New Turtles (Chelonia) from the Late Eocene Through Late Miocene of the Panama Canal Basin

Abstract

Four distinct fossil turtle assemblages (Chelonia) are recognized from the Panama Canal Basin. The oldest, from the late Eocene–early Oligocene Gatuncillo Formation, is dominated by podocnemidid pleurodires. The early Miocene Culebra Formation includes both podocnemidids and trionychids. The early to middle Miocene Cucaracha Formation includes taxa classified in Geoemydidae (including Rhinoclemmys panamaensis n. sp.), Kinosternidae (represented by Staurotypus moschus n. sp.), large testudinids, trionychids, and podocnemidids, and finally, the late Miocene Gatun Formation records cheloniid sea turtles. These fossils include the oldest known representatives of Rhinoclemmys, the oldest record of kinosternids in Central America with a more extensive southern paleodistribution for Staurotypus and staurotypines in general, early occurrences of giant tortoises in the Neotropics, the oldest occurrence of soft-shell turtles in the tropics, the oldest late Eocene–early Oligocene Neotropical occurrences of podocnemidids. The Panamanian fossil turtles represent clades that are primarily endemic to North America, showing their very early arrival into the Neotropics prior to the complete emergence of the Isthmus of Panama, as well as their first contact with Caribbean-South American pleurodires by the early Miocene.

Full text

NEW TURTLES (CHELONIA) FROM THE LATE EOCENE THROUGH
LATE MIOCENE OF THE PANAMA CANAL BASIN
EDWIN CADENA,
1,2
JASON R. BOURQUE,
1
ALDO F. RINCON,
1
JONATHAN I. BLOCH,
1
CARLOS A. JARAMILLO,
2
AND BRUCE J. MACFADDEN
1
1
Florida Museum of Natural History, Dickinson Hall, Gainesville, FL 32611, USA,
,eacadena@ncsu.edu.; ,jbourque@flmnh.ufl.edu.; ,jbloch@flmnh.ufl.edu.; and
2
Smithsonian Tropical Research Institute, Panama, AA 34002-
0948, Panama, ,jaramilloc@si.edu.
ABSTRACT—Four distinct fossil turtle assemblages (Chelonia) are recognized from the Panama Canal Basin. The
oldest, from the late Eocene–early Oligocene Gatuncillo Formation, is dominated by podocnemidid pleurodires. The
early Miocene Culebra Formation includes both podocnemidids and trionychids. The early to middle Miocene
Cucaracha Formation includes taxa classified in Geoemydidae (including Rhinoclemmys panamaensis n. sp.),
Kinosternidae (represented by Staurotypus moschus n. sp.), large testudinids, trionychids, and podocnemidids, and
finally, the late Miocene Gatun Formation records cheloniid sea turtles. These fossils include the oldest known
representatives of Rhinoclemmys, the oldest record of kinosternids in Central America with a more extensive
southern paleodistribution for Staurotypus and staurotypines in general, early occurrences of giant tortoises in the
Neotropics, the oldest occurrence of soft-shell turtles in the tropics, the oldest late Eocene–early Oligocene
Neotropical occurrences of podocnemidids. The Panamanian fossil turtles represent clades that are primarily
endemic to North America, showing their very early arrival into the Neotropics prior to the complete emergence of
the Isthmus of Panama, as well as their first contact with Caribbean-South American pleurodires by the early
Miocene.
INTRODUCTION
E
XTANT TURTLES from North and Central America are
exclusively represented by cryptodires, whereas in South
America pleurodires are dominant and cryptodires less diverse.
This modern distribution of turtles in the New World is the
result of a complex and poorly documented biogeographical
history resulting from endemism, diversification, interchange
and mixing of lineages from different geographic sources, all
these events potentially influenced by geologic history.
The most recent geological event with profoun d implica-
tions in the dispersal and interchange of biota between
North-Centra l America and South Ame rica, also ca using the
isolation betw een Carib bean and Pacific ma rine faunas, was
the formation of th e Isthmus of Panama about 4 Myr ago
(Barley et al., 2010; Coates et al., 2004; Coates and Obando,
1996; Herr era et al., 2008; Webb, 1985). The emer gence of the
Isthm us of Panama allowed for the dispersal of some cryp-
todires from North to South America, including members of
the Kinosternidae, Chelydridae, Emydidae, and Geoemydi-
dae (Pritchard, 1984). Molecular data recognizes at least
three different dispersal events of the Geoemyididae (Bata-
guridae) from North to South America through Panama after
the emergence of the isthmus (Le and McCord, 2008). In
contrast, the arrival to South America by members of the
Trionychidae and Testudi nidae (including giant tortoises)
may have taken place from Florida through the Antilles
(Head et al., 2006; Pritchard, 1984).
In the early Miocene (around 20 Myr) North-Central
America was separated from South America by a narrow
strait (Kirby et al., 2008; Montes et al., in press) (Fig. 1.1).
During the beginning of the middle Miocene (around 15 Myr)
the Culebra strait was closed, expanding the Central American
Peninsula farther to the east. However, the Atrato seaway still
separated the peninsula from South America (Kirby et al.,
2008) (Fig. 1.2). The opening and closing of narrow straits at
the easternmost tip of the Central American Peninsula
persisted until the rise of eastern Panama and final closure
of the Atrato Seaway by the Pliocene (4 Myr) that connected
South with Central America (Kirby et al., 2008) (Fig. 1.3).
Recent fieldwork by geologists and paleontologists from the
Florida Museum of Natural History–University of Florida,
the Smithsonian Tropical Research Institute, and the Panama
Canal Authority was conducted in order to relocate previously
collected sites (Whitmore and Stewart, 1965) and to prospect
for new fossil localities. The later has been facilitated by the
recent excavations associated with the widening of the Panama
Canal resulting a once-in-a-century opportunity to explore
fresh exposures of fossiliferous rocks. The result is that many
new mammal, crocodile, snake, turtle, fish, crab, mollusk, and
plant fossils have been recovered. One of the primary
conclusions coming from the early-middle Miocene mamma-
lian fossil record is that all of the recovered taxa have exclusive
affinities with taxa from North America (MacFadden, 2006;
MacFadden, 2009; MacFadden et al., 2010).
Previously the known fossil record of turtles from southern
Central America has been sparse, based on fragmentary
specimens that include: 1) Geochelone costarricensis from the
Oligocene–Miocene of Costa Rica (Coates, 1999; Segura, 1944);
2) carapace pieces of Rhinoclemmys sp. and shells of Geochelone
sp. from the late Miocene of Honduras (D. Webb and Perrigo,
1984); 3) isolated costal bones of a trionychid from the Pliocene of
Costa Rica (Laurito et al., 2005); 4) Rhinoclemmys nicoyana from
thelatePleistoceneCostaRica(Acun˜a, 1996); and 5) undescribed
Miocene fossil turtles from outcrops along the Panama Canal
(MacFadden, 2006; Whitmore and Stewart, 1965).
Here we describe fossil turtles initially reported by
Whitmore and Stewart (1965) and MacFadden (2006) as well
as new specimens recently collected along the Panama Canal
area and the Panama Colon Highway (Fig. 2). The fossils are
from the: 1) late Eocene–early Oligocene Gatuncillo Forma-
tion (Montes et al., in press); 2) early Miocene Culebra and
Cucaracha formations cropping out along the Gaillard Cut
(Kirby et al., 2008; Montes et al., in press); and 3) late
Miocene Gatun Formation (Coates, 1999).
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Journal of Paleontology, 86(3), 2012, p. 539–557
Copyright ’ 2012, The Paleontological Society
0022-3360/12/0086-0539$03.00
539

The available geochronology for these lithostratigraphic
units includes: 1) Sr
87
/Sr
86
chemostratigraphy for the Culebra
Formation (Kirby et al., 2008); 2) magmatic U/Pb dates (19.3
6 0.4 Ma) calculated for the lowermost part of the Culebra
Formation (Montes et al., in press); 3) biostratigraphy for the
Gatuncillo (Montes et al., in press); and 4) the proposed ages
for the late Miocene Gatun Formation (Coates, 1999).
SYSTEMATIC PALEONTOLOGY
Institutional abbreviations.—AMNH, American Museum of
Natural History, Division of Vertebrate Paleontology, New
York, U.S.A.; CRI, Peter Pritchard’s collection at the
Chelonian Research Institute, Oviedo, Florida, U.S.A.;
UMNH, The University of Utah, Utah Museum of Natural
History, Salt Lake City, U.S.A.: UF, University of Florida,
Florida Museum of Natural History Vertebrate Paleontology
Collection, Gainesville, U.S.A.; UF (H), University of Florida,
Florida Museum of Natural History Herpetology Collection,
Gainesville, U.S.A.; USNM, Smithsonian National Museum of
Natural History, Paleobiology, Washington, U.S.A.
T
ESTUDINES (Batsch, 1788)
C
RYPTODIRA (Cope, 1868)
G
EOEMYDIDAE (Theobald, 1868)
R
HINOCLEMMYS (Fitzinger, 1836)
Included species.—R. punctularia punctularia (Daudin,
1801), R. punctularia flammigera (Paolillo, 1985), R. annulata
(Gray, 1860), R. aereolata (Dume´ril et al., 1851), R. diademata
Journal of Paleontology pleo-86-03-13.3d 21/2/12 10:38:08 540 Cust # 11-106R
FIGURE 1—Early Miocene to Present paleogeographical reconstructions of Central America, modified from Kirby et al. (2008). Numbers represent
localities with fossil turtles. 1, early Miocene, hypothetical distribution of North American cryptodires (trionychids) and Caribbean-South American
pleurodires (podocnemidids); 2, middle-late Miocene arrival of cryptodires (trionychids and testudinids) into South America; 3, Pliocene–Present arrival of
kinosternids and geoemydids into South America, and complete retreat of trionychids toward North-Central America and podocnemids into more fluvial
environments of South America; 4, geologic history of the Panama Canal Basin modified from Kirby et al., 2008, showing the fossil record content for each
formation. Past (hypothetical) and modern distribution for pleurodires is shown by the gray dash-gap-dash line enclosing the land areas and costal margins,
and for cryptodires is shown by the gray dash-dash line. A, Panama Canal Basin; podocnemidids, trionychids; B, Pubenza Locality, Colombia;
podocnemidids ‘unpubl. data’ Cadena; C, Panama Canal Basin; trionychids, testudinids, geoemydids, and kinosternids; D, La Venta, Colombia;
podocnemidids, testudinids, (Wood. 1997); E, Costa Rica; geoemydids (Segura, 1944); F, Honduras; geoemydids and testudinids (Web & Perrigo, 1984);G,
Urumaco, Venezuela; geoemydids and trionychids (Sanchez-Villagra & Scheyer, 2010); H, Costa Rica; trionychids (Laurito et al., 2005); I, Costa Rica;
geoemydids (Acun˜a and Laurito, 1996); J, Pubenza locality, Colombia; kinosternids (Cadena et al., 2007). Past (hypothetical) and modern distribution for
pleurodires is shown by the gray dash-gap-dash line enclosing the land areas and costal margins, and for cryptodires is shown by the gray dash-dash line.
540 JOURNAL OF PALEONTOLOGY, V. 86, NO. 3, 2012

(Mertens, 1954), R. funerea (Cope, 1875), R. melanosterna
(Gray, 1861), R. nasuta (Boulenger, 1902), R. pulcherrima
pulcherrima (Gray, 1855), R. pulcherrima incisa (Bocourt,
1868), R. pulcherrima manni (Dunn, 1930), R. pulcherrima
rogerbarbouri (Ernst, 1978), R. rubida rubida (Cope, 1870), R.
rubida perixantha (Mosimann and Rabb, 1953), and R.
panamaensis n. sp.
R
HINOCLEMMYS PANAMAENSIS new species
Figure 3.1–3.5
Holotype.—UF 237887, articulated shell, missing only the
posterior carapace and both xiphiplastra.
Referred material.—UF 257068 (Fig. 3.5), neural 3, UF
257066 (right epiplastron); UF 257067 (left epiplastron).
Etymology.—Named for its provenance, the country of
Panama.
Occurrence.—Recovered from a sandstone in the upper part
of the early to middle Miocene, Cucaracha Formation located
just under the west side of the Centenario Bridge, in the
Panama Canal Basin (MacFadden et al., 2010; more detailed
locality information available at the Division of Vertebrate
Paleontology, Florida Museum of Natural History).
Diagnosis.—Differs from all species of Rhinoclemmys in its
large adult size (estimated carapace length of between 38 to
41 cm); elongate carapace and plastron; dorsum of carapace
smooth and acarinate to only faintly keeled medially; anterior
peripherals 1–3 relatively flat with little to no step along the
visceral marginal scale sulci; and an anteriorly wide gular
scute.
Comparative description.—The type specimen of Rhinoclem-
mys panamaensis n. sp. is a partial articulated shell that is
30.5 cm long (from the anterior margin of the shell to the right
hypo-xiphiplastral suture) by 25.4 cm wide. Recovered
elements include: the nuchal, neurals 1–4, paired costals 1–4
and partial costal 5, paired peripherals 1–6 and right
peripheral 7, paired epiplastra, the entoplastron, paired
hyoplastra, and paired partial hypoplastra. The shell is
rectangularly ovoid in shape, with the anterior margins of
the carapace and plastron straight and smooth. The shell is
flattened, distorted by fractures and crushing, particularly
along the midline of the plastron. The plastral forelobe is
almost rectangular in shape, similar to that seen in other
geoemydids including other species of Rhinoclemmys.
Rhinoclemmys panamaensis n. sp. is the largest known
member of the genus and is morphologically most similar to
extant R. funerea. It is recognized as a geoemydid by the
presence of inguinal and axillary musk duct foramina
(Character 36; Claude and Tong, 2004) with a discernible
axillary pore on the ventral side of the left peripheral 3. It is
further classified in Rhinoclemmys based on the following
shared characters: an almost smooth transition from the
dorsal margin of the gular scale to the visceral surface of the
epiplastron; presence of a very small axillary scale; nuchal with
a strong posteromedial concavity on the ventral surface;
visceral gular margin very narrow at the midline, gradually
widening or flaring posteriorly and overlapping or nearly
overlapping the humeral along the margin of the epiplastron;
moderately keeled nuchal with narrow and elongate semi-
triangular cervical scale; nuchal with very broad sutural
contact with neural 1 (Carr, 1991; Hutchinson, 2006).
Carapace.—The specimen lacks distinct dorsolateral keels
along the costals as in other Rhinoclemmys species and
Echmatemys which is the typical condition in these two New
World genera (Claude and Tong, 2004). Whereas R. pana-
maensis lacks distinct lateral carination on the carapace in the
form of thickened bony ridges, it does possess a faintly
discernible medial keel restricted to the nuchal and anterior
neural 1. It is possible that a keel was present along the
posterior-most neurals as in modern Rhinoclemmys species,
however the rear portion of the fossil carapace is not
preserved. The faint anterior medial keel (or hump) exhibited
in UF 237887 is similar to the condition observed in R.
funerea.
Nuchal.—The nuchal bone is wider than long with a shallow
embayment along the anterior margin. In this way, it
resembles that of other extant species of Rhinoclemmys,as
well as other Miocene aged nuchals referred to Rhinoclemmys
sp. from Panama (UF 237892) and Honduras (UF 46671;
Webb and Perrigo, 1984). The cervical scale is trapezoidal in
shape and narrower than wide. In contrast, the Eocene
geoemydid Bridgeremys pusilla Hutchison, 2006 has a wider
than long cervical, and primitive geoemydids such as
Palaeoemys testunidiformis and Echmatemys septaria have a
small squared cervical. Posteriorly on the nuchal, there is a
very broad sutural contact with neural 1 typical of other
Rhinoclemmys species.
Neurals.—Neural 1 of R. panamaensis is oval in shape,
broadly contacts the nuchal and does not contact costal set 2.
In other Rhinoclemmys where the neural 1 contacts costal set
2, the shape tends to be hexagonal. Neural 2 of R. panamaensis
is octagonal and contacts costal set 1 by way of shortened
anterolateral sides. This character is shared with R. funerea
and some R. punctularia
(e.g., CRI 03190, CRI 0796, CRI
1813, CRI 3706) (UMNH 11440, 11445 sensu (Hutchinson,
2006; personal observation) in which at least one of the
anterolateral sides contacts at least one of the first costals. All
other Rhinoclemmys tend to have a hexagonal neural 2 that is
widest posteriorly, and lacks contact with costal set 1. Contact
Journal of Paleontology pleo-86-03-13.3d 21/2/12 10:38:33 541 Cust # 11-106R
FIGURE 2—Map of the Panama Canal Basin. Star symbols indicate
early Miocene fossil localities from the Culebra and Cucaracha
Formations (Lirio East, Lirio West, Lirio Norte, Centenario Bridge,
Cartagena Hill), one late Miocene site from the Gatun Formation (Banco
Este), and one from the Oligocene, Gatuncillo Formation (Gatuncillo site).
CADENA ET AL.—EOCENE–MIOCENE TURTLES FROM PANAMA 541

of neural 2 with costal set 1 is considered primitive and shared
with Bridgeremys pusilla (Hutchinson, 2006), Echmatemys
septaria (Hay, 1892), Palaeoemys testudiniformis, and P.
hessiaca (Claude and Tong, 2004). Neural 3 of R. panamaensis
is nearly rectangular in shape but is distorted due to crushing.
Costals.—Costals 1–5 of R. panamaensis are preserved on
both sides of the carapace, and the right costal 5 only partially
preserved. Vertebral scales 1–3 are slightly longer than wide,
and vertebral 2 is the longest. Presence of longer than wide
vertebral scales is primitive and seen in the fossils Palaeoemys
testudiniformis and Echmatemys septaria. Vertebrals that are
as long as wide or wider than long are typical for most derived
geoemydids. Pleural scales 1–2 are preserved on both sides of
the carapace, as well as the anterior-most portion of the right
pleural 3. Contact between pleural 2 and marginal 4 has been
suggested as derived for geoemydids (Claude and Tong, 2004);
however after examination of a considerable number of
specimens for eight of the nine extant species of Rhinoclemmys
(see Appendix 1) we conclude that this condition is highly
variable and in most of the cases marginal 4 only contacts
pleural 1. This is also the condition in R. panamaensis and the
primitive geoemydids Palaeoemys testudiniformis and Bridger-
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FIGURE 3—Rhinoclemmys panamaensis holotype, UF 237887. 1, 2, carapace in dorsal view; 3, 4, plastron and carapace in ventral view; 5, dorsal view,
R. panamaensis, UF 257068, neural 3. Abbreviations: abd5abdominal scute; ce5cervical scute; cos5costal; ent5entoplastron; epi5epiplastron;
fem5femoral scute; gul5gular scute; hum5humeral scute; hyo5hyoplastron; hyp5hypoplastron; intg5intergular scute; ma5marginal scute;
ne5neural; pec5pectoral scute; pe5peripheral; pl5pleural; ve5vertebral scute.
542 JOURNAL OF PALEONTOLOGY, V. 86, NO. 3, 2012

emys pusilla. The marginal scales 1–7 are discernible on the left
side of the carapace, whereas on the right side only marginal
1–3 are clearly delimited.
Peripherals.—Peripherals 1–6 are preserved on the left side
of the carapace, and peripherals 1–7 on the right. On the
ventral surface, the transition or slope between scaled and
unscaled parts of peripheral 1–2 is moderate and almost
smooth, as is the condition for all species of Rhinoclemmys
(Hutchinson, 2006). On the ventral side of peripheral 3, the
anterior musk duct pore is situated along the axillary scale
sulcus just proximal to the marginal 3–4 sulcus.
Plastron (epiplastron).—The epiplastron, is overall similar
to that of Rhinoclemmys and other geoemydids. The medial
contact between the epiplastra is very short, and their lateral
margins are only slightly convex to relatively straight as in the
primitive geoemydid Palaeoemys testudiniformis. Typically
geoemydids have slightly convex lateral margins on the
epiplastra. Rhinoclemmys panamaensis has a triangular shaped
gular scute with its posterior tip overriding a very small area
on the anterior portion of the entoplastron. The degree of
gular overlap on the entoplastron is highly variable within
geoemydids, and can range from being restricted to the
epiplastra, to considerably overlapping the anterior portion of
the entoplastron. On the dorsal surface of the epiplastra of UF
237887 (Fig. 4.1), the transition from the most posterolateral
margin of the gular to the visceral surface is marked by an
almost smooth step, and the medial contact between the gulars
is very short as is R. nassuta and R. melanosterna. The dorsal
humeral scale overlap is also more reduced than is typical for
other geoemydids.
An epiplastron with straight anterior and lateral edges, and
without a distinct notch at the contact between the gular and
humeral scales, is primitive and seen in the early Eocene
Palaeoemys testudiniformis and Paleoemys hessiaca. Bridger-
emys pusilla and Echmatemys septaria from the middle Eocene
show a derived epiplastron with slightly convex lateral edges
and a faint tuberosity along the distal margin of the gulars. In
these taxa the dorsal surface the gular and humeral scales is
very wide, and the gulars are squared with their posterior
margins straight and perpendicular to the midline near the
entoplastron. Rhinoclemmys panamaensis (Fig. 4.1) possesses
an epiplastron with straight lateral edges (plesiomorphic);
however, the notch at the contact between the gular and
humeral is well defined as in all other derived geoemydids.
Extant species of Rhinoclemmys display a considerable variety
of shapes and scale arrangements on the anterior plastral lobe
with a marked increase in ornamentation and distinction
between the gular and the humeral scales, particularly on the
dorsal surface (Fig. 4). For instance, R. annulata and R.
areolata have a rectangular gular scale that is wider than long
with the posterior edge straight and perpendicular to the
midline of the plastron. This is similar to Bridgeremys pusilla
and Echmatemys septaria; however, in these taxa the gular is
slightly longer than wide. R. diademata, R. funerea, R.punc-
tularia, and R. pulcherrima all have triangular gular scales with
short medial contact. A diagnostic character for R. pulcher-
rima is the presence of a series of narrow ridges on the dorsal
surface of the humeral scale being parallel to the gular-
humeral sulcus. In R. nassuta the diagnostic character is a
dorsally concave gular scale, for R. punctularia gulars that are
restricted to the epiplastron andnot overlapping onto the
entoplastron as in other species. Epiplastron comparisons for
extant species of Rhinoclemmys and R. panamaensis are
illustrated in Figure 4, showing the arrangement of scales in
dorsal view.
Plastron (entoplastron).—The entoplastron of R. pana-
maensis is bell-shaped, slightly longer than wide, with straight
anterior sides and convex posterior sides typical of other
geoemydids. The humeral-pectoral sulcus crosses the posterior
portion of the entoplastron and the anterior hyoplastra as in
most geoemydids. This sulcus overlaps the posteriomedial
portion of the epiplastra in R. diademata (Carr, 1991) and
occasionally in R. funerea and
R. punctularia.
Plastron (hyo-hypoplastra).—The medial contact between
the hyoplastra is shorter than the contact between the
hypoplastra. The axillary and inguinal buttresses are well
developed (despite being slightly crushed), and both the
anterior and posterior musk duct foramina are discernible
on the left side of the plastron. The presence of paired axillary
and inguinal musk duct foramina is considered synapo-
morphic for all turtles (Hirayama, 1984; Le and McCord,
2008).
The pectoral-abdominal sulcus is anterior to the hyo-
hypoplastral suture as in other geoemydids. Conversely, in
Bridgeremys pusilla this sulcus is medially very close to the
hyo-hypoplastral suture. A very small axillary scale is present
at the most anterolateral corner of both hyoplastra, in contrast
to Bridgeremys pusilla, Echmatemys septaria and Palaeoemys
sp., which lack this scale. As in all species of Rhinoclemmys the
axillary is enclosed by the pectoral scale.
R
HINOCLEMMYS sp.
Here we include other fossil material, most of them isolated
bones that lack of the diagnostic characters defined above for
R. panamaensis, but they share enough morphological
similarities to be considered as Rhinoclemmys.
Referred material.—UF 242075, USNM PAL171020A,
USNM PAL171020C, USNM PAL171021, UF 237888, UF
237889, UF 237892, UF 237895, UF237897, UF237995.
Occurrence.—The UF specimens are from the same locality
and age as for Rhinoclemmys panamaensis. Additionally,
USNM PAL171020 (A–C) and USNM PAL171021 were
collected by Whitmore and Stewart at the Culebra Reach,
Station 1998 + 00, 600 feet W of center line of Panama Canal,
Cucaracha Formation, early to middle Miocene in age (Kirby
et al., 2008).
Description and remarks.—UF 242075 (Fig. 5.1, 5.2) pygal.
This specimen possesses a small medial notch along the
margin, and is longer than wide. It resembles juvenile
specimens of Rhinoclemmys funerea and Rhinoclemmys areo-
lata (Gray, 1860), e.g., UF(H) 54199 (Fig. 5.3, 5.4).
USNM PAL171021 (Fig. 5.5, 5.6) right costal 1; UF
237888, left costal 1; and UF 237889, left costal 1. These
specimens represent different individuals of similar size,
approximately two times smaller than the costal 1 in the
holotype of Rhinoclemmys panamaensis. A distinct sulcus
between vertebrals 1 and 2, and between these vertebrals and
pleural 1 is present on the dorsal surface of these specimens.
On the ventral and distal surfaces of the costal, the axillary
buttress scar is preserved and well developed as in other
geoemydids.
UF 237892 (Fig. 5.7, 5.8), nuchal. This specimen has a very
narrow cervical scale on the dorsal surface, accompanied by
thickened bone in this area. The sulcus between marginal 1
and vertebral 1 is visible at the anterior-most portion of the
nuchal. There is a small portion of pleural 1 that contacts
vertebral 1 along the lateral sutures. Ventrally, the visceral
sulcal margin is adjacent to a deep medial concavity. In most
aspects UF 237892 and UF 46671 (Fig. 4.9) from the late
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Miocene of Honduras (Webb and Perrigo, 1984) resemble the
nuchals of Rhinoclemmys spp.
UF 237881 (Fig. 5.10), neural 5. This specimen has well-
discerned intervertebral sulci, and resemble other species of
Rhinoclemmys spp.
USNM PAL171020A (Fig. 5.11, 5.12), left xiphiplastron.
This specimen possesses a rounded posterior tip. The femoral-
anal sulcus is strongly incised on the ventral surface and the
anal scale overrides almost 60% of the bone. On the dorsal
surface, the visceral sulcus of the femoral and anal scales is
marked by a shallow groove that runs almost parallel to the
lateral margin of the xiphiplastron. USNM PAL171020A
resembles the xiphiplastra of Rhinoclemmys spp.
USNM PAL171020C (Fig. 5.13, 5.14), right xiphiplastron.
This specimen resembles USNM PAL171020A in many
aspects, with the exception of the posterior tip of the element
having a wide shallow embayment, in all aspects is similar to
the xiphiplastron of Rhinoclemmys spp.
UF 237895 (Fig. 5.15, 5.16), left epiplastron; UF 237895
(Fig. 5.17, 5.18), right epiplastron. Both specimens resemble
the epiplastra of Rhinoclemmys diademata, R. funerea,
R.punctularia, and R. pulcherrima in both dorsal and ventral
views, having the same arrangement of scales and contacts
between them.
UF 237795 (Fig. 5.19–5.21), left dentary. This specimen is
the only turtle cranial element recovered from the Panama
Canal Basin. It is 2.8 cm long which is much larger than that
of R. funerea (1.5 to 2 cm long average) and other living
species of Rhinoclemmys (1 to 1.5 cm long average). Due to its
large size, this specimen may be attributable to R. panamaen-
sis. In dorsal view the lingual and labial ridges are almost
parallel, and rounded at the symphysis as in most living
species of Rhinoclemmys. In lateral view, the labial ridge is
slightly higher than the lingual as in all living species of
Rhinoclemmys. The sulcus cartilaginis meckelii ends at the
symphysis in a sinuous path, in contrast to living species in
which this sulcus is nearly straight from the posterior to the
symphysis. The sulcus cartilaginis meckelii is very wide,
similar to that of R. funerea. This sulcus is narrower in other
living species of Rhinoclemmys.The foramen alveolare inferius
is clearly visible on UF 237795.
K
INOSTERNIDAE (Hay, 1892)
S
TAUROTYPUS (Wagler, 1830)
S
TAUROTYPUS MOSCHUS new species
Figure 6.1–6.3
Holotype.—UF 242076, left peripheral 2.
Occurrence.—Same locality and age as for Rhinoclemmys
panamaensis.
Etymology.—From Latin moschus for ‘‘musk’’ or ‘‘musky’’
in reference to the well-developed anterior musk duct groove
present in this species.
Diagnosis.—Presence of a well-developed deeply incised
anterior musk duct groove that runs parallel with and closely
adjacent to the visceral scale margin for marginals 1 and 2
(differs from S. triporcatus and S. salvinii which possess a very
faintly incised anterior musk duct groove that is further from
the marginal 1–2 sulcus); terminus for the anterior musk duct
groove situated anteriorly on peripheral 2 or on the peripheral
1–2 suture (differs from S. triporcatus and S. salvinii in which
the anterior musk duct terminus lies on peripheral 1); marginal
scales 1 and 2 relatively narrow to the height of the peripheral
in dorsal aspect; costiform process of the nuchal with slight
contact to the anterior-most portion of peripheral 2 at the
peripheral 1–2 suture.
Description and comparisons.—UF 242076 (Fig. 6.1–6.3) is
from an adult individual with fully-formed sutures, compara-
ble in size to extant members of the genus Staurotypus
examined here with a carapace length between 24 and 27 cm.
On the dorsal surface, marginals 1–2 are moderately bulbous
and narrow, with a strong notch at the intermarginal sulcus
along the lateral margin of the peripheral. Such a notch is
absent to only slightly-developed in S. salvinii and S.
triporcatus. The outer rim of the marginals is notably thick
and squared, with a distinct lip (or crest) along the dorsal edge.
The outer marginal rim is typically much thinner and tapered
in that of modern Staurotypus. Arguably, the most significant
feature on the element is the presence of the deeply incised
musk duct on the visceral face and the atypical position of its
terminal point along the peripheral 1–2 suture. The presence
of an anterior musk duct groove is a synapomorphy for
Kinosternidae and generally spans from peripheral 4 to its
terminus well into peripheral 1 in Staurotypus (Hutchinson,
1991).
Staurotypus moschus n. sp. is the most southern-occurring
staurotypine yet known. Its extant congeners are S. salvinii
from the Pacific lowlands of Southern Mexico to Guatemala
and S. triporcatus from the base of the Yucatan Peninsula
(Bonin et al., 2006; Iverson, 1985). Interestingly, these three
species each exhibit different degrees of development of the
anterior musk duct groove that appear to represent a
morphocline, with S. moschus being the most deeply incised,
S. salvinii being only moderately to weakly so, and S.
triporcatus very weakly developed. This character has been
coded as ‘‘weakly incised’’ for the genus Staurotypus in
previous phylogenetic analyses (Hutchinson, 1991). A weakly
incised musk duct groove has been interpreted as primitive due
to its absence in early kinosternoids (Hutchison, 1991). The
antiquity of S. moschus and its possession of a seemingly
derived deeply incised musk duct groove could provide
evidence that a weakly developed musk duct in the extant
taxa S. salvinii and S. triporcatus represents a secondary
character state reversal. Alternatively, extant Staurotypus
could represent a more basal lineage than S. moschus. The
distance of this groove from the visceral marginal sulcus
represents a morphocline, with the groove being closest to the
marginal sulcus in S. moschus, moderately close in S. salvinii,
and farthest from the sulcus in S. triporcatus. For reasons
discussed here, including depth and distance of the musk duct
from the ventral marginal sulcus, S. moschus is tentatively
interpreted here as having its closest affinities with
S. salvinii.
A small pit that would receive the distal-most tip of the
costiform process is just visible on the visceral face of UF
242076, at the peripheral 1–2 suture, indicating that the
costiform process would have extended across the entire
visceral face of peripheral 1. Just posterior to this pit, there is a
slight swelling of the element, also present in S. salvinii and S.
triporcatus. However, this pit is more substantial in the latter
Journal of Paleontology pleo-86-03-13.3d 21/2/12 10:39:04 545 Cust # 11-106R
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FIGURE 4—Epiplastra of species of Rhinoclemmys in dorsal view. 1, R. panamaensis UF 237887; 2, R. annulata CRI 3638; 3, R. areolata CRI 8318; 4,
R. diademata CRI 1516; 5, R. funereal CRI 1730; 6, R. melanosterma CRI 4198; 7, R. nasuta CRI 2638; 8, R. pulcherrima CRI 1839; 9, R. punctularia CRI
2702. Abbreviations: gul5gular scute; hum5humeral scute; visc5visceral surface.
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FIGURE 5—Rhinoclemmys sp., UF 242075, most posterior portion of the pygal. 1. dorsal view; 2, ventral view, Rhinoclemmys areolata UF(H) 54199
complete pygal; 3, dorsal view; 4, ventral view, Rhinoclemmys sp. USNM PAL 171021, right costal 1; 5, ventral view; 6, dorsal view, Rhinoclemmys sp., UF
237892, nuchal bone; 7, dorsal view; 8, ventral view, Rhinoclemmys sp., UF 46671, nuchal bone referred in Web and Perrigo (1984), from late Miocene of
Honduras; 9, dorsal view, Rhinoclemmys sp., UF 237881, neural 5; 10, dorsal view, Rhinoclemmys sp., USMN PAL171020A, left xiphiplastron. 11. Dorsal
view, 12, ventral view, Rhinoclemmys sp., USMN PAL171020C, right xiphiplastron; 13, ventral view; 14, dorsal view, R. panamaensis, UF257066, left
epiplastron; 15, dorsal view; 16, ventral view, R. panamaensis, UF257067, right epiplastron; 17, ventral view; 18, dorsal view, R. panamaensis, UF 257195,
left dentary; 19, dorsal view; 20, lateral view; 21, medial view. Scale bar53 cm for 1–12, 14–18; 1 cm for 13, 14; 2 cm for 19–21.
546 JOURNAL OF PALEONTOLOGY, V. 86, NO. 3, 2012

two taxa, with the costiform process usually more intrusive on
peripheral 2 (Hutchinson, 1991; Fig. 5).
T
ESTUDINIDAE (Gray, 1825)
I
NCERTAE SEDIS
Referred material.—USNM V23180, USNM PAL171017,
USNM PAL171020, USNM V23146, UF 242091.
Occurrence.—All USNM specimens were collected by
Whitmore and Stewart at the Culebra Reach, Station 1998 +
00, 600 feet W of center line of the Panama Canal, Cucaracha
Formation. The age for the Cucaracha Formation is early to
middle Miocene sensu (Kirby et al., 2008).
Description and remarks.—USNM V23180 (Fig. 7.1, 7.2),
right coracoid. This specimen is 19 cm long by 14 cm wide
(maximum values) and missing a portion of the posterolateral
margin and most of the dorsal and ventral margins of the
proximal articular area. The medial edge is thinner than the
lateral, and the blade is relatively short, with slightly rounded
and flared distal margin. Proximally, the most central part of
the glenoid surface is preserved, being slightly concave for
articulation with the humerus. The sutural contact with the
scapula is not preserved.
USNM PAL171017 (Fig. 7.3, 7.4), complete right ulna.
This specimen measures l3 cm long by 4 cm wide (maximum
values). The olecranon process is rounded and well developed.
The proximal articular surface is curved and the bicipital
tubercle is poorly preserved. Proximally, the ulna is convex
where it articulates with the humerus, and the most
dorsolateral margin is missing.
USNM PAL171017 and USNM V23180 resemble the ulna
and the coracoid of giant tortoises, such as members of the
Galapagos Island complex Chelonoidis elephantopus USMN
59867 (Fig. 7.5–7.8). These elements could belong to the
Caribbean/Central-South American genus Chelonoidis, or
perhaps represent another extinct giant tortoise clade such as
Hesperotestudo (or Caudochelys).
USNM V23146 (Fig. 7.9), osteoderm. This specimen is
highly curved and elongate, typical of osteoderms found on
the distal forelimbs, hind-quarters, or posterior pedes in
modern tortoises.
USNM PAL171020 (Fig. 7.10, 7.11), right xiphiplastron.
This element is relatively small (5 cm long by 4 cm wide,
maximum values). Ventrally, it possesses a very small anal
scale, which is trapezoidal in shape and short medially. On the
dorsal surface the transition from the margins of the femoral
and anal scales to the visceral surface is strongly marked by a
deep concavity. USNM PAL171020 resembles the xiphiplas-
tron of Chelonoidis.
UF 242091 (Fig. 7.12–7.14), right epiplastron. This speci-
men is 6.8 cm long by 5.9 cm wide. Dorsally it possesses a
broad gular margin, with a deep step posteriorly. The gular-
humeral margins are very thick, being thickest at the posterior
gular. The gular is highly convex in both dorsal and ventral
aspects. Ventrally, the gular-humeral sulcus terminates at the
epi-entoplastral suture, indicating that the posteromedially
terminus of the gulars would have been on the anterior portion
of the entoplastron. In these features it resembles Hesper-
otestudo specimens examined.
C
HELONIIDAE (Gray, 1825)
I
NCERTAE SEDIS
Referred material.—UF 244433 (Fig. 8.1, 8.4), right
costal 4.
Occurrence.—Recovered fro m the lower part of the
middle-late Miocene Gatun Formation (Coates, 1999). The
locality is located on the E astern side of the Gatun Locks,
area of Colon , Northern area of the Panama Can al Basin
(more detailed locality information available at the Division
of Vertebrate Paleontology, Florida Museum of Natural
History).
Description and remarks.—This specimen is 35 cm long by
19 cm wide). Dorsally, the sulci between pleural 2–3 and
between these and vertebral 3 are discernible. In many aspects
this costal resembles the costal bones of cheloniids, particu-
larly of Caretta caretta (Fig. 8.5, 8.6), one of the most
abundant modern sea turtles in the neotropics.
T
RIONYCHIDAE (Baur, 1893)
I
NCERTAE SEDIS
Referred material.—UF 242108 (Fig. 8.7), right costal 1; UF
242088 (Fig. 8.8), costal; UF 242106 (Fig. 8.9), costal.
Occurrence.—UF 242088 is from the same locality and age
as for Rhinoclemmys panamaensis, Cucaracha Formation. UF
242108 and UF 242106 were collected at the Lirio Norte site,
west margin of the Panama Canal, upper member of the
Culebra Formation, early Miocene (Fig. 2; more detailed
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FIGURE 6—Staurotypus moschus holotype, UF 242076, left peripheral 2. 1, dorsal view; 2, ventral view; 3, anterior sutural view. Abbreviations: cp5pit
that accepts the costiform process; mdg5anterior musk duct groove.
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locality information available at the Division of Vertebrate
Paleontology, Florida Museum of Natural History)
Description and remarks.—All of the costals exhibit
ornamentation consisting of ridges and large pits typical of
trionychids or soft shell turtles. The nature of the dorsal
pitting on the fossils shows similarities to the genus Apalone.
Apalone is the only extant North-Central American trionychid
(Meylan, 1987) and is absent from South America today.
P
LEURODIRA (Cope, 1874)
P
ELOMEDUSOIDES (Cope, 1868)
P
ODOCNEMIDIDAE (Cope, 1868)
I
NCERTAE SEDIS
Referred material.—UF 242276, UF 242170, UF 244763,
anterior plastral lobe; UF 242174, right xiphiplastron; UF
242160; right costal 6; UF 242150, left costal 2; UF 242165,
right side of a pelvic girdle; UF 242171, distal and proximal
portions of a right humerus; UF 242097, proximal portion of a
left femur; UF 242111, right peripheral 2; UF 242158, left
peripheral 8; UF 242168, neural 3 or 5?, UF 242161, right
partial xiphiplastron. UF 257070, right epiplastron, UF
242168, left peripheral.
Occurrence.—UF 242097 and UF 242170 were collected at
the Lirio East site (Fig. 2), east margin of the Panama Canal,
conglomeratic sandstones, Culebra Formation, early Miocene.
UF 242161 was collected at the Gatuncillo site, Gatuncillo
Formation, middle to late Eocene (Graham, 1985). UF
257070, was collected at Lirio West, UF 242168 was collected
at the lower segment of Cucaracha Formation, Centenario
Bridge site. All other UF specimens were collected at the Lirio
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FIGURE 7—Testudinidae incertae sedis, USNM V23180, right coracoid. 1, ventral view; 2, dorsal view, USNM PAL 171017, right ulna; 3, ventral
view; 4, dorsal view, Chelonoidis elephantopus, USMN 59867, right coracoids; 5, dorsal view; 6, ventral view, Ch. Elephantopus, USMN 59867, right ulna;
7, dorsal view; 8, ventral view, USNM V23146, osteoderm; 9, lateral view, USNM PAL 171020, right xiphiplastron; 10, ventral view; 11, dorsal view, UF
257065, right epiplastron; 12, 13, dorsal view; 14, medial sutural view, UF 257069, right hypoplastron; 15, dorsal view.
FIGURE 8—Cheloniidae incertae sedis, UF 24443, right costal. 1, 2, dorsal view; 3, 4, ventral view, Caretta caretta, UF 200374, right costal 4; 5, dorsal
view; 6, ventral view, Trionychidae incertae sedis, UF 212108, left costal 1; 7, dorsal view; 8, dorsal view, UF 242088, costal bone; 9, dorsal view, UF
242106, costal bone. Scale bar510 cm for 1–6, 5 cm for 7, 8. Abbreviations: pl5pleural scute; phr5proximal head of the rib; v5vertebral scute.
CADENA ET AL.—EOCENE–MIOCENE TURTLES FROM PANAMA 549

Norte site, west margin of Panama Canal, from conglomerate
channels belonging to the Culebra Formation (more detailed
locality information available at the Division of Vertebrate
Paleontology, Florida Museum of Natural History).
Description and remarks.—UF 24217 (Fig. 9.1, 9.2), artic-
ulated left hyoplastron, hypoplastron and mesoplastron. The
mesoplastron is almost circular in shape and laterally
positioned between the hyoplastron and hypoplastron, as in
all other pelomedusoides (Gaffney et al., 2006). On the ventral
surface the pectoral scales contact the epiplastra as in almost
all other podocnemidids (Gaffney et al., 2011), variable in
Peltocephalus dumerilianus and Podocnemis sextuberculata
(Cadena et al., in press). Also, the pectoral-abdominal sulcus
is anterior to the mesoplastron, which is typical but not
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FIGURE 9—Podocnemididae incertae sedis, UF 242176, left articulated hyoplastron, hypoplastron and mesoplastron. 1, 2, ventral view, UF 242170,
neural 2; 3, dorsal view, UF 242170, left and right xiphiplastra, together with the right hypoplastron; 4, 5, dorsal view, UF 257070, right epiplastron; 6,
ventral view; 7, dorsal view, UF 244763, anterior plastral lobe; 8, 9, ventral view, UF 242160, right costal 6; 10, dorsal view, UF 342150, left costal 2; 11,
dorsal view, UF 242165, right side of a pelvic girdle; 12, lateral view, Podocnemis expansa, AMNH 62942, right side of the pelvic girdle; 13, lateral view,
UF 242171, distal and proximal ends of a right humerus; 14, lateral view, UF 242158, left peripheral 8; 15, dorsal view, UF 242161, right partial
xiphiplastron; 16, dorsal view; 17, ventral view, UF 242168, left peripheral; 18, dorsal view; 19, ventral view. Abbreviations: abd5abdominal scute;
ent5entoplastron; epi5epiplastron; fem5femoral scute; gul5gular scute; hum5humeral scute; hyo5hyoplastron; hyp5hypoplastron; int5intergular
scute; isch sc5ischium scar; mes5mesoplastron; pec5pectoralscute; pub sc5pubic scar.
550 JOURNAL OF PALEONTOLOGY, V. 86, NO. 3, 2012

exclusive of podocnemidids. The abdominal-femoral sulcus
ends laterally at the hypoplastral notch.
UF 242170 (Fig. 9.3–9.5), associated left and right xiphi-
plastron, right hypoplastron, neural 2, costal 3 or 5?, and at
least ten undifferentiated fragments of costals. It is difficult to
discern any scute sulci on the ventral plastron. The open U-
shaped anal notch and concave outline of the mesoplastron at
the anterolateral margin of the hypoplastron are the most
diagnostic features ventrally. Dorsally, both xiphiplastra
preserve the pubic and ischial scars, indicating a strongly
sutured pelvis to the plastron, a synapormophy for Pleurodira
(Gaffney and Jenkins, 2010). The pubic scar is oval-shaped,
oriented almost parallel to the medial margin of the
xiphiplatron. The ischial scar is triangular in outline and ends
medially at an acute point very close to the sutural contact
between both xiphiplastra typical of podocnemidids. Neural 2
is hexagonal and elongate with short anterolateral sides and
lacking a vertebral sulcus. There are two lateral horn-like
projections at the most anteroventral margin where it
articulates with neural 1. The presence of neural bones in
UF 242170 excludes it from being related to Bairdemys
venezuelensis a late Miocene Caribbean-Tropical South
American podocnemoid that lacks the entire neural bone
series (Scheyer et al., 2008).
UF 257070 (Fig. 9.6, 9.7), right epiplastron. On the dorsal
surface, the gular scale is small and triangular, as for most
podocnemidids. UF 244763 (Fig. 9.8, 9.9) is a complete
anterior plastral lobe that is as long as wide, as in all other
podocnemidids. In contrast, bothremydids possess a forelobe
that is much wider than long. The entoplastron is diamond-
shaped and located anterior to the bridge as in other
podocnemidids. This differs from bothremydids in which the
most posterior tip of the entoplastron generally reaches the
level of the bridge. The gular scale is triangular and only
overlaps the epiplastron, whereas than the intergular is very
wide and overlaps the most anterior corner of the entoplas-
tron. The humeral-pectoral sulcus in UF 244763 is situated
anterior to the epi-hyoplastral suture and crosses the
entoplastron at its maximum width as in all other podocne-
midids (Gaffney et al., 2011) except Peltocephalus dumerilianus
and occasionally in Podocnemis sextuberculata (Cadena et al.,
in press). In the latter two taxa this sulcus is posterior to the
epi-hyoplastral suture, and crosses the entoplastron in its
posterior region. In contrast, most bothremydids have a
humeral-pectoral sulcus that is situated well-posterior to the
epiplastron-hyoplastron suture, slightly crossing or touching
the most posterior margin of the entoplastron. The long
anterior plastral lobe of UF 244763 differs from Bairdemys
venezuelensis, which has a shorter anterior plastral lobe
(Gaffney et al., 2006, fig. 275).
UF 242160 (Fig. 9.10), right costal 6. This specimen is
moderately curved dorso-ventrally. Dorsally, the sulcus
between pleural 3 and 4 is visible, as well as the sulcus
between vertebral 4 and pleural 3 and 4.
UF 242150 (Fig. 9.11) left costal 2. The anterior and
posterior margins are almost parallel, with a short postero-
lateral side, and well preserved sulci between pleurals 1 and 2,
and vertebral 2 and pleurals 1 and 2. UF 242160 and UF
242150 were found associated, and presumably belong to the
same individual.
UF 242165 (Fig. 9.12), right side of a pelvic girdle, including
a complete ilium, pubis, and the most distal portion of the
ischium. The acetabulum capsule is almost oval with sutural
contacts between the ilium, pubis, and ischium typical to that
of all pleurodires. It is particularly similar to the pelvic girdle
of the genus Podocnemis, e.g., P. expansa AMNH 62947
(Fig. 9.13).
UF 242171 (Fig. 9.14), partial right humerus. Only the
distal and proximal portions are preserved, with the central
portion missing. Proximally, the lateral process is missing and
the medial process is large and posteroproximally projected.
The articular hemisphere is rounded and lacks a shoulder, a
synapomorphy for pleurodires (Gaffney, 1990). The distal
portion has a rounded and slightly convex capitellum for
articulation with the ulna and radius, and a distinct
ectepicondylar foramen on the anterodistal margin, charac-
teristic of all turtle humeri.
UF 242097, proximal left femur. Dorsally, the acetabular
head is oval and slightly elongate and inclined clockwise with
respect to the longitudinal axis of the bone. The major and
minor trocanters are almost at the same horizontal plane with
prominent lateral projections. Ventrally, the interthrocanteric
fossa is very shallow with a long scar on its posterior edge. In
all aspects UF 242097 resembles the femur of podocnemidids.
Other isolated carapace elements include: UF 242111, right
peripheral 2; UF 242158 (Fig. 9.15), peripheral 8(10 cm long
3 8 cm wide, maximum values); and UF 242168 neural 3 or
5?, in all aspects resembling podocnemidids.
UF 242161 (Fig. 9.16, 9.17), partial right xiphiplastron
(anterolateral-most portion). The anterior-most part of the
sutural scar for the ilium is well-discerned on the dorsal
surface. Ventrally, the posterior-most portion of the femoral-
anal sulcus is preserved, in all features resembling the
xiphiplastron of podocnemidids.
UF 242168 (Fig. 9.18, 9.19), left peripheral. This specimen
exhibits a well-defined visceral marginal step similar to all
other podocnemidids.
DISCUSSION
The turtle record, in contrast to mammals from the same
localities (MacFadden, 2006; MacFadden et al., 2010), shows
that during the early Miocene lineages of both North-Central
American cryptodires and Caribbean- South America pleur-
odires were present in the Panamanian isthmus, previous to its
complete emergence during the early Pliocene.
Four stratigraphically distinct fossil turtle assemblages are
recognized from Panama that lend insight into the evolution
of the group in southern Central America: 1) the late Eocene–
early Oligocene fossils from Gatuncillo Formation; 2) the
early Miocene fossils from Culebra Formation; 3) the early to
middle Miocene fossils from Cucaracha Formation; and 4) the
late Miocene fossils from Gatun Formation.
Assemblage 1 (late Eocene–early Oligocene).—The occur-
rence of podocnemidids, from the late Eocene–early Oligocene
Gatuncillo Formation together with the Paleocene record of
podocnemidids from Colombia (Cadena et al., 2010; Cadena
et al., in press) and the Oligocene record from South Carolina
(Weems, 2009), indicates a very wide distribution for
podocnemidids in the Paleogene neotropics, particularly in
the Caribbean and its coastal regions. It also shows the earliest
northern migration of South American fresh-water turtles
across the Panamanian seaway.
Assemblage 2 (early Miocene).—Turtles from the early
Miocene Culebra Formation are represented by cryptodires
(trionychids) and pleurodires (podocnemidids), inhabiting
deltaic or estuarine environments, a paleodeposition suggested
for the upper segment of the Culebra Formation (Kirby et al.,
2008; Montes et al., in press) Most of the fossils from the
Culebra Formation are complete but disarticulated shell
elements, with very few non-shell elements, and abundant
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evidence of bioerosion produced by mollusks. This suggests
that turtle bones were exposed to pre-burial transport by
streams. This assemblage is the earliest evidence of interaction
between North American trionychids and Caribbean-South
American podocnemidids in the Neotropics.
New World trionychids are represented by only one genus,
Apalone, which is restricted today to temperate–subtropical
regions of North America (Bonin et al., 2006). The early
Miocene occurrence of trionychids from the Panama Canal
basin represents the southernmost record for this family in the
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FIGURE 10—Geographical distributions for extant species of Rhinoclemmys and Staurotypus, after Iverson 1992. 1, R. areolata; 2, R. funereal; 3, R.
pulcherrima; 4, R. rubida; 5, R. punctularia; 6, R. annulata; 7, R. diademata; 8, R. melanostena; 9, R. nasuta; 10, the genus Staurotypus.
FIGURE 11—Gular scute measurements for species of Rhinoclemmys (see specimens examined in Appendix 2). Rhinoclemmys panamaensis has the
widest anterior gular in the genus Rhinoclemmys. 1, anterior gular width (X) versus posterior gular (Y) width; 2, midline gular length (X) versus anterior
gular width (Y). All measurements in mm. Abbreviations: g5gular scute; h5humeral scute.
CADENA ET AL.—EOCENE–MIOCENE TURTLES FROM PANAMA 553

New World, and together with the record from the Castillo
Formation (late Miocene) in North-western Venezuela (San-
chez-Villagra et al., 2004) represents the earliest arrival of
trionychids to the tropics of Central and South America, even
being able to cross the Panamanian seaway before its final
closure. The record suggests that trionychids inhabited a much
wider range in the Americas including tropical climates.
Podocnemidids from Culebra Formation indicate their
continuous occurrence at the most eastern tip of Central
America Peninsula after the Paleogene. Exhibiting large size of
the shell, which we estimate to have been approximately
100 cm long at the midline, based on the size of some isolated
elements including the costals UF 242160 (Fig. 9.10) and UF
242150 (Fig. 9.11), and the peripheral UF 242158 (Fig. 9.15),
which are slightly larger than the largest size reported for
modern Podocnemis specimens, which is 80 cm in P. expansa
(Valenzuela, 2001).
Assemblage 3 (early to middle Miocene).—Represented
primarily by cryptodires, including a trionychid (present in
both the Culebra and Cucaracha formations), and kinoster-
nid, and perhaps multiple species of geoemydid, testudinid,
and podocnemidid. Many of these lineages first occur in
Panama at the lower segment of Cucaracha Formation. This
formation has been interpreted as a paleodeltaic plain (Kirby
et al., 2008). In contrast to the turtles from the Culebra
Formation, the turtles from the Cucaracha Formation are
represented by more complete and articulated specimens, e.g.,
the holotype of Rhinoclemmys panamaensis n. sp., indicating
relatively little pre-burial transport.
The occurrence of Rhinoclemmys panamaensis and Rhino-
clemmys sp. during the early to middle Miocene in the Panama
Canal basin not only represent the earliest record of
geoemydids in Central America but also support the
hypothesis suggested by Le and McCord (2008), based on
molecular data, of an initial dispersal event of the lineage that
contains R. nasuta from Central to South America during the
early Miocene. The strong morphological similarities between
R. panamaensis and R. funerea suggest that the fossil species
was probably largely aquatic.
Closely related to geoemydids are testudinids (Lourenco et
al., 2012). It has been suggested that land tortoises arrived to
Central and South America by passive flotation from North
America and the Antilles during the Miocene (Pritchard,
1984), favored by their adaptations for over-water dispersal
(Meylan and Sterrer, 2000). The fossil testudinids described
here are, together with the Oligocene?–Miocene record of
Geochelone costarricensis described by Segura (1944), the
earliest records of land tortoises from Central America. Some
of the fossil material described here is slightly larger than
modern giant tortoise material, including Chelonoidis elephan-
topus USNM 59867 from the Galapagos Islands and
Dipsochelys gigantea USNM 222495 from the Aldabran
Islands, and indicates the presence of one of the giant tortoise
lineages (perhaps of the Chelonoidis or Hesperotestudo groups)
in Central America during the early Miocene.
The family Kinosternidae is endemic to North America, with
the oldest fossils being from northern Mexico (Brinkman and
Rodriguez De La Rosa, 2006) and the northwestern United
States (Hutchison, 1991). The occurrence of Staurotypus
moschus in Panama represents the oldest record of kinosternids
in Central America. It also indicates a significant former
southeastern range extension for the genus and for the
subfamily Staurotypinae as a whole. Today, this subfamily
comprises only the extant taxa Staurotypus triporcatus,
Staurotypus salvinii, and Claudius angustatus, all with relatively
small ranges in middle Central America. Staurotypines have
been hypothesized as the most primitive living kinosternids
(Hutchison and Bramble, 1981) yet almost nothing is known
about their fossil record. Pleistocene fossils from the Rio de la
Pasion, Guatemala (AMNH 13989, 24176, 24177, 24179)
probably represents
Staurotypus (JRB, personal observation).
Assemblage 4 (late Miocene).—Represented by a sea turtle
(Cheloniidae), from the marine deposits of the Gatun
Formation, which likely represented a shallow marine strait
that connected the Pacific Ocean and the Caribbean Sea
during the late Miocene (Coates and Obando, 1996).
ACKNOWLEDGMENTS
Funding for this project came from National Science
Foundation grants OISE 0638810, EAR 0642528, and EAR
0824299, PIRE 0966884 (OISE, EAR, DRL), the Smithsonian
Institution, the Panama Canal Authority, M. Tupper,
SENACYT, R. Perez SA, the Florida Museum of Natural
History Miss Lucy Dickinson Fellowship. Thanks go to the
Authority of the Panama Canal for access to the Panama
Canal areas. Thanks for access to collections to K. de Querioz,
J. Jacobs (Smithsonian National Museum of Natural History,
Washington, U.S.A); E. Gaffney and C. Mehling, (Fossil
Amphibians, Reptiles, and Birds Collections, Division of
Paleontology, American Museum of Natural History, New
York, U.S.A.); P. Pritchard (Chelonian Research Institute,
Oviedo, Florida U.S.A.); R. Rowe (Utah Museum of Natural
History, Salt Lake City, Utah U.S.A.), K. Krysko (Florida
Museum of Natural History, Herpetology collection, Gaines-
ville, Florida U.S.A.), R. Hulbert, Jr. (Florida Museum of
Natural History, Vertebrate Paleontology collection, Gaines-
ville, Florida U.S.A.). Special thanks to N. Cannarozzi (who
found Rhinoclemmys panamaensis holotype), S. Moron, A.
Gomez, L. Grawe, and all other paleontologists and geologists
working at the Panama Canal Basin.
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ACCEPTED 13 JANUARY 2012
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CADENA ET AL.—EOCENE–MIOCENE TURTLES FROM PANAMA 555

APPENDIX 1
Specimens studied by direct examination.
Institutional Abbreviations
AMNH, American Museum of Natural History, Division of Vertebrate
Paleontology, New York, USA
CRI, Peter Pritchard’s collections at Chelonian Research Institute,
Oviedo, Florida, USA
ICN, Instituto de Ciencias Naturales, Universidad Nacional de
Colombia, Bogota´, Colombia
UMNH, The University of Utah, Utah Museum of Natural History,
Salt Lake City, USA
UF, University of Florida, Florida Museum of Natural History
Vertebrate Paleontology Collections, Gainesville, USA
UF (H), University of Florida, Florida Museum of Natural History
Herpetology Collections, Gainesville, USA
USNM, Smithsonian National Museum of Natural History, Paleobi-
ology, Washington, USA
Rhinoclemmys
R. annulata: CRI 2962, CRI 0049, CRI 0048, CRI 3638, CRI 1132, CRI
2774, CRI 1222, CRI 2773, UMNH 10241, UMNH 11393. USNM 7252,
USNM 8217, USNM 562917. R. areolata: CRI 8318, CRI 2775, CRI 2963,
UMNH 6480, UF(H) 54199, USNM 46002, USNM 46299, USNM 59934.
R. diademata: CRI 1339, CRI 3656, CRI 3826, CRI 1517, CRI 1476, CRI
1516. R. funerea: CRI 4380, CRI 2961, CRI 5494, CRI 4579, CRI 0348, CRI
4578, CRI 5250, CRI 7699, CRI 1730, UMNH 3050, UMNH 8274, UMNH
8277, USNM 59934, USNM 8218, USNM 8273, USNM 8270 R.
melanosterna: CRI 2441, CRI 2434, CRI 2433, CRI 4198. R. nasuta:CRI
2443, CRI 2638, CRI 2448, CRI 2442. R. pulcherrima: CRI 2155, CRI 1839,
CRI 1433, CRI 1090, CRI 7597, CRI 1444, CRI 2895, CRI 6625, CRI 1478,
UMNH 11516, UMNH 11517, USNM 46308, USNM 46309, USNM
46310, USNM 102889, USNM 104626. R. punctularia: CRI 2877, CRI
2871, CRI 3706, CRI 2514, CRI 0345, CRI 4581, CRI 3191, UMNH 9982,
UMNH 11440, USNM 28978, USNM 85022, USNM 86861, USNM
65108. R. rubida: CRI 1234, USNM 45615, USNM 222433.
Staurotypus
S. triporcatus: CRI 1488, CRI 0166, CRI 1105, CRI 0157, CRI 0167,
CRI 0161. S. salvini: CRI 1104, CRI 3622, CRI 1142.
Chelonoidis
C. abingdoni: USMN 222479. C. carbonaria: ICN 7619, ICN 7644, ICN
1733. C. elephantopus: USMN 015192, USMN 59867, USMN 284682,
USMN 284685, USMN 284693, USMN 284694. C. gigantea: USMN
222495, USMN 107644.
Journal of Paleontology pleo-86-03-13.3d 21/2/12 10:40:23 556 Cust # 11-106R
APPENDIX 2—Rhinoclemmys measurements, Peter Pritchard Collection, Oviedo, FL, July 2009.
Species
Epiplastron measurements (mm)
Midline gular width Anterior gular width Posterior gular width Sex
Rhinoclemmys funerea CRI 4380 29.5 30.1 8.9 F
Rhinoclemmys funerea CRI 2961 26.3 28.3 7.4 M
Rhinoclemmys funerea CRI 5494 25.6 30 4.3 F
Rhinoclemmys funerea CRI 4579 18.4 23.7 6.7 F
Rhinoclemmys funerea CRI 0348 27.4 27 6.6 F
Rhinoclemmys funerea CRI 4578 27 24.1 7.8 F
Rhinoclemmys funerea CRI 5250 21 33.1 7.9 F
Rhinoclemmys funerea CRI 7699 21.7 25 8.2 F
Rhinoclemmys funerea CRI 1730 27.7 26.4 10.8 M
Rhinoclemmys punctularia CRI 2877 23 16.5 0 F
Rhinoclemmys punctularia CRI 2871 16.9 17.2 2 F
Rhinoclemmys punctularia CRI 3706 22 22.2 0 F
Rhinoclemmys punctularia CRI 2514 17.1 20.5 3.2 F
Rhinoclemmys punctularia CRI 0345 12.8 16.5 2.1 F
Rhinoclemmys punctularia CRI 4581 16.3 17.6 0 F
Rhinoclemmys punctularia CRI 3191 18.2 17 0 F
Rhinoclemmys punctularia CRI 0296 17 16.5 0 F
Rhinoclemmys punctularia CRI 0796 19.5 17 0 F
Rhinoclemmys punctularia CRI 1813 17.2 15 0 F
Rhinoclemmys punctularia CRI 2702 15.4 16.6 1.2 F
Rhinoclemmys punctularia CRI 5481 10.1 11.5 1.1 F
Rhinoclemmys nasuta CRI 2443 11.9 16.3 2 F
Rhinoclemmys nasuta CRI 2638 13 20.6 3 M
Rhinoclemmys nasuta CRI 2448 12.7 17.1 4 M
Rhinoclemmys nasuta CRI 2442 11 15.4 3.1 F
Rhinoclemmys annulata CRI 2962 18 20.5 6 M
Rhinoclemmys annulata CRI 0049 11.8 16 4.5 F
Rhinoclemmys annulata CRI 0048 12 17.1 4.3 M
Rhinoclemmys annulata CRI 3638 11 20 5 F
Rhinoclemmys annulata CRI 1132 11.7 12.7 3 M
Rhinoclemmys annulata CRI 2774 12.1 19.4 4 M
Rhinoclemmys annulata CRI 1222 16 20 2.5 F
Rhinoclemmys annulata CRI 2773 7.5 11.8 3 F
Rhinoclemmys pulcherrima manni CRI 2155 15 20 2 M
Rhinoclemmys pulcherrima manni CRI 1839 16 21 5 M
Rhinoclemmys pulcherrima manni CRI 1483 17.6 21.4 5 F
Rhinoclemmys pulcherrima manni CRI 1090 11 12.8 3.5 F
Rhinoclemmys pulcherrima manni CRI 7597 13 15.5 4 M
Rhinoclemmys pulcherrima manni CRI 1444 15.5 20.7 6 F
Rhinoclemmys pulcherrima manni CRI 2895 14 13.5 4 M
Rhinoclemmys pulcherrima manni CRI 6625 14.7 18 3 M
Rhinoclemmys pulcherrima manni CRI 1478 14.9 15.4 2.1 F
Rhinoclemmys areolata CRI 8318 13.5 16.2 4.4 F
Rhinoclemmys areolata CRI 2775 14.2 13.1 3 F
Rhinoclemmys areolata CRI 2963 13.8 12.5 1 M
Rhinoclemmys melanosterna CRI 2441 22.6 25 7.5 F
Rhinoclemmys melanosterna CRI 4198 17.6 20.3 5 F
Rhinoclemmys melanosterna CRI 2434 16.2 19.4 3 F
556 JOURNAL OF PALEONTOLOGY, V. 86, NO. 3, 2012

Journal of Paleontology pleo-86-03-13.3d 21/2/12 10:40:24 557 Cust # 11-106R
Species
Epiplastron measurements (mm)
Midline gular width Anterior gular width Posterior gular width Sex
Rhinoclemmys melanosterna CRI 2433 13.5 15 2 F
Rhinoclemmys rubida CRI 1234 13.5 10.8 0 F
Rhinoclemmys diademata CRI 1339 30 27.2 6 F
Rhinoclemmys diademata CRI 3656 22.3 20.7 2 F
Rhinoclemmys diademata CRI 3826 14.6 18.2 4 F
Rhinoclemmys diademata CRI 1517 15 22 4 F
Rhinoclemmys diademata CRI 1476 12.1 14.2 3 F
Rhinoclemmys diademata CRI 1516 20 23 5 F
Rhinoclemmys panamaensis 24.22 37.83 3.14 .
APPENDIX 2—Continued.
CADENA ET AL.—EOCENE–MIOCENE TURTLES FROM PANAMA 557