay from the specimens; subse-
(Benammi et al. 2006); this age assignment is supported by quently, immersion in a solution of 3 per cent acetic acid
the presence of the ammonite genera Hysteroceras Hyatt, for 5 min proved necessary to get rid of more consolidated
1900, Mortoniceras Meek, 1876 and cf. Anisoceras Pictet, carbonate encrustations, although this procedure slightly
1854 (Cantú-Chapa 1987; Applegate et al. 2006). Finally, damaged the ophiuroid skeletal microstructure.
the Upper Member (40 m thick) comprises grey dolomitic Five specimens were carbon coated for SEM observa-
limestones dated as Cenomanian on the basis of the tion, while the sixth, which is preserved on a slab together
miliolid foraminiferan Dicyclina schlumbergeri Munier- with an aspidorhynchid fish, was left uncoated. In the
Chalmas, 1887 (Fernández-Becerra 1985). descriptions, the use of morphological terms follows
The ophiuroid material described herein originates Stöhr (2005), while higher-level classification is adopted
from the Middle Member which is also referred to by the from Smith et al. (1995).
informal name ‘Tlayúa Quarry’, and it is these levels that
have produced numerous algae, plants, foraminifera, Institutional abbreviations. FCMP, Museum of Palaeontology,
sponges, cnidarians, molluscs, insects, spiders, crabs, Faculty of Sciences, Universidad Nacional Autónoma de México
echinoderms, reptiles and osteichthyan fish. The last- (UNAM), Mexico City; IGM, National Collection of the Institute
named group is represented best amongst macrofossil taxa of Geology, Universidad Nacional Autónoma de México
(UNAM), Mexico City.
and accounts for 80 per cent in current collections (>6500
specimens) from Tlayúa Quarry (Applegate et al. 2006).

SYSTEMATIC PALAEONTOLOGY
MATERIAL AND METHODS
Order OPHIURIDA Müller and Troschel, 1840
Suborder OPHIURINA Müller and Troschel, 1840
The faunule studied comprises six articulated specimens,
Family OPHIACTIDAE Matsumoto, 1915
all preserved on slabs of lithographic limestone. Seven addi-
tional specimens are known, but these are not considered
Genus OPHIACTIS Lütken, 1856
further because of insufficient preservation. Unfortunately,
the exact provenance of the specimens studied within the
Type species. Ophiactis krebsii Lütken, 1856 (= Ophiolepis sav-
quarry has not been noted and consequently, we cannot
ignyi Müller and Troschel, 1842), by subsequent designation of
determine whether this lot represent isolated, scattered H. L. Clark (1915).
occurrences or was found concentrated at one or more
 MARTIN-MEDRANO ET AL.: NEW ALBIAN (EARLY CRETACEOUS) OPHIUROIDS 85

T E X T - F I G . 2 . Generalized
lithostratigraphic section of the Tlayúa
Ravine (modified after Martin-Medrano
2006).

Ophiactis applegatei sp. nov. Locality and horizon. Tlayúa Quarry (Puebla, Mexico); litho-
Plates 1–4 graphic limestone of the Middle Member of the Tlayúa Forma-
tion of middle to late Albian age.
Derivation of name. In honour of Dr Shelton Pleasants Apple-
gate (1928–2005), for his indefatigable enthusiasm and promo- Diagnosis. Five-armed ophiactid with indented disc; radial
tion of palaeontology in Mexico and for the great vision and shields contiguous, except distally; short, blunt and
creativity that brought him to lead the Tlayúa Quarry research slightly flattened spines at least in ventral interradial areas
project from 1981 until his death. close to disc edge; one tiny plus one large operculiform
scale covering second tentacle pore in angle of mouth slit;
Types. Holotype is IGM 9298; paratypes are IGM 9290 and three to four flat, elongate oral papillae; single tentacle
FCMP 602.
scale on all pores, up to four tapering and erect spines
not exceeding the length of an arm segment.
Other material. Three additional specimens, preserved on slabs;
IGM 9296 (ventral side exposed; disc diameter 4.97 mm), IGM
Description of holotype. Disc round to subpentagonal with
9299 (ventral side exposed; disc diameter 4.78 mm, and IGM
indented interradial areas; disc diameter 5.2 mm. Dorsal side of
9301 (ventral side exposed, disc diameter 4.07 mm).
86 PALAEONTOLOGY, VOLUME 52

disc not exposed; ventral side with small, rounded plates, bear- uous, separated close to their distal tip by a gap forming an
ing short, blunt and flattened spines at least on disc margin (Pl. acute angle between radial shields. The gap is occupied by at
1, figs 1–2); no bursal slit observable. Oral shield approximately least one plate, sunken probably as a result of preservation, con-
as long as broad, arrow shaped, with slightly obtuse proximal sistent with the dorsal arm plates of the first segments outside
angle and rounded convex distal edge. Adoral shields abutting in the disc.
front of oral shield, not extending around lateral angles of oral Dorsal arm plates rather thick, fan shaped, with convex distal
shield or hardly so (Pl. 2, figs 1–2). Apical papilla very large, edge and slightly concave proximal edges forming an obtuse
heart shaped and flat (Pl. 2, figs 1–3). Second tentacle pore angle. Adjacent dorsal arm plates contiguous in proximal arm
opens within the mouth slit and bears a tiny distal scale and a segments (Pl. 4, fig. 5) separated by lateral arm plates in distal
large, operculiform proximal one encompassed by the rounded arm segments.
pentagonal first ventral arm plate and the angle formed by the Specimen FCMP 602 exposes the ventral disc, and generally
oral plate and adoral shield (Pl. 2, figs 1–2). Three to four flat, fits with the description of the holotype well; however, a few
elongate oral papillae occur on the lateral edges of the oral details are better preserved and provide supplementary morpho-
plates, increasing in size distally. Five arms, more or less com- logical data. Disc indented, scales of interradial areas with small,
plete and comprising >60 segments, total length of the longest blunt and slightly flattened spines, smaller than spines on the
arm as preserved 26.1 mm. Tentacle pores developed throughout edge of the interradial area. Larger spines of disc edge slightly
the arm (Pl. 2, figs 4–5), with one flat, rounded tentacle scale on extending into interradial notch. Bursal slit visible, abutting oral
all pores. Ventral arm plate rather thick, widest distally and gen- frame but not reaching edge of disc; no genital papillae discern-
erally longer than wide, except in proximalmost arm segments; ible. Three flat and blunt oral papillae on oral plate, flanked by
with convex distal edge, strongly concave lateral indentations for large round scale of second oral tentacle pore. Five arms, more
the tentacle pore, and narrower and straight proximal edge (Pl. or less complete (Pl. 3, fig. 3), maximum arm length (as pre-
2, figs 4–5). Adjacent ventral arm plates are contiguous at least served) is 17.5 mm. Ventral arm plates contiguous throughout
in proximal arm segments. Lateral arm plate of stout aspect, the arm. Tentacle pores present on all arm segments.
with distinct notch for tentacle pore. One slightly displaced
median lateral arm plate, higher than wide, shows coarse outer Discussion. Early Cretaceous ophiuroid taxa described to
surface, a slight constriction parallel to the strongly concave date clearly differ from O. applegatei sp. nov., in particu-
proximal edge of the plate, and large spine articulations in shal- lar with regard to disc appendages and plating of the oral
low notches of the distal edge of the plate, with two parallel hor-
frame. Ophiura graysonensis (Alexander, 1931; lower
izontal articulation ridges (Pl. 2, figs 4–5). Three to four, thick
Cenomanian of Texas), Ophiura texana (W. B. Clark,
and tapering arm spines, coarsely striated, widest at their base,
pointed, erect, not exceeding the length of one arm segment (Pl. 1893; upper Albian of Texas), Ophiolancea swartkopensis
2, figs 4–5). Some spines, especially the ventralmost, seem to be Shone, 1986 (Valanginian of South Africa), Ophiopeza
slightly flattened, but it cannot be ruled out though that this is a buehleri Hess, 1970 (upper Hauterivian of Switzerland)
preservational artifact. and Xanthamphiura hauteriviensis Hess, 1970 (upper
Two of the arms present a sudden and more or less obvious Hauterivian of Switzerland) have different mouth plating
change in arm segment size, suggesting that arm tips were in the and lack spines on the ventral side of the disc. The poorly
process of regeneration (Pl. 3, fig. 1). The two completely pre- known Ophiura? straini Cornell et al., 1991 (Albian of
served arms show a distinct curvature in their median portion New Mexico) has clearly separated radial shields, while
which clearly exceeds the maximum horizontal coiling capacity Geocoma libanotica König, 1825 (Cenomanian of Leba-
of the arms.
non) is not clearly defined (Hess 1960; Kutscher 1997).
Of other ophiuroid species from mid- and Upper
Paratype supplements and variation. Specimen IGM 9290 exposes
Mesozoic deposits, of which discs are known, hardly any
the dorsal disc and attribution to O. applegatei sp. nov. is based
on similarities in disc indentation and arm structure (Pl. 3, fig.
bears a closer resemblance to O. applegatei sp. nov. with
2). Recrystallization tends to blur details of disc plating; disc respect to mouth plating, disc indentation and append-
subpentagonal, slightly indented. No spines or granules are pres- ages. In addition, very few of these have been attributed
ent on disc, their absence presumably being due to insufficient to the Ophiactidae. Hess (1965) described isolated arm
preservation. The radial shield is of stout aspect, length about a ossicles and arm segments from the lower Oxfordian of
quarter of the disc diameter, proximal tip covered by small Switzerland as Ophiopholis? trispinosa; this species has
imbricating disc scales (Pl. 4, fig. 6). Paired radial shields contig- only up to three arm spines and, at least on median arm

EXPLANATION OF PLATE 1
Figs 1–2. Ophiactis applegatei sp. nov. from the middle–late Albian Tlayúa Formation, Tlayúa Quarry, Puebla (Mexico). 1, Drawing of
ventral disc and proximal arm segments. 2, FCMP 602 (paratype; SEM), ventral disc and proximal arm segments.
Abbreviations: LAP, lateral arm plates; OS, oral shield; S, arm spines; SED, spines on edge of disc; TS, tentacle scales; VAP, ventral
arm plates.
 PLATE 1

SED

OS

LAP VAP
TS
1

OS

500 µm
2

MARTIN-MEDRANO et al., Ophiactis applegatei

88 PALAEONTOLOGY, VOLUME 52

segments, ventrally abutting lateral arm plates. Ophiactis? PALAEOECOLOGY AND TAPHONOMY
sulcatus Kutscher and Jagt (in Jagt 2000) (lower Maas-
trichtian of Rügen, and possibly upper Campanian of Palaeoecological models published for these lithographic
north-east Belgium) is based on isolated lateral arm plates limestone and its exceptionally well-preserved biota
which had up to five arm spines; median lateral arm favour one of two interpretations: deposition took place
plates are elongated. Ophiopholis sp. 1 of Ishida (2004, p. in an open-marine basin under storm-dominated condi-
37), from the Miocene of Japan, has larger oral shields. tions and restricted bottom water circulation (Kashiyama
Thus, the erection of a new species for the Mexican mate- et al. 2004), or it occurred in a shallow-water, protected
rial appears warranted. back reef lagoon (Applegate 1987). The most convincing
Attribution to the Ophiactidae is based mainly on the piece of evidence comes from well-documented cyanobac-
presence of disc spinelets, a single, cordiform apical terial mats, which imply that water depths did not exceed
papilla, short erect spines and conspicuous radial shields. 60 m (Applegate et al. 2006). In their evaluation of the
There is a certain affinity with members of the Ophiacan- palaeoecology of different biotic groups encountered in
thidae, but the arm spines in the new species are atypi- the Tlayúa lithographic limestone, Applegate et al. (2006)
cally short and the lateral arm plates rather massive and highlighted the considerable percentage of allochthonous
not as strongly constricted as in most ophiacanthids. Sim- forms, originating from either reefal environments, the
ilarities to the Amphiuridae are only superficial; the Mex- open sea or nearby terrestrial settings, with life inside the
ican specimens lack the typical paired infradental papillae lagoon assumed to have been restricted to nektonic
of amphiurids. organisms and occasional occurrences of benthic scaveng-
According to Paterson (1985, p. 76), the Recent Oph- ers (Espinosa-Arrubarrena and Applegate 1996; Guerrero-
iactis abyssicola (Sars, 1861) has a round or indented Arenas 2004). There is ample evidence of stagnant
disc, spinelets of variable distribution on interradial disc conditions inside the lagoon, which led to hypersaline
areas, a cordiform apical papilla and a single tentacle and anoxic conditions (Applegate et al. 2006). Miliolids
scale. As far as number and arrangement of oral papil- (Trocholina lenticularis Henson, 1948, and the genus
lae are concerned, there are greater similarities to the Quinqueloculina d’Orbigny, 1826; (C. Rosales-Domı́nguez,
extant ophiactid Histampica duplicata (Lyman, 1875) pers. comm. 2006) are found at some levels in Tlayúa
(see Paterson 1985, fig. 32), even though other charac- Quarry (Alvarado-Ortega et al. 2007), like on slab FCMP
teristics (i.e., number of tentacle scales and disc 602 and corroborate hypersalinity.
appendages) clearly differ. Ophiopholis has dorsal arm All ophiuroids studied show hardly any sign of decay;
plates surrounded by small plates and thus clearly dif- only the arm tips of IGM 9299 (Pl. 4, fig. 2) show slight
fers from our specimens. The closest affinities are disintegration. Despite their small size, the dislocated arm
shared with five-armed species of the genus Ophiactis, plates were not subjected to transport and this substanti-
which justifies the attribution to this genus. However, ates stagnant conditions at the time of decay, but the role
as long as dorsal disc plating (and possible appendages) of possible cyanobacterial mats as sediment traps prevent-
and internal disc structure (especially shape of ambula- ing the plates from further transport must also be consid-
cral part of the oral plate) remain unknown, this ered. Burial of these specimens was probably fairly rapid
assignment is tentative. (Allison 1991; Ausich 2001), as the distalmost segments
Ophiactis applegatei sp. nov. is the oldest known of an ophiuroid arm are amongst the first skeletal parts
member of the Ophiactidae, and being based on to disintegrate (BT, pers. obs.). Most specimens reveal
well-preserved, articulated specimens, it is reasonably conspicuous contortions of the arms; the tips and median
well-defined. The question of the origin of hexamery, parts of the arms are often unnaturally curled. In IGM
frequently encountered in Ophiactis, remains unan- 9298, the distal halves of two arms are broken but still
swered, as all individuals of the new species have five adhere to the proximal half (Pl. 3, fig. 1). It is unclear
arms. whether these contortions and injuries are because of

EXPLANATION OF PLATE 2
Figs 1–5. Ophiactis applegatei sp. nov. from the middle–late Albian Tlayúa Quarry, Puebla (Mexico). IGM 9298 (holotype). 1,
Drawing of fig. 2, showing jaw and proximal arm plates. 2, Jaw and proximal arm plates (SEM). 3, Mouth frame (SEM). 4,
Proximalmost arm segments (SEM). 5, median arm segments (SEM).
Abbreviations: AO, adoral shield; AP, apical papilla; DP, dental plate; LAP, lateral arm plates; OP, oral papillae; OS, oral shield;
S, arm spines; S*, arm spine on oral shield, not drawn as its position is probably an artifact of preservation; TS, tentacle scale;
VAP1, first ventral arm plate; VAP2, second ventral arm plate.
 PLATE 2

2 3

4 5

MARTIN-MEDRANO et al., Ophiactis applegatei

90 PALAEONTOLOGY, VOLUME 52

unfavourable conditions shortly prior to the animals’ Guzmán-Mendoza (Instituto de Investigaciones en Materiales,
death or rather constitute a post mortem effect caused by UNAM) and S. Espinosa-Matı́as (Facultad de Ciencias, UNAM)
hypersalinity and comparable to contortions seen in fish for advice and technical assistance during SEM sessions; G. Gon-
and reported on by Applegate et al. (2006) for Tlayúa zález-Mancera (Facultad de Quı́mica, UNAM) for technical sup-
port in carbon coating; H. Hernández-Campos (professional
and by Viohl (1990) for Solnhofen. One specimen (IGM
photographer) for photographs; F. Solı́s-Marı́n (Instituto de
9296) (Pl. 4, fig. 1) displays an arrangement of the arms
Ciencias del Mar y Limnologı́a, UNAM) for assistance given
comparable to the posture of a moving ophiuroid using during review of the National Collection of Echinoderms; G.
one leading arm (Ishida and Fujita 2001). Another speci- Hendler (Museum of Natural History of Los Angeles County,
men (IGM 9301) has its ventral side exposed and the Los Angeles) for advice given and for allowing access to collec-
arms partly flexed on the dorsal side in a way comparable tions of extant brittlestars in his care, D. Navarro-Santillán and
to the arm-coiling posture mentioned by Emson and Wil- I. Nieto-López (Facultad de Ciencias, UNAM) for technical sup-
kie (1981) (Pl. 4, figs 3–4). Up to now, a total number of port.
thirteen ophiuroid specimens are known from the Tlayúa
Quarry. Other echinoderm classes are represented by few
specimens only. As the lithographic limestone are being REFERENCES
intensely explored for fossils under the supervision of the
Geological Institute of UNAM, the paucity of echinoderm A L E N C Á S T E R , G. 1973. Una nueva especie de Toucasia en el
remains cannot be explained by a lack of collecting. It Cretácico medio de los Estados de Oaxaca y Puebla. Universi-
seems more probable that the environment was not dad Nacional Autónoma de México, Instituto de Geologı́a, Pale-
ontologı́a Mexicana, 36, 4–20.
favourable for echinoderms, which implies that their
A L E X A N D E R , C. I. 1931. A new Lower Cretaceous ophiuroid.
remains, inclusive of the brittlestars described herein,
Journal of Paleontology, 5, 152–153.
must all be allochtonous, having been transported from A L L I S O N , P. A. 1991. Variation in rates of decay and disartic-
reefal or open marine settings into the hostile deposi- ulation of Echinodermata: implications for the application of
tional environment and mixed with autochthonous, actualistic data. Palaios, 5, 432–440.
open-marine, reefal, freshwater and terrestrial organisms. A L V A R A D O - O R T E G A , J. 2005. Sistemática de los peces Ich-
Recent species of Ophiactis are often associated with coral thyodectiformes de la Cantera Tlayúa, Puebla, México. Unpub-
(S. Stöhr, pers. comm. 2008). Thus, it is quite possible lished PhD thesis, Facultad de Ciencias, Universidad Nacional
that these specimens were transported into the lagoon Autónoma de México, México, 302 pp.
after their coral hosts had been broken up during a —— E S P I N O S A - A R R U B A R R E N A , L., B L A N C O , A.,
storm, because remains of gorgonians have been recorded V E G A , F. J., B E N A M M I , M. and B R I G G S , D. E. G. 2007.
Exceptional preservation of soft tissues in cretaceous fishes
from the Tlayúa lithographic limestones (Applegate et al.
from the Tlayúa Quarry, Central México. Palaios, 22, 682–685.
2006). Whether the ophiuroids reached their final desti-
A P P L E G A T E , S. P. 1987. A preliminary study of the Tlayúa
nation dead or alive is uncertain; the possible moving Quarry near Tepexi de Rodrı́guez, Puebla. Sociedad Mexicana
posture of specimen IGM 9296 (Pl. 4, fig. 1) suggests that de Paleontologı́a, Revista, 1, 40–50.
at least some individuals were still alive. As the exact —— 1996. An overview of the Cretaceous fishes of the quarries
provenance of the specimens remains unknown, it cannot near Tepexi de Rodrı́guez, Puebla, México. 529–538. In
be determined whether all originate from a single or from A R R A T I A , G. and V I O H L , G. (eds). Mesozoic fishes. Sys-
several such transportation events. The mode of occur- tematics and paleoecology. F. Pfeil, München, 576 pp.
rence of these ophiuroids fits the model of a back reef —— and E S P I N O S A - A R R U B A R R E N A , L. 1982. Litho-
lagoon with at least temporarily stagnant water and hy- graphic limestone-like deposits in Tepexi de Rodrı́guez,
persaline conditions well. Puebla, México. Society of Vertebrate Paleontology, Universidad
Nacional Autónoma de México, Instituto de Geologı́a, Field Trip
Guidebook, 42nd Annual Meeting, 39 pp.
Acknowledgements. We thank J. W. M. Jagt (Natuurhistorisch
—— G O N Z A L E Z - R O D R Í G U E Z , K. and A L V A R A D O -
Museum Maastricht [SCZ]) and S. Stöhr (Naturhistoriska ri-
O R T E G A , J. 2000. Fish fauna of the Tlayúa quarries. 97–
ksmuseet, Stockholm) for their valuable comments and thorough
130. In E S P I N O S A - A R R U B A R R E N A , L., M O N T E L -
reviews of the manuscript; M. C. Perrilliat-Montoya (Instituto
L A N O - B A L L E S T E R O S , M. and A P P L E G A T E , S. P.
de Geologı́a, UNAM) for the loan of material; M. Piña-Barba, J.

EXPLANATION OF PLATE 3
Figs 1–3. Ophiactis applegatei sp. nov. from the middle–late Albian Tlayúa Formation, Tlayúa Quarry, Puebla (Mexico). 1, IGM 9298
(holotype), ventral surface; the regenerating arm tip is marked by a thick arrow; the two arms broken in the distal half are
marked by thin arrows. 2, IGM 9290 (paratype), dorsal surface. 3, FCMP 602 (paratype), ventral surface. Scale bar represents
2 mm.
 PLATE 3

MARTIN-MEDRANO et al., Ophiactis applegatei

92 PALAEONTOLOGY, VOLUME 52

(eds). Paleontological highlights of the Mixteca Poblana in Cen- bearing beds in the Tlayúa quarries, near Tepexi de Rodrı́guez,
tral Mexico. Society of Vertebrate Paleontology, Universidad Puebla, México. 539–550. In A R R A T I A , G. and V I O H L , G.
Nacional Autónoma de México and Universidad Autónoma (eds). Mesozoic fishes. Systematics and paleoecology. F. Pfeil,
del Estado de Hidalgo, Field Trip Guidebook, 60th Annual München, 576 pp.
Meeting. Universidad Autónoma del Estado de Hidalgo, Pub- F E L D M A N N , R. M., V E G A , F., A P P L E G A T E , S. P. and
licatión Especial, 172 pp. B I S H O P , G. 1998. Early Cretaceous arthropods from the
—— E S P I N O S A - A R R U B A R R E N A , L., A L V A R A D O - Tlayúa Formation at Tepexi de Rodrı́guez, Puebla, México.
O R T E G A , J. and B E N A M M I , M. 2006. Revision of recent Journal of Paleontology, 72, 79–90.
investigations in the Tlayúa Quarry. 275–304. In V E G A , F. J., F E R N Á N D E Z - B E C E R R A , S. 1985. Levantamiento a detalle
N Y B O R G , T. G., P E R R I L L I A T , M. C., M O N T E L L A N O - de columnas estratigráficas en el área de Tepexi de Rodrı́guez,
B A L L E S T E R O S , M., C E V A L L O S - F E R R I Z , S. R. S. and Puebla-‘‘Cantera Tlayúa’’. Petróleos Mexicanos, Superinten-
Q U I R O Z - B A R R O S O , S. A. (eds). Studies on Mexican dencia General de Exploración, Zona Centro, Distrito Poza
paleontology, 24. Springer, Berlin, 308 pp. Rica, 31 pp.
A U S I C H , W. I.. 2001. Echinoderm taphonomy. 171–227. In G U E R R E R O - A R E N A S , R. 2004. Icnofósiles de invertebrados
J A N G O U X , M. and L A W R E N C E , J. M. (eds). Echinoderm de la Formación Tlayúa en Tepexi de Rodrı́guez, Puebla, Méxi-
studies, 6. A. A. Balkema, Rotterdam, 348 pp. co. Unpublished MS thesis, Facultad de Ciencias, Universidad
BENAMMI, M., ALVARADO-ORTEGA, J. and Nacional Autónoma de México, México, 49 pp.
U R R U T I A - F U C U G A U C H I , J. 2006. Magnetostratigraphy H E N S O N , F. R. S. 1948. Larger imperforate foraminifera of
of the Lower Cretaceous strata in Tlayúa Quarry, Tepexi de south-western Asia. Families Lituolidae, Orbitolinidae and
Rodrı́guez, State of Puebla, México. Earth Planets Space, 58, Meandropsinidae. British Museum (Natural History), London,
1295–1302. 127 pp.
B U I T R Ó N , B. E. and M A L P I C A - C R U Z , R. 1987. Tepexi de H E S S , H. 1960. Neubeschreibung von Geocoma elegans (Ophiu-
Rodrı́guez, Puebla, una localidad fosilı́fera de México. Sociedad roidea) aus dem unteren Callovien von La Voulte-sur-Rhône
Mexicana de Paleontologı́a, Field Trip Guidebook, Distrito (Ardèche). Eclogae geologicae Helvetiae, 53, 335–385.
Federal, México, 24 pp. —— 1965. Mikropaläontologische Untersuchungen an Ophiuren.
—— S O L Í S M A R I N , F. A., M I R A N D A , J. C. and IV. Die Ophiuren aus dem Renggeri-Ton (Unter-Oxford) von
M I R A N D A , J. B. 1994. El hallazgo de un Ofiuroidea Pliocénico Chapois (Jura) und Longecombe (Ain). Eclogae geologicae
de la Región de Vizcaı́no, Baja California Sur. Memorias de la XII Helvetiae, 58, 1059–1082.
Convención Geológica Nacional, Sociedad Geológica Mexicana, —— 1970. Schlangensterne und Seesterne aus dem oberen Hau-
México, D.F., 26–27. terivien «Pierre jaune» von St-Blaise bei Neuchâtel. Eclogae
C A N T Ú - C H A P A , A. 1987. Las amonitas del Albiano Superior geologicae Helvetiae, 63, 1069–1091.
de Tepexi de Rodrı́guez, Puebla. Sociedad Mexicana de Paleon- —— 1975. Die fossilen Echinodermen des Schweizer Juras.
tologı́a, Revista, 1, 159–160. Veröffentlichungen aus dem Naturhistorischen Museum Basel, 8,
C L A R K , H. L. 1915. Catalogue of Recent ophiurans: based on 1–130, pls 1–48.
the collection of the Museum of Comparative Zoology. Mem- H Y A T T , A. 1900. Cephalopoda. 502–604. In V O N Z I T T E L ,
oirs of the Museum of comparative Zoölogy, Harvard College, K. A. (ed). Textbook of palaeontology. Macmillan, London ⁄
25, 163–376. New York, 890 pp.
C L A R K , W. B. 1893. The Mesozoic Echinodermata of the Uni- I S H I D A , Y. 2004. Ophiuroids. The Association for the Geologi-
ted States. Bulletin of the United States Geological Survey, 97, cal Collaboration in Japan, Monograph, 51, 1–80. [in Japanese,
1–207, pls 1–50. English summary].
C O R N E L L , W. C., L e M O N E , D. V. and N O R L A N D , W. D. —— and F U J I T A , T. 2001. Escape behavior of epibenthic
1991. Albian ophiuroids from Cerro de Cristo Rey, Dona Ana ophiuroids buried in the sediment: observations of extant
County, New Mexico. Journal of Paleontology, 65, 1009–1012. and fossil Ophiura sarsii sarsii. 285–292. In B A R K E R , M.
E M S O N , R. H. and W I L K I E , I. C. 1981. The arm coiling (ed.). Echinoderms 2000. A.A. Balkema Publishers, Lisse, 590
response of Amphipholis squamata (Delle Chiaje). 11–18. In pp.
L A W R E N C E , J. (ed.). Echinoderms: Proceedings of the Inter- J A G T , J. W. M. 2000. Late Cretaceous-Early Palaeogene echino-
national Conference. A. A. Balkema, Rotterdam, 234 pp. derms and the K ⁄ T boundary in the southeast Netherlands
E S P I N O S A - A R R U B A R R E N A , L. and A P P L E G A T E , S. and northeast Belgium – Part 3. Ophiuroids; with a chapter
1996. A possible model for the paleoecology of the vertebrate on: Early Maastrichtian ophiuroids from Rügen (northeast

EXPLANATION OF PLATE 4
Figs 1–6. Ophiactis applegatei sp. nov. from the middle–late Albian Tlayúa Formation, Tlayúa Quarry, Puebla (Mexico). 1, IGM 9296,
ventral surface. 2, IGM 9299, ventral surface. 3–4, IGM 9301 (part and counterpart), ventral surface. Scale bar represents 2 mm.
5, IGM 9290 (paratype; SEM), median arm segments. 6, IGM 9290 (paratype; SEM), radial shields and proximalmost arm
segments.
Abbreviations: DAP, dorsal arm plates; LAP, lateral arm plates; P, plate between radial shields; R, radial shields.
 PLATE 4

1 2

3 4

5 6

MARTIN-MEDRANO et al., Ophiactis applegatei

94 PALAEONTOLOGY, VOLUME 52

Germany) and Møn (Denmark) by Kutscher and J. W. M. —— 1842. System der Asteriden. F. Vieweg & Sohn, Braun-
Jagt. Scripta Geologica, 121, 1–179. schweig, xx + 134 pp.
K A S H I Y A M A , Y., F A S T O V S K Y , D. E., R U T H E F O R D , M U N I E R - C H A L M A S , E. 1887. Sur la Cyclolina et trois nou-
S., K I N G , J. and M O N T E L L A N O , M. 2004. Genesis of a veaux genres de foraminifères de couches à Rudistes. Cyclopsi-
locality of exceptional fossil preservation: paleoenvironments na, Dicyclina et Spirocyclina. Complete Rendus des Séances,
of Tepexi de Rodrı́guez (mid-Cretaceous, Puebla, México). Société Géologique de France, 1887, xxx–xxxi.
Cretaceous Research, 25, 153–177. N I E T O - L Ó P E Z , I. and G A R C Í A - B A R R E R A , P. 2006. Cre-
K Ö N I G , C. 1825. Icones Fossilium Sectiles. The author, London, taceous echinoids of Mexico. 101–114. In V E G A , F. J., N Y -
44 pp. B O R G , T. G., P E R R I L L I A T , M. C., M O N T E L L A N O -
K U T S C H E R , M. 1997. Bemerkungen zu den Plattenkalk- B A L L E S T E R O S , M., C E V A L L O S - F E R R I Z , S. R. S. and
Ophiuren, insbesondere Geocoma carinata (v. Münster, 1826). Q U I R O Z - B A R R O S O , S. A. (eds). Studies on Mexican pale-
Archaeopteryx, 15, 1–10. ontology, 24. Springer, Berlin, 308 pp.
—— and J A G T , J. W. M. 2000. Early Maastrichtian ophiuroids O R B I G N Y , A. D. d’. 1826. Tableau méthodique de la classe
from Rügen (northeast Germany) and Møn (Denmark). 45– des Céphalopodes. Annales des Sciences naturelles, 7, 245–314.
107, pls 22–36. In J A G T , J. W. M. (ed.). Late Cretaceous– P A N T O J A - A L O R , J. 1992. Geologı́a y paleoambiente de la
Early Palaeogene echinoderms and the K ⁄ T boundary in the cantera Tlayúa, Tepexi de Rodrı́guez, Estado de Puebla. Revi-
southeast Netherlands and northeast Belgium- Part 3: Ophiu- sta del Instituto de Geologı́a, Universidad Nacional Autónoma
roids. Scripta Geologica, 121, 1–179, 36 pls. de México, 9, 156–169.
L Ü T K E N , F. C. 1856. Bidrag til Kundskab om Slangestjern- P A T E R S O N , G. L. J. 1985. The deep-sea Ophiuroidea of the
erne. III. Bidrag til Kundskab om Ophiurerne ved Central- North Atlantic Ocean. Bulletin of the British Museum of Natu-
amerikas Vestkyst. Videnskabelige Meddelelser fra det ral History (Zoology), 49, 1–162.
Naturhistoriske Forenings i Kjobenhavn, 1856, 20–26. P I C T E T , F. J. 1854. Traité de paléontologie, Céphalopodes, 2,
L Y M A N , T. 1875. Zoological results of the Hassler Expedition. 583–716. J.-B. Baillière, Paris, 896 pp.
II. Ophiuridae and Astrophytidae. Illustrated Catalogue of the Q U I R O Z - B A R R O S O , S. A. and S O U R - T O V A R , F. 1995.
Museum of Comparative Zoology, 8, 1–34. Nuevo registro de Ophiuroidea (Ophiurinidae) para el Pensi-
M A R T I N - M E D R A N O , L. 2003. El registro de ofiuroideos lvánico de América del Norte, proveniente de la Formación
fósiles de México y la descripción de dos nuevas formas para el Ixtaltepec, Oaxaca. Memorias del V Congreso Nacional de Pale-
Cretácico y Terciario. Unpublished Bachelor¢s thesis, Facultad ontologı́a, Sociedad Mexicana de Paleontologı́a, México, D.F.,
de Ciencias, Universidad Nacional Autónoma de México, 31.
México, 55 pp. S A R S , M. 1861. Oversigt of Norges Echinodermer. Christiania, vi
—— 2006. Análisis paleontológico de los ofiuroideos fósiles de Méx- + 160 pp.
ico. Unpublished MSc thesis, Facultad de Ciencias, Universi- S H O N E , R. W. 1986. A new ophiuroid from the Sundays River
dad Nacional Autónoma de México, México, 140 pp. Formation (Lower Cretaceous), South Africa. Journal of Pale-
—— and G A R C Í A - B A R R E R A , P. 2006. Fossil ophiuroids of ontology, 60, 904–910.
Mexico. 115–131. In V E G A , F. J., N Y B O R G , T. G., P E R - S M I T H , A. B., P A T E R S O N , G. L. J. and L A F A Y , B. 1995.
R I L L I A T , M. C., M O N T E L L A N O - B A L L E S T E R O S , Ophiuroid phylogeny and higher taxonomy: morphological,
M., C E V A L L O S - F E R R I Z , S. R. S. and Q U I R O Z - molecular and palaeontological perspectives. Zoological Journal
B A R R O S O , S. A. (eds). Studies on Mexican paleontology, 24. of the Linnean Society, 114, 213–243.
Springer, Berlin, 308 pp. S T A N T O N , T. W. 1901. Chondrodonta, a new genus of ostrei-
M A T S U M O T O , H. 1915. A new classification of the Ophiu- form mollusks from the Cretaceous, with descriptions of the
roidea: with descriptions of new genera and species. Proceed- genotype and a new species. Proceedings of the United States
ings of the Academy of Natural Sciences of Philadelphia, 67, Natural History Museum, 24 (1257), 301–307.
43–92. S T Ö H R , S. 2005. Who’s who among baby brittle stars (Echino-
M E E K , F. B. 1876. A report of the invertebrate Cretaceous and dermata, Ophiuroidea): postmetamorphic development of
Tertiary fossils of the upper Missouri country. In H A Y D E N , some North Atlantic forms. Zoological Journal of the Linnean
F. V. (ed.). Report of the United States Geological Survey of the Society, 143, 543–576.
Territories, 9, i–xvi + 629 pp. V I O H L , G. 1990. Solnhofen lithographic limestone. 285–289.
M Ü L L E R , J. and T R O S C H E L , F. H. 1840. Uber die In B R I G G S , D. E. G. and C R O W T H E R , P. R. (eds). Palae-
Gattungen der Ophiuren. Archiv für Naturgeschichte, 6, 327– obiology, a synthesis. Blackwell Scientific Publications, Oxford,
330. 583 pp.

View publication stats