Cretaceous Research 106 (2020) 104247

Contents lists available at ScienceDirect

Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

Short communication

First Plesiosaurian record (Diapsida; Sauropterygia) from the La

Herradura formation, (ValanginianeHauterivian), Morro Solar, Peru
Iva
lez a, *, Jose

n Meza-Ve
P. O'Gorman b
a
Departamento de Paleontología de Vertebrados, Museo de Historia Natural e UNMSM, Lima, Peru
b
n Paleontología Vertebrados, Museo de La Plata, Universidad Nacional de La Plata, Paseo del Bosque s/n., B1900FWA, La Plata, Argentina-CONICET:
Divisio
Consejo Nacional de Investigaciones Científicas y T

ecnicas, Argentina

a r t i c l e i n f o a b s t r a c t

Article history: We report the first plesiosaur remains (Diapsida, Sauropterygia) from Peru, recovered from Lower
Received 2 May 2019 Cretaceous rocks cropping out in Morro Solar, within Lima city (La Herradura Formation, Valanginian
Received in revised form eHauterivian). The material is restricted to an undetermined propodium, fragmentary vertebrae and
12 August 2019
ilium, being identified as a Plesiosauria, with putative affinities to Leptocleididae or Elasmosauridae. This
Accepted in revised form 9 September 2019
Available online 16 September 2019
record provides further data on the group in the Southeastern Pacific.
© 2019 Elsevier Ltd. All rights reserved.
Keywords:
Plesiosauria
Lower Cretaceous
La Herradura formation
Morro Solar
Peru

1. Introduction
ramo-Fonseca, 2015; Pa

Plesiosaurians are a monophyletic group of Mesozoic diapsid
reptiles adapted to aquatic life (Benson and Druckenmiller, 2014).
Plesiosaurians achieved a cosmopolitan distribution, being recor-
ded on all continents including Antarctica (Welles, 1952; Gasparini
et al., 1984; O'Keefe, 2004a; Carpenter, 1999; O'Gorman, 2012;
Otero et al., 2014; Bardet et al., 2014; Brown, 1981; Benson and
Druckenmiller, 2014). This clade is characterized by using the
anterior and posterior limbs in locomotion (O'Keefe and Carrano,
2005). Plesiosaurians appeared towards the end of the Triassic
and diversified during the Jurassic and the Cretaceous until the K/
Pg mass extinction (Benson and Druckenmiller, 2014). The clade
Plesiosauria developed two morphotypes (plesiosauromorph and
pliosauromorph; O’Keefe, 2002). The greatest number of specimens
and diversity of Cretaceous plesiosaurs come from North America
(Welles, 1943; Bakker, 1993; Carpenter, 1999; O'Keefe, 2004b, 2008;
Sato, 2003; Bardet et al., 2014; Druckenmiller and Russell, 2008).
The record of South American plesiosaurs has increased in the last
decades; they have been recorded in Colombia (Hampe, 2005;
* Corresponding author. Av. Arenales 1256, Jesús María, Lima 11, Peru.
E-mail addresses: imv2999@gmail.com (I. Meza-Ve
lez), joseogorman@fcnym.
unlp.edu.ar (J.P. O'Gorman).
Pa
ramo-Fonseca et al., 2019), Venezuela
(Colbert, 1949), Chile (Otero et al., 2015a) and Argentina (Gasparini
and Spalletti, 1990; Gasparini and De La Fuente, 2000; Gasparini
and Salgado, 2000; Gasparini et al., 2003; O'Gorman, 2016).
Despite this recent increase in the record of South American
Cretaceous plesiosaurs, the plesiosaurian record from Peru is
almost null. Given the few fossil records of Mesozoic vertebrates in
Peru and the scarcity of records of cretaceous plesiosaurs in the
South Pacific (Otero et al., 2015a, b), the Morro Solar plesiosaur
record will add valuable biogeographic information of the Creta-
ceous in Peru and marine reptiles cretaceous of the South American
Pacific. In 2000, Manuel Rojas Aquije and Iva
n Meza-Ve
lez collected
material referable to Plesiosauria indet. (MUSM 3735) in levels of
the La Herradura Formation, Morro Solar locality, Peru (Fig. 1). The
specimen was donated by the latter to the collection of the
Department of Vertebrate Paleontology of the Natural History
Museum of the UNMSM. In this contribution MUSM 3735 is
described and its possible affinities are discussed.
Anatomical abbreviation: cap, capitulum; cr, caudal rib; tub/tro,
tuberosity/trochanter.
Institutional abbreviations: GPMM, Geomuseum der Universita €t
Münster, Westfalen, Germany; KUVP, University pf Kansas Natural
History Museum and Biodiversity Research Center; UNMSM, Uni-
versidad Nacional Mayor de San Marcos; MUSM, Museo de Historia

https://doi.org/10.1016/j.cretres.2019.104247
0195-6671/© 2019 Elsevier Ltd. All rights reserved.
2 I. Meza-V
elez, J.P. O'Gorman / Cretaceous Research 106 (2020) 104247
Fig. 1. Locality where the MUSM 3735 specimen was collected. A, geological map (modified from Ferna
3735 was collected.
Natural de la Universidad Nacional Mayor de San Marcos, Lima,
Peru; RA and VL, paleontological collections of the Museo Geo-

gico Jose
lo
Royo y Go

mez, Servicio Geolo
gico Colombiano; SAM,
South Australian Museum, Adelaide.
2. Geological setting
The Morro Solar promontory is a topographic height of 281 m
and occupies an area of 7.48 km2. Morro Solar is located south of the
Chorrillos locality, Lima Province, Peru (Fig. 1). The sedimentary
sequence of Morro Solar comprises 450 m of quartz-rich sands,
shales and sandstones of ValanginianeHauterivian age (Ferna
ndez-
Concha, 1958). The original Morro Solar-San Lorenzo Formation
described by Lisson (1907) was later re-redefined as a group by
Ferna
ndez-Concha (1958). Following the latter author The Morro
Solar Group can be subdivided into a basal Salto del Fraile Forma-
tion; in the mid-section, the shales and sandstones of the La Her-
radura Formation, where the material described in the present
contribution was collected; finally the uppermost Marcavilca For-
mation (Ferna
ndez-Concha, 1958).
Following Ferna
ndez-Concha (1958), the La Herradura Forma-
tion, which crops out in the central part of Morro Solar, is subdivided
into two members: the basal La Virgen Member and the upper La
Herradura Member (Rosenzweig, 1953; Ferna
ndez-Concha, 1958).
The better exposed outcrops of La Herradura Formation are located
in the sea coast and along the road that connects Chorrillos with La
Herradura Beach (Nún ~ ez del Prado and Cha
vez, 1989). The base and
top of this formation are transitional with the Salto Del Fraile and
Marcavilca formations, respectively (Aleman et al., 2006). The La

ndez-Concha, 1958). B, general map; C, photo of the locality where MUSM
Virgen Member, where MUSM 3735 was collected, consists of 80 m
of grayish brown to reddish brown oxidized shale, with subordinate
layers of siltstones, fine-grained sandstones and gypsum near the
base; the shale is laminated and contains abundant calcareous
nodules and ferruginous concretions; the siltstones are sandy,
including calcareous thin layers, and lack any internal structures.
Layers of sandstone are present near the base and are continuous for
a great distance (Aleman et al., 2006).
The La Herradura Member, which takes its name from the local
beach, comprises 114 m of sandstones and shales; the sandstones
tend to increase in thickness towards the top, coinciding with the
presence of a massive cross-bedding (Aleman et al., 2006). The
lower 49 m have fine to medium-grained gray-brown sandstones
interbedded with laminated shales, while the upper part consists of
65 m of laminated shales and siltstones in thin strata (Aleman et al.,
2006).
The La Virgen Member includes the ammonoids Raimondiceras
raimondii, Raimondiceras pfluckeri, Lissonia riveroi, Favrella lorensis
and Argentiniceras pardoi (Romero et al., 1995; Aleman et al., 2006)
and the bivalves Buchotrigonia paradisensis, Buchotrigonia lorenti
and Anditrigonia lorenti (Romero et al., 1995; Aleman et al., 2006).
Romero et al. (1995) reports these same taxa for La Herradura
Member. There is no previous record of marine reptiles in any
formation of the Morro Solar Group.
3. Material and methods
Because the remains are found in together in the same strati-
graphic and therefore interpreted as part of a single individual. The
 I. Meza-V
elez, J.P. O'Gorman / Cretaceous Research 106 (2020) 104247 3

Fig. 2. MUSM 3735, Plesiosauria indet. A, caudal vertebrae in articular view; B, C, interminata vertebrae in articular view; D, ilium in posterior? view; E-G, propodium in E, dorsal, F,
proximal and G, ventral, views. Scale bar ¼ 20 mm.

material was collected partially clean, especially the propodium were taken using a digital caliper (error 0.5 mm) directly on the
and the ilium, they were prepared with pneumatic equipment and specimens. Photos were taken using a Canon digital camera SX 130
consolidating substances (e.g. Paraloid and B-72). Measurements IS and direct edition on Photoshop CS 5.
4 I. Meza-V
elez, J.P. O'Gorman / Cretaceous Research 106 (2020) 104247

Fig. 3. Comparison of outlines of several propodial elements. A, MUSM 3735; B, C (VLI 17052004-1, Stenorhynchosaurus munozi Pa
ramo-Fonseca, Go
mez-Pe
rez, Noe

, Etayo-Serna,
2016), B, right humerus in dorsal view and C, left femur in dorsal view. D, E (SAM P 15980, Leptocleidus sp.), D, left humerus in dorsal view and E, left femur in dorsal view. F, G
(GPMM A3 B4, Brancasaurus brancai Wegner, 1914), F, right humerus in dorsal view and G, right femur in dorsal view. H, I (RA 4425, Callawayasaurus colombiensis (Welles)
Carpenter, 1999), H, left humerus in dorsal view and I, left femur in dorsal view, J (KUVP 40002, Dolicorhynchops bonneri (Adams, 1997) O'Keefe, 2008) right humerus in dorsal view.
Scaled at equal shaft wide. (Redrawn from Welles, 1962; Kear, 2007; O'Keefe, 2008; Sachs et al., 2016; Pa
ramo-Fonseca et al., 2016).

4. Systematic paleontology
Sauropterygia Owen, 1860
Plesiosauria de Blainville, (1835)
Plesiosauria indet.
(Fig. 2)
Materials. MUSM 3735. A probable caudal vertebra, two uniden-
tified vertebrae; a partially preserved ilium, a propodium (Fig. 2).
Locality and horizon. Morro Solar, district of Chorrillos, Lima, Peru.
La Herradura Formation, La Virgen Member. Lower Cretaceous
ValanginianeHauterivian (Ferna
ndez-Concha, 1958, Fig. 1)
5. Description
Only three vertebrae are preserved. A single caudal vertebra could
be identified; it preserves half of the vertebral centrum. Both artic-
ular faces are slightly amphycoelic. The lateral surface preserves a
prominent facet compatible with a parapophysis strongly projected
laterally towards the center of the vertebral body. The presence of a
(Fig. 2A), a single and circular rib facet located on the lateral surface
of vertebral central allows to identify the vertebra as caudal as this
morphology is shared by caudals of elasmosaurids (Welles, 1943);
leptocleidids (Druckenmiller and Russell, 2006; Sachs et al., 2016).
The ventral foramina are nor observed as the ventral surface is
damage. Two other fragmentary vertebrae are preserved; both are
damaged. Most anatomical features are lost, however, it can be noted
that both articular facets are slightly concave.
Only a half ilium is preserved and part of the proximal end is
eroded, making it difficult to determine its laterality. The piece is
slender and the medial part of the body is curved (Fig. 2D). The
propodium preserves the tuberosity/trochanter, capitulum, the
diaphysis and part of the distal expansion (Fig. 2E-G). The preserved
part of the shaft is 380 mm in length (chapter and column), with a
column 90 mm at neck level. The capitulum is strongly convex and
asymmetric and is covered by transphyseal foramina. The cross
section of the shaft is sub-elliptical and no marked muscular scar is
present on the ventral surface. The capitulum and tuberosity are
mostly confluent. The expanded distal end is only partially pre-
served; it was impossible to determine the original expansion.
dorsal trochanter and distal expansion (Rhodin, 1985; Benson and
Druckenmiller, 2014). The incompleteness of the specimen pre-
cludes determining whether it is a humerus or a femur. The ques-
tion about the familial affinities of this material is complex due to
the incompleteness of the specimen. In order to determine possible
affinities of MUSM 3735, the propodial was compared to other
plesiosaurians from the Late Cretaceous. The propodium of bra-
chauchenine pliosaurids have relatively longer and gracile shafts
(see Hampe, 2005: fig. 8, but see Pa
ramo-Fonseca et al., 2016 for an
alternative identification of VL17052004-1). Also, among the lep-
tocleidids, the outline of the propodial of Leptocleidus sp. differs in
the almost straight anterior margin (Kear, 2007, Fig. 3). However,
other specimens referred to Leptocleidus show a more concave
anterior margin (Cruickshank and Long, 1997: Figs. 14, 15). The
leptocleidid Brancasaurus brancai shows an outline more compat-
ible with MUSM 3735 (Sachs et al., 2016). Although elasmosaurids
propodial outlines are variable in shape (Welles, 1952, 1962; Sato,
2003; Lomax and Wahl, 2013; O'Gorman; et al., 2015; Kubo et al.,
2012) additionally some polycotylids shows humerus with out-
lines similar to the MUSM 3735 (O'Keefe, 2008; Schmeisser
McKean, 2012), some are consistent with the MUSM 3735
morphology (Fig. 3). Therefore at least two groups, Leptocleididae
and Elasmosauridae show morphologies compatible with the ma-
terial preserved on the MUSM 3735. The MUSM 3735 also has a
biochron compatible with MUSM 3735. As no other diagnostic el-
ements are preserved, the specimen is referred as Plesiosauria
indet. This is the first record of Plesiosauria from the Morro Solar
Group and the second Early Cretaceous record of plesiosaurians
from the Pacific margin of South America (O'Gorman et al., 2015).
7. Conclusions
The overall bone morphology of MUSM 3735 indicates that it
belongs to Plesiosauria, as characterized by the propodium. MUSM
3735 shows affinities with at least two plesiosaur groups, Lep-
tocleididae, Elasmosauridae and Polycotylidae. This is the first re-
cord of Plesiosauria from Peru and the second Early Cretaceous
record of plesiosaurians from the Pacific margin of South America.
Acknowledgements

6. Discussion
MUSM 3735 shows features that clearly indicate plesiosaurian
affinities, such as propodium with the presence of transphyseal
foramina on capitulum, dorso-ventral compression, presence of a
The authors thank the anonymous reviewers of this paper for
their valuable contributions and observations. We also thank
Departamento de Paleontología de Vertebrados, Museo de Historia
Natural (UNMSM). Thank Alí Altamirano for take the photos, A.
Benites, R. Varas and L. Tejada for provide bibliography (the first
 I. Meza-V
elez, J.P. O'Gorman / Cretaceous Research 106 (2020) 104247
three of them from the afore mentioned institution and L. Tejada

gico Minero y Metalúrgico).
from Instituto Geolo
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