# A South American sebecid from the Miocene of Hispaniola documents the presence of apex predators in early West Indies ecosystems

Lázaro Viñola López, Jorge Vélez-juarbe, Philippe Munch, Juan N. Almonte

Milan, Pierre-Olivier Antoine, Laurent Marivaux, Osvaldo Jimenez-Vasquez,

Jonathan I. Bloch

► To cite this version:

Lázaro Viñola López, Jorge Vélez-juarbe, Philippe Munch, Juan N. Almonte Milan, Pierre-Olivier Antoine, et al.. A South American sebecid from the Miocene of Hispaniola documents the presence of apex predators in early West Indies ecosystems. Proceedings of the Royal Society of London. Series B, Containing papers of a Biological character. Royal Society (Great Britain), 2025, 292 (2045), pp.20242891. 10.1098/rspb.2024.2891. hal-05046395

HAL Id: hal-05046395

https://hal.science/hal-05046395v1

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Distributed under a Creative Commons Attribution 4.0 International License
A South American sebecid from the Miocene of Hispaniola
documents the presence of apex predators in early West Indies
ecosystems

Lazaro W. Viñola-Lopez<fnref n="1" />,<fnref n="2" />,* , Jorge Vélez-Juarbe<fnref n="3" />, Philippe Münch<fnref n="4" />, Juan N. Almonte Milan<fnref n="5" />,
Pierre-Olivier Antoine<fnref n="6" />, Laurent Marivaux<fnref n="6" />, Osvaldo Jimenez-Vasquez<fnref n="7" />, Jonathan I. Bloch<fnref n="1" />

Keywords: Caribbean | extinction | Sebecus | paleobiogeography | biodiversity museum

<footnote n="1">¹ Florida Museum of Natural History, University of Florida, P.O. Box 117800, Gainesville, FL 32611, USA</footnote>

<footnote n="2">² Negaunee Integrative Research Center, Field Museum of Natural History, Chicago, IL, USA</footnote>

<footnote n="3">³ Department of Mammalogy, Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, CA 90007, USA</footnote>

<footnote n="4">⁴ Géosciences Montpellier (UMR 5243, CNRS/UM/Université des Antilles), c.c. 060, Université de Montpellier (UM), Place Eugène Bataillon, F-34095 Montpellier Cedex 05, France</footnote>

<footnote n="5">⁵ Museo Nacional de Historia Natural “Prof. Eugenio de Jesús Marcano”, Santo Domingo, Dominican Republic</footnote>

<footnote n="6">⁶ Institut des Sciences de l'Évolution de Montpellier, Université de Montpellier (UM), CNRS, IRD, cc64, Place Eugène Bataillon, F-34095 Montpellier Cedex 05, France</footnote>

<footnote n="7">⁷ Gabinete de Arqueología de la Oficina del Historiador de La Habana, Cuba</footnote>

<footnote>*Lazaro W. Viñola-Lopez</footnote>

<footnote>Email: lwvl94@gmail.com</footnote>
1 A South American sebecid from the Miocene of Hispaniola

2 documents the presence of apex predators in early West Indies

3 ecosystems

4

5 Lazaro W. Viñola-Lopez¹,², Jorge Vélez-Juarbe³, Philippe Münch⁴, Juan N. Almonte Milan⁵,

6 Pierre-Olivier Antoine⁶, Laurent Marivaux⁶, Osvaldo Jimenez-Vasquez⁷, Jonathan I. Bloch¹

7

8 ¹Florida Museum of Natural History, University of Florida, P.O. Box 117800, Gainesville, FL

9 32611, USA

10 ²Negaunee Integrative Research Center, Field Museum of Natural History, Chicago, IL, USA

11 ³Department of Mammalogy, Natural History Museum of Los Angeles County, 900 Exposition

12 Blvd., Los Angeles, CA 90007, USA

13 ⁴Géosciences Montpellier (UMR 5243, CNRS/UM/Université des Antilles), c.c. 060, Université

14 de Montpellier (UM), Place Eugène Bataillon, F-34095 Montpellier Cedex 05, France

15 ⁵Museo Nacional de Historia Natural “Prof. Eugenio de Jesús Marcano”, Santo Domingo,

16 Dominican Republic

17 ⁶Institut des Sciences de l'Évolution de Montpellier, Université de Montpellier (UM), CNRS, IRD,

18 cc64, Place Eugène Bataillon, F-34095 Montpellier Cedex 05, France

19 ⁷Gabinete de Arqueología de la Oficina del Historiador de La Habana, Cuba

20 *Lazaro W. Viñola-Lopez

21 Email: lwvl94@gmail.com

22

23 Keywords: Caribbean | extinction | *Sebecus* | paleobiogeography | biodiversity museum

24

25 Abstract

26 Absence of terrestrial apex predators on oceanic islands lead to the evolution of endemic

27 secondary apex predators like birds, snakes, and crocodiles and loss of defense mechanisms

28 among species. These patterns are well documented in modern and Quaternary terrestrial

29 communities of the West Indies, suggesting that biodiversity there assembled similarly through

30 overwater dispersal. Here, we describe fossils of a terrestrial apex predator, a sebecid

31 crocodyliform with South American origins from the late Neogene of Hispaniola that challenge this

32 scenario. These fossils, along with other putative sebecid specimens from Cuba and Puerto Rico,
show that deep-time Caribbean ecosystems more closely resembled coeval localities in South
American than those of today. We argue that Plio-Pleistocene extinction of apex predators in the
West Indies resulted in mesopredator release and other evolutionary patterns traditionally
observed on oceanic islands. Adaptations to a terrestrial lifestyle documented for sebecids and
the chronology of West Indian fossils strongly suggest that they reached the islands in the
Eocene-Oligocene through transient land connections with South America or island hopping.
Furthermore, sebecids persisted in the West Indies for at least five million years after their
extinction in South America, preserving the last populations of notosuchians yet recovered from
the fossil record.

42

# 1. Introduction

44

The West Indies had a high diversity of endemic terrestrial vertebrates during the late Quaternary, resulting from millions of years of dispersal events (largely from South America) and subsequent in situ evolutionary radiations across the Cenozoic. The mode and tempo of vertebrate colonizations have been debated for over a century (1-2). Some studies argue that the Core Greater Antilles (Cuba-Hispaniola-Puerto Rico, CGA) were colonized via a short-lived land connection or string of islands known as GAARlandia around the Eocene/Oligocene transition (EOT) (e.g., 3-8). Geologic reconstructions of GAARlandia suggest that it coincided with a drop in eustatic sea level and the uplift of the Aves Ridge at the EOT, connecting part of the CGA with northern South America for nearly two million years (4). Results from recent geotectonic studies in the Lesser Antilles suggest that large islands in this area were exposed several times between the Eocene and the Pliocene, and that the distance between Puerto Rico and northern South America was significantly shorter than previously estimated, potentially facilitating dispersals from the continent to the islands (9-11). Meanwhile, other studies argue that the island communities, which lack carnivores, marsupials, ungulates, and several clades of frogs and reptiles, were assembled as a result of colonization via long-distance overwater dispersal (12-17). The absence of specialized terrestrial carnivores on islands has long-lasting ecological effects that ripple down through terrestrial ecosystems, often associated with mesopredator release and secondary flightlessness among birds (18-20). These effects are well documented in modern and late Quaternary communities in the Caribbean, where crocodiles and birds acted as top predators with secondary adaptations to a more terrestrial lifestyle (21-23), and numerous clades of birds evolved flightlessness and terrestrial specializations independently (e.g., 21, 24-26).

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In the late Eocene-early Oligocene, the time interval proposed for the GAARlandia landspan, the terrestrial carnivore guild in South America included metatherian mammals (Sparassodonta), large snakes (Madtsoiidae), large birds (Phorusrhacidae), and notosuchian crocodyliforms (Sebecidae) (27-28). Sebecids had distinctive labio-lingually compressed and serrated
(ziphodont) teeth and postcranial adaptations to terrestriality. They inhabited Europe in the late
Cretaceous and South America through most of the Cenozoic until the early Late Miocene (28-
30). Fossil ziphodont crocodyliform teeth have previously been recovered from the Early Miocene
of Cuba (22), but due to the independent evolution of this dental morphology in notosuchians and
at least two other neosuchian clades, taxonomic attribution to a specific group has been lacking
(31). Here, we report the first unambiguous record of a sebecid outside of South America during
the Cenozoic and argue that these South American terrestrial apex predators were likely a
dominated part of the food web in the West Indies for much of the Neogene.

78
79

**2. Materials and Methods**

81

82 (a) Institutional abbreviations.

83 FLMNH, Florida Museum of Natural History, Gainesville, FL, USA; LACM, Vertebrate
84 Paleontology Collection, Natural History Museum of Los Angeles County, Los Angeles, CA, USA;
85 MNHNCu, Paleontology collection, Museo Nacional de Historia Natural de Cuba, La Habana,
86 Cuba; MNHNSD, Museo Nacional de Historia Natural "Prof. Eugenio de Jesús Marcano", Santo
87 Domingo, Dominican Republic; UM, Museum of Paleontology, University of Michigan, Ann Arbor,
88 IL, USA.

89

90 (b) Specimens observed.

91 *Boverisuchus vorax* (LACM 21260-64 56184-86; FMNH PR399, PR479; UM uncat.); *Sebecus
92 *icaeorhinus* (FMNH PR 861 cast of holotype); *Sebecidae* ( FMNH PR 2828, PR 5109; LACM
93 uncat); *Borealosuchus wilsoni* (FMNH PR 1674); *Alligator mississippiensis* (FLMNH 39106);
94 *Crocodylus rhombifer* (MNHNSD FOS 23.325); *Crocodylus acutus* (MNHNSD); *Gavialidae*
95 (MNHNSD FOS 23.003; 23.005; 23.1029, 23.1030).

96

97 (c) Micro-Computed Tomography Scanning and imaging.

98 We produced high-resolution X-ray computed tomography (micro-CT) scans for two vertebrae
99 (MNHNSD FOS 23.1323, 23.1324) and three teeth (MNHNSD FOS 23.1325, MNHNCu P3035,
100 P3115) using the Phoenix v|tome|x M dual tube system at the University of Florida's Nanoscale
101 Research Facility. Scanning was carried out using a 240 kV x-ray tube and a tungsten target at
102 100 kV and 200 mA with an exposure time of 200 ms, a composite average of three images per
103 view, and a 1mm copper filter. The resulting x-ray data were converted into volumetric data using
104 GE's proprietary datos|x software and reconstructed in three dimensions using VG Studio Max
105 4.1.2023. High-resolution X-ray computed tomography (CT) scan for LACM 162454 was
106 performed using a Bruker Skyscan 1273. Scanning was executed using a 130 kV source, an
107 exposure time of 592 ms, and a 1 mm aluminum filter. Digital images of the sebecid and
108 planocraniid teeth were taken with Keyence VHX-7000 Digital Microscope using a Zs-20 lens.

109

## 3. Systematic Paleontology

111

112 Crocodylomorpha Walker, 1970; Crocodyliformes Hay, 1930; Notosuchia Gasparini, 1971 (*sensu*
113 Ruiz et al., 2021); Sebecosuchia Simpson, 1937 (*sensu* Leardi et al., 2024); Sebecidae Simpson,
114 1937 (*sensu* Leardi et al., 2024); *Sebecus* Simpson, 1937

115

116 cf. *Sebecus* sp.

117

118 (a) **Material.** Third(?) cervical (MNHNSD FOS 23.1323) and anterior caudal (MNHNSD FOS
119 23.1324) vertebrae of adult specimens and an isolated tooth (MNHNSD FOS 23.1325).

120

121 (b) **Locality and age.** The specimens described here were collected in the locality Paleo Pond 1,
122 on km 57 along the Juan Pablo II (RD7) highway on Sabana Grande de Boya, Dominican
123 Republic (Figs. S1-2). On this roadcut, sediments late Miocene-early Pliocene in age (7.14–4.57
124 Ma) of the Yanigua/Los Haitises Formation crop out (Core, 2015; Viñola-Lopez et al., 2022) (See
125 electronic supplementary material for detailed biostratigraphic context).

126

127 (c) **Description and comparison.** The cervical vertebra (MNHNSD FOS 23.1323) preserves
128 most of its structures, but the surface cortical bone is highly fractured and eroded, obscuring
129 some characters (Figs. 1d, S3a). In some instances, the surface is completely eroded away,
130 leaving a carbonate internal mold that outlines the general anatomy. The centrum is
131 amphicoelous, anteroposteriorly longer than high or wide, and the posterior articular surface area
132 is larger than the anterior surface area. The centrum has a rectangular outline in lateral view, and
133 the anterior and posterior articular surfaces are vertical and shallowly concave (nearly
134 approaching an amphiplatyan condition) (32-37) with the anterior articular surface being higher
135 than wide (anterior centrum height: 29 mm; anterior centrum width: 26 mm) (Fig. 1d, S3a). The
136 ventral surface of the centrum has a low median keel. The diapophysis is large, projects
137 ventrolaterally, and is located on the anterior half of the vertebra resembling the condition
138 observed in the third cervical of *Sebecus icaeorhinus* and *Sahitisuchus fluminensis* (37; Kellner et
139 al., 2013REF [38]), whereas in other basal mesoeucrocodylians (e.g., *Simosuchus clarki*,
140 *Yacarerani boliviensis*), it is placed closer to the anterior margin (34, 39). The general outline of
141 the parapophysis is preserved, from which it can be inferred that it was restricted to the anterior
142 half of the vertebral centrum and that its main axis was oriented longitudinally and nearly
143 horizontal, as in *S. icaeorhinus* and other mesoeucrocodylians (e.g., *S. clarki*, *Y. boliviensis*) (Fig.
144 1d, S3a). The diapophysis and parapophysis are separated by a deep longitudinal groove on the lateral side of the centrum as in *S. icaeorhinus* (Pol et al., 2012). The region where the hypapophysis would be located is eroded and missing in the specimens. The dorsal surface of the centrum has a relatively deep longitudinal groove along the midline. The neural canal is oriented anterodorsally to the horizontal plane. The anterior opening of the neural canal is nearly circular (17 mm wide, 12 mm high), contrasting with the tall and oval shape of the posterior opening (14 mm wide, 18 mm high) (S3a).

151 Although most of the neural spine is missing, the base of the spine is posteriorly located, as in *Sebecus*, and unlike the centrally-located spine of *Notosuchus* (35-36) (Fig. 1d, S3a). The base of the spine is anteroposteriorly short (15 mm long; about one-third the length of the vertebra) as in *Sebecus*, *Notosuchus*, *Baurusuchus*, and *Simosuchus* (34, 37). On the neural arch, anterior to the spine, there is a prespinal fossa resembling the condition in some mesoeucrocodylians (e.g., *Mahajangasuchus insignis*, *Baurusuchus albertoi*) and *Sebecus icaeorhinus*, although in MNHNSD FOS 23.1323, it is deeper and more developed as in the latter taxon (37, 40-41) (S3a). This fossa is located on the midpoint between the spine and the prezygapophyses, and there is a longitudinal canal leading to it that starts at the anterodorsal margin of the arch between the prezygapophyses. The prezygapophyses are projected dorsally, as in most mesoeucrocodylians (e.g., *Mahajangasuchus*, *Simosuchus*, *Notosuchus*, *Baurusuchus*) and *Sebecus* (34, 37, 40-41), and the anterior margin of the prezygapophyseal process has a small anteriorly oriented ridge. This anterior ridge resembles the prezygapophyseal bulge, resembling the condition observed in *Sebecus* and *Notosuchus*, but differs from other crocodylians with a convex process (37). It further resembles *Sebecus* in that the articular facets of the prezygapophyses are flat, subcircular, and oriented 45 degrees with respect to the main axis of the vertebra (Figs. 1d, S3a).

168 The anterior caudal vertebra (MNHNSD FOS 23.1324) is nearly complete and well preserved (Figs. 1b, S3b). The neural spine is a long rectangular and medio-laterally compressed blade slightly inclined posteriorly, its base is anteroposteriorly broad (16.53 mm), and its width decreases apically. The neural spine is located on the postero-medial region of the neural arch, a condition shared with *Sebecus huilensis*, *Yacarerani boliviensis*, *Mahajangasuchus insignis*, *Campinasuchus dinizi*, and *baurusuchids* (33, 42). The prezygapophyses are widely separated, projecting anterodorsally with the articular surface oriented dorsomedially. There is a deep prespinal fossa at the midline of the vertebra, between the prezygapophysis and anterior to the base of the neural spine. The postzygapophyses are short, project ventrolaterally, and are much more inclined than the prezygapophysis (Fig. S3b). The transverse processes are oval in cross-section as in *Sebecus huilensis*, whereas they are smaller and dorsoventrally compressed in the advanced notosuchians *Marialiasuchus* and *Caipirasuchus* (33, 43-44). The transverse
processes are located just ventral to the level of the pre- and post-zygapophyses, and project posterolaterally and slightly dorsal relative to the main axis of the vertebra (Figs. 1b, S3b). On each side of the dorsal surface of the vertebrae, between the base of the transverse processes and the neural spine, there is a depression, which is also found in *Notosuchus terrestris* (36). The neural canal is large, oval in cross-section anteriorly, and nearly circular posteriorly (Fig. S3b). The neural arch is fused to the centrum, thereby suggesting the specimen reached skeletal maturity (Fig. 1b).

In lateral view, the centrum is elongated and tall, and its lateral walls are excavated (Fig. 1b, S3b). The centrum is amphicoelous and longer (24.14 mm) than wide (9.88 mm). In lateral view, the anterior articular surface has a circular outline and is anteroventrally inclined, whereas the posterior surface is subrectangular and slightly posterodorsally oriented. The ventral margin of the centrum forms an arch, with the posterior region of the centrum projecting more ventrally than the anterior one, resembling the prominent hemapophyses in the anterior caudals of *Simosuchus clarki*, *Mahajangasuchus insignis* and *Baurusuchus albertoi* (34; 40-41). The hemapophysis is inclined posteroventrally like in other sebecids, mesoeucrocodylians (*Mahajangasuchus*), sphagesaurians (e.g., *Mariliasuchus*), and baurusuchids (*Campinasuchus*) (30, 33, 40-43, 45). In ventral view, the midsection of the centrum is mediolaterally compressed, whereas the anterior and posterior surfaces expand laterally. The ventral surface of the centrum is excavated at the midline, forming two low parasagittal ridges.

MNHNSD FOS 23.1325 is the upper portion of a labiolingually flattened tooth (Fig. S4d). The preserved portion of the tooth is ~12 mm high; its base has a mesiodistal length of 8.11 mm and a labiolingual width of 3.55 mm (Fig. S4d, S5b). As such, the length/width ratio of 2.3 is similar to that observed in *Sebecus* (46). The lingual and labial surfaces are convex and smooth. In lingual and labial views, the tooth is nearly symmetrical along its longitudinal axis. The mesial and distal surfaces are narrow and bear a well-defined carinae with true serrations formed by clefts in the enamel (Fig. S5b). The denticles are rectangular to convex, and the ones located in the middle portion of the teeth are approximately 0.2 mm in length, with a density of ~5 denticles per mm.

(d) **Other ziphodont teeth.** Three other isolated teeth were collected from the early Miocene Domo de Zaza locality in Cuba (Museo Nacional de Historia Natural de Cuba, MNHNCu P3035, P3115) and the early Oligocene Yauco locality in Puerto Rico (Los Angeles County Museum, LACM 162454) (Figs. S1, S4-S5). The three teeth are labiolingually compressed and slightly recurved mesially and lingually. The carinae are worn to different degrees but still show well-defined serrations formed of rectangular to convex denticles with a density of ~5 denticles per
215 mm (See electronic supplementary material for detailed description of specimens and
216 biostratigraphic data).

217
218 **4. Discussion**
219

220 (a) Sebecids and ziphodont crocodyliforms in the West Indies.

221 The oldest ziphodont crocodyliform recorded in the West Indies corresponds to an isolated tooth
222 from the middle Eocene Seven Rivers locality in Jamaica (47). During the middle Eocene,
223 Western Jamaica was connected to the Nicaragua Rise, a southern extension of North America.

224 Vertebrates thus far recovered from this locality have affinities with the fauna from similar age
225 deposits in North America (48-49). Given the biogeographical affinities of fossils from Jamaica,
226 the ziphodont tooth from Seven Rivers most likely represents a planocraniid rather than a sebecid
227 (31, 50).

228 Other ziphodont crocodyliform teeth in the West Indies come from Domo de Zaza, a
229 Lower Miocene (17.5-18.5 Mya) locality of the Lagunitas Formation in central Cuba (22, 31).

230 Because at least three groups of Cenozoic crocodyliforms, including sebecosuchian,
231 planocraniids, and mekosuchines, independently evolved ziphodont dentition, assigning the
232 Cuban isolated teeth to a specific group was initially problematic. Affinities with Mekosuchinae are
233 unlikely because all members of this clade are part of a mostly endemic radiation from the
234 Eocene-Pleistocene of Australia (51-53), and only a later Holocene dispersal into the Pacific
235 Islands (54). Meanwhile, a close relationship with planiocraniids was not entirely discarded as this
236 clade inhabited North America and Eurasia, but the group became extinct in the Eocene (50).

237 Sebecosuchians are the other known clade in the Americas with ziphodont teeth, but their New
238 World fossil record was, until now, restricted to South America in the Cenozoic (28, 31).

239 Nonetheless, the amphicoelous vertebrae and the ziphodont tooth from the Late Miocene-Early
240 Pliocene of the Dominican Republic show that sebecids inhabited the West Indies. Therefore, it is
241 likely that the Early Miocene Cuban specimens as well as the early Oligocene ziphodont tooth
242 from Puerto Rico also belong to insular sebecids (see electronic supplementary material for
243 specimens description).

244 In South America, the Paleogene records of Sebecidae are best documented at high
245 latitudes, but isolated specimens from Eocene deposits at low latitudes revealed that the family
246 had a pancontinental distribution (28, 45) (Fig. 2). Their range in the Miocene was restricted to
247 the more tropical zones at mid and low latitudes, consistent with other groups of terrestrial
248 vertebrates in South America (28, 45). How sebecids colonized the Greater Antilles is unclear,
249 but the source population was likely in northern South America. Considering their terrestrial
250 adaptations, their dispersal may have (i) been either facilitated by some ephemeral terrestrial
251 connection or string of large and closely spaced islands or (ii) occurred on a natural raft. The
252 fossil from the early Oligocene of Puerto Rico is consistent with the age and location where it
253 would be expected to find a South American carnivore that dispersed via GAARlandia or the
254 Lesser Antilles around the Eocene-Oligocene transition. Once in the CGA, the founding
255 population would have become isolated as islands separated with the opening of the Mona
256 passage in the Oligocene and the Mind passage in the Early Miocene (6, 55). This is consistent
257 with their fossil record on the island, limited so far to the CGA. Similar temporal and spatial
258 colonization patterns have been documented for other clades with South American affinities in the
259 West Indies, like sloths, caviomorph rodents (chinchiloids), and anurans (7-8, 56-58).
260

261 (b) The “missing” apex carnivores and their asynchronous extinction

262 Absence of native terrestrial carnivores in the late Quaternary fossil record and modern
263 ecosystems in the West Indies has been cited as evidence against biogeographical hypotheses
264 that propose the existence of a land connection with the mainland. This is not only reflected in the
265 lack of taxa from the well-documented Quaternary fossil record in the region, but also in the
266 endemic radiation of large nocturnal and diurnal raptorial birds that filled this ecological niche (21,
267 26, 59) along with *Crocodylus rhombifer*, one of the most terrestrially adapted crocodylians alive
268 today (22, 60). These predators were at the top of the food chain in the CGA, which included
269 more than 51 land mammal species (sloths, rodents, primates, and island shrews), along with
270 numerous other now extinct reptiles and birds. Fossils of these species are often found in
271 predator roosts showing evidence of bitemarks and digestion (22-23, 26). Furthermore, several
272 clades of birds, including rails, cave rails, cranes, ibis, as well as predatory owls, hawks, and
273 falcons, evolved adaptations to a terrestrial lifestyle and, in some cases, secondary flightlessness
274 likely related to the lack of terrestrial predators (24, 26). These adaptations are likely fairly recent,
275 following the disappearance of sebecids in the West Indies in post-Miocene times (Fig. 2). As it
276 has been documented on other islands, extinction and occasional dispersal can change the
277 structure and composition of an insular fauna originally assembled via vicariance into resembling
278 a truly oceanic one (61).

279 In South America, these notosuchians were apex predators adapted to terrestrial
280 ecosystems. The postcranial morphology of *Sebecus icaeorhinus* shows several adaptations to
281 terrestriality with erect limb posture (35), while the neuroanatomy and cranial morphology of
282 several species further support this habitat preference and suggest that prey acquisition and
283 manipulation likely occurred on land (62-64). Results from multiproxy isotope analyses (δ¹³C, δ¹⁸O, δ⁴³/⁴²Ca) of tooth enamel derived from Paleocene fossils from Bolivia suggest that sebecids
284 occupied a top position on the C3-based food web in a local dry tropical environment (65).
285 Sebecids in the West Indies were likely also apex predators in terrestrial ecosystems for more
286 than 22 million years. Their remains have been found in association with fossils of sloths and
287
288 other terrestrial (e.g., rodents, primates) or semiaquatic vertebrates (e.g., pelomedusoid turtles;
289 electronic supplementary material, Table S1).

290 The last unequivocal records of Sebecidae from South America are from the early Late
291 Miocene, including *Sebecus huilensis* (Peru and Colombia), *Barinasuchus arveloi* (Venezuela
292 and Peru), and an undescribed large form known from the Villavieja Formation (12.58–10.52
293 Mya) in La Venta, Colombia (28, 66-70). The geographic range construction and subsequent
294 extinction of sebecids has been associated with the progressive retreat of tropical climate to lower
295 latitudes in South America through the Cenozoic, and habitat changes in the earliest Late
296 Miocene caused by the disintegration of the Pebas wetland system (28, 45). In the West Indies,
297 the last occurrence of sebecids dates from the late Miocene–early Pliocene (7.14–4.57 Mya) (71),
298 at least 5 Ma later than on the mainland (28, 69). The Hispaniolan fossils described here
299 document not only the last known sebecid, but also the last appearance of the clade Notosuchia.
300 This highly diverse group of crocodyliforms with a wide array of ecological and morphological
301 specialization had a nearly global distribution in the Cretaceous, of which only Sebecosuchia
302 survived the Cretaceous/Paleogene boundary in Europe and South America 72-73).

303 Islands (physical and ecological) around the world often act as biodiversity museums,
304 hosting relics of old lineages that once had a broad distribution but that today are threatened by
305 extinction (74-75). These lineages survived and, in some instances, diversified in isolation on the
306 islands for millions of years while the remaining populations on the mainland disappeared (74).
307 The West Indies are well known as the last refugium of numerous lineages of plants,
308 invertebrates, and vertebrates threatened by extinction or extinct elsewhere (e.g., 76-79). The
309 isolation of sebecids on the islands buffered them from macroecological changes recorded in
310 South America, allowing them to survive longer than their relatives on the mainland. Moreover,
311 the fauna associated with these last sebecids included other crocodyliforms (alligatoroids and
312 gryposuchine gavialoids), pelomedusoid turtles, caviomorph rodents, and megalocnid sloths (71,
313 electronic supplementary material Table S1), suggesting that the Cenozoic Greater Antillean
314 vertebrate fauna more closely resembled that of the South American mainland than previously
315 recognized, with multiple lineages of crocodyliforms as macropredators (80). The relatively recent
316 decline and demise of these multi-species crocodyliform assemblages likely paved the way for
317 the origination of late Quaternary and modern-day trophic structures in the West Indies.
318

319 **Data accessibility.** All the data used in this study is available on the manuscript and the
320 electronic supplementary material.

321 **Acknowledgments.** We want to extend our gratitude to our colleagues Evelin Gabot, America
322 Sánchez, Arturo Leon at the Museo Nacional de Historia Natural "Prof. Eugenio de Jesus
323 Marcano" in the Dominican Republic who participated in the field seasons collecting fossils.
Special thanks to Hernán Santos, Wilson Ramírez (University of Puerto Rico, Mayagüez Campus), and Elson Core for their earlier efforts collecting and disentangling the geologic setting of the Yanigua/Los Haitises section. Many thanks to Stephanie Abramowicz (NHMLAC) for her assistance scanning LACM 162454, and Niko Vuckovich for the Keyence images of LACM 162454. Our gratitude to William Simpson and Rachel Narducci for access to specimens under their care at the Field Museum of Natural History and the Florida Museum of Natural History.

Thanks to Yasmani Ceballos and Kenneth Angielczyk for their comments and suggestions on the early versions of the manuscript and to Mitchell Riegler for discussions on character evolution of ziphodont crocodilians. The manuscript was enhanced thanks to the revisions of the editor Locke Rowe and two anonymous reviewers.

334

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<figure><img src="image_2.png" /><figcaption>Figure 1. Generalized relationship of crocodilians that independently evolved ziphodont dentition and the general anatomy of nuchal and caudal vertebrae in *Notosuchia* and *Eusuchia* (A). Comparison of caudal (MNHNSD FOS 23.1324) (B) and nuchal (MNHNSD FOS 23.1323) (D) vertebrae of *Sebecus* sp. from the late Miocene-early Pliocene of Dominican Republic (Scale = 15 mm). *Sebecid* teeth from the early Miocene of Cuba (MNHNCu P3115) (D, Scale = 10 mm) and late Miocene-early Pliocene of Dominican Republic (MNHNSD FOS 23.1325) (E, Scale = 15 mm).</figcaption></figure>

166x191mm (300 x 300 DPI)
<figure><img src="image_3.png" /><figcaption>Figure 2. Maps of South America and the Caribbean region during the Paleogene (A), Neogene (B), and late Quaternary (C) show the generalized distribution of Sebecidae in circles. The silhouettes correspond to the terrestrial apex predator groups present in the region during each period, with native South American predators (Sebecidae, Madtsoiidae, Phorusrhacidae, Sparassodonta) in blue, late Cenozoic invasive predators (Canidae, Felidae, Ursidae) in red, and endemic secondary terrestrial predators in the West Indies (Strigiformes, Accipitridae, Crocodylus) in green. Localities of Sebecidae based on electronic supplementary material Table S2.</figcaption></figure>