Isotopic and anatomical evidence of an herbivorous diet in the Early Tertiary giant bird Gastornis. Implications for the structure of Paleocene terrestrial ecosystems

The mode of life of the early Tertiary giant bird Gastornis has long been a matter of controversy. Although it has often been reconstructed as an apex predator feeding on small mammals, according to other interpretations, it was in fact a large herbivore. To determine the diet of this bird, we analyze here the carbon isotope composition of the bone apatite from Gastornis and contemporaneous herbivorous mammals. Based on ¹³C-enrichment measured between carbonate and diet of carnivorous and herbivorous modern birds, the carbonate δ¹³C values of Gastornis bone remains, recovered from four Paleocene and Eocene French localities, indicate that this bird fed on plants. This is confirmed by a morphofunctional study showing that the reconstructed jaw musculature of Gastornis was similar to that of living herbivorous birds and unlike that of carnivorous forms. The herbivorous Gastornis was the largest terrestrial tetrapod in the Paleocene biota of Europe, unlike the situation in North America and Asia, where Gastornis is first recorded in the early Eocene, and the largest Paleocene animals were herbivorous mammals. The structure of the Paleocene terrestrial ecosystems of Europe may have been similar to that of some large islands, notably Madagascar, prior to the arrival of humans.     

Earliest and first Northern Hemispheric hoatzin fossils substantiate Old World origin of a “Neotropic endemic”

The recent identification of hoatzins (Opisthocomiformes) in the Miocene of Africa showed part of the evolution of these birds, which are now only found in South America, to have taken place outside the Neotropic region. Here, we describe a new fossil species from the late Eocene of France, which constitutes the earliest fossil record of hoatzins and the first one from the Northern Hemisphere. Protoazin parisiensis gen. et sp. nov. is more closely related to South American Opisthocomiformes than the African taxon Namibiavis and substantiates an Old World origin of hoatzins, as well as a relictual distribution of the single extant species. Although recognition of hoatzins in Europe may challenge their presumed transatlantic dispersal, there are still no North American fossils in support of an alternative, Northern Hemispheric, dispersal route. In addition to Opisthocomiformes, other avian taxa are known from the Cenozoic of Europe, the extant representatives of which are only found in South America. Recognition of hoatzins in the early Cenozoic of Europe is of particular significance because Opisthocomiformes have a fossil record in sub-Saharan Africa, which supports the hypothesis that extinction of at least some of these “South American” groups outside the Neotropic region was not primarily due to climatic factors.     

First ceratosaurian dinosaur from Australia

The basal theropod dinosaur clade Ceratosauria, and its subclade Abelisauroidea, is characteristic of late Mesozoic terrestrial vertebrate faunas in western Gondwana (South America, Africa, Madagascar, and India) and Europe. Yet unambiguous records of ceratosaurs have hitherto been absent from Australia, where the theropod assemblage appears to include several typically Laurasian clades. Here, we report the first evidence of ceratosaurs (and potentially abelisauroids) from eastern Gondwana––a diagnostic astragalocalcaneum from the Aptian (121–125?Ma) of Victoria, Australia. Ceratosauria thus occurred in both western and eastern Gondwana during the Early Cretaceous. This fossil adds to the poorly known dinosaur fauna of Australia, a major clade of basal theropods, emphasising that its mid-Cretaceous theropod diversity was surprisingly cosmopolitan despite relative geographic isolation, including clades that have been thought to be typical of both Gondwana and Laurasia––Ceratosauria, Spinosauridae, Carcharodontosauria, Tyrannosauroidea, and Deinonychosauria. Such a contemporaneous association of theropod clades is unknown from other Gondwanan continents and questions the views that the late Mesozoic dinosaur fauna of Australia was dominated by Gondwanan or Laurasian elements, extreme isolation, relictualism, and/or novelty as a ‘centre of origin’. The cosmopolitan theropod fauna of Australia probably reflects the global distribution of these clades early in their history, prior to significant continental breakup.     

The oldest European fossil songbird from the early Oligocene of Germany

We report on the oldest European songbird (Passeriformes), from the early Oligocene (30–34 million years ago) of Frauenweiler in Germany. The specimen represents the earliest associated remains of an early Tertiary passerine described so far. It ties the first appearance of Passeriformes in Europe to a minimum age of 30 million years. Passeriform birds are absent in Eocene deposits that yielded abundant remains of small land birds and apparently dispersed into Europe around the Eocene/Oligocene boundary (about 34 million years ago), not at the Oligocene/Miocene boundary (about 24 mya) as hitherto thought. This possibly relates the appearance of songbirds in Europe to a well-known major faunistic break at the Eocene/Oligocene boundary, called the grande coupure. The Frauenweiler songbird most notably differs from extant Passeriformes in having a larger processus procoracoideus on the coracoid and appears to be outside Eupasseres, the taxon which includes Oscines (all modern European and most Old World songbirds) and Suboscines (most South and Central American songbirds). It shows that there were earlier dispersal events of non-oscine songbirds into Europe before the arrival of Oscines from the Australian continental plate towards the late Oligocene.     

An ‘ameridelphian’ marsupial from the early Eocene of Australia supports a complex model of Southern Hemisphere marsupial biogeography

Recent molecular data strongly support the monophyly of all extant Australian and New Guinean marsupials (Eomarsupialia) to the exclusion of extant South American marsupials. This, together with available geological and fossil evidence, has been used to argue that the presence of marsupials in Australia is simply the result of a single dispersal event from South America during the latest Cretaceous or Palaeocene, without subsequent dispersals between the two continents. Here, I describe an isolated ankle bone (calcaneus) of a metatherian from the early Eocene Tingamarra Local Fauna in northeastern Australia. Strikingly, this specimen, QM F30060, lacks the ‘continuous lower ankle joint pattern’ (CLAJP), presence of which is a highly distinctive apomorphy of the marsupial clade Australidelphia, which includes Eomarsupialia, the living South American microbiotherian Dromiciops and the Tingamarran fossil marsupial Djarthia. Comparisons with a range of marsupials and stem-metatherians strongly suggest that the absence of the CLAJP in QM F30060 is plesiomorphic, and that this specimen represents the first unequivocal non-australidelphian (‘ameridelphian’) metatherian known from Australia. This interpretation is confirmed by phylogenetic analyses that place QM F30060 within (crown-group) Marsupialia, but outside Australidelphia. Based on these results, the distribution of marsupials within Gondwana cannot be explained by simply a single dispersal event from South America and Australia. Either there were multiple dispersals by marsupials (and possibly also stem-metatherians) between South America and Australia, in one or both directions, or, alternatively, there was a broadly similar metatherian fauna stretching across southern South America, Antarctica and Australia during the Late Cretaceous–early Palaeogene.     

Euarchontan affinity of Paleocene Afro-European adapisoriculid mammals and their origin in the late Cretaceous Deccan Traps of India

The controversial family Adapisoriculidae, a group of shrew-sized Paleocene mammals, had proposed relationships with insectivores, marsupials and more recently to plesiadapiforms. Adapisoriculid remains are numerous in the early Paleocene locality of Hainin, Belgium, and allow us a test of these different phylogenetic hypotheses. Here, we identify the first tarsal bones of adapisoriculid mammals. The highly specialised bones indicate an arboreal mode of life with euarchontan affinity. Moreover, the tarsal bones are morphologically very close to those of the late Cretaceous Deccanolestes from the Deccan intertrappean beds of India, and also share several characters with the Paleocene plesiadapiforms and the extant cynocephalid dermopterans. The adapisoriculid affinities of Deccanolestes are also confirmed by tooth morphology, indicating that Deccanolestes is a primitive member of this family. These phylogenetic affinities suggest a paleobiogeographic scenario for the family with dispersal either via East Africa or across the Tethys area.     

A structural intermediate between triisodontids and mesonychians (Mammalia, Acreodi) from the earliest Eocene of Portugal

A new mammal, Mondegodon eutrigonus gen. et sp. nov., is described from the earliest Eocene locality of Silveirinha, Portugal. This species shows dental adaptations indicative of a carnivorous diet. M. eutrigonus is referred to the order Acreodi and considered, along with the early Paleocene North American species Oxyclaenus cuspidatus, as a morphological intermediate between two groups of ungulate-like mammals, namely, the triisodontids and mesonychians. Considering that triisodontids are early to early-late Paleocene North American taxa, Mondegodon probably belongs to a group that migrated from North America towards Europe during the first part of the Paleocene. Mondegodon could represent thus a relict genus, belonging to the ante-Eocene European mammalian fauna. The occurrence of such a taxon in Southern Europe may reflect a period of isolation of this continental area during the Paleocene/Eocene transition. In this context, the non-occurrence of closely allied forms of Mondegodon in the Eocene North European mammalian faunas is significant. This strengthens the hypothesis that the mammalian fauna from Southern Europe is characterized by a certain degree of endemism during the earliest Eocene. Mondegodon also presents some striking similarities with an unnamed genus from the early Eocene of India which could represent the first Asian known transitional form between the triisodontids and mesonychians.     

The oldest African bat from the early Eocene of El Kohol (Algeria)

The Afro-Arabian Paleogene fossil record of Chiroptera is very poor. In North Africa and Arabia, this record is limited, thus far, to a few localities mainly in Tunisia (Chambi, late early Eocene), Egypt (Fayum, late Eocene to early Oligocene), and Sultanate of Oman (Taqah, early Oligocene). It consists primarily of isolated teeth or mandible fragments. Interestingly, these African fossil bats document two modern groups (Vespertilionoidea and Rhinolophoidea) from the early Eocene, while the bat fossil record of the same epoch of North America, Eurasia, and Australia principally includes members of the ??Eochiroptera.?? This paraphyletic group contains all primitive microbats excluding modern families. In Algeria, the region of Brezina, southeast of the Atlas Mountains, is famous for the early Eocene El Kohol Formation, which has yielded one of the earliest mammalian faunas of the African landmass. Recent fieldwork in the same area has led to the discovery of a new vertebrate locality, including isolated teeth of Chiroptera. These fossils represent the oldest occurrence of Chiroptera in Africa, thus extending back the record of the group to the middle early Eocene (Ypresian) on that continent. The material consists of an upper molar and two fragments of lower molars. The dental character association matches that of ??Eochiroptera.?? As such, although very fragmentary, the material testifies to the first occurrence of ??Eochiroptera?? in Algeria, and by extension in Africa. This discovery demonstrates that this basal group of Chiroptera had a worldwide distribution during the early Paleogene.     

<Emphasis Type="BoldItalic">Osedax</Emphasis> borings in fossil marine bird bones

The bone-eating marine annelid Osedax consumes mainly whale bones on the deep-sea floor, but recent colonization experiments with cow bones and molecular age estimates suggesting a possible Cretaceous origin of Osedax indicate that this worm might be able grow on a wider range of substrates. The suggested Cretaceous origin was thought to imply that Osedax could colonize marine reptile or fish bones, but there is currently no evidence that Osedax consumes bones other than those of mammals. We provide the first evidence that Osedax was, and most likely still is, able to consume non-mammalian bones, namely bird bones. Borings resembling those produced by living Osedax were found in bones of early Oligocene marine flightless diving birds (family Plotopteridae). The species that produced these boreholes had a branching filiform root that grew to a length of at least 3 mm, and lived in densities of up to 40 individuals per square centimeter. The inclusion of bird bones into the diet of Osedax has interesting implications for the recent suggestion of a Cretaceous origin of this worm because marine birds have existed continuously since the Cretaceous. Bird bones could have enabled this worm to survive times in the Earth’s history when large marine vertebrates other than fish were rare, specifically after the disappearance of large marine reptiles at the end-Cretaceous mass extinction event and before the rise of whales in the Eocene.     

A new Dasypodini armadillo (Xenarthra: Cingulata) from San Gregorio Formation,Pliocene of Venezuela: affinities and biogeographic interpretations

We describe Pliodasypus vergelianus gen. et sp. nov., a Dasypodini armadillo from the middle Pliocene of Venezuela (Vergel Member, San Gregorio Formation). Although scarce, the remains are remarkable because of their geochronologic proximity to the main phase of Great American Biotic Interchange (GABI). The cladistic analysis conducted reveals that Pliodasypus groups with Dasypus and both are sister taxa of Propraopus, whereas Anadasypus is at a basal position. With respect to the records of tribe Dasypodini, after its oldest representative (Anadasypus, middle and late Miocene), the chronologically subsequent form is Pl. vergelianus (middle Pliocene), followed by Dasypus bellus in higher northern latitudes (late Pliocene), and then by widespread occurrences in the Pleistocene of North America (D. bellus) and South America (Propraopus, Dasypus punctatus, and Dasypus novemcinctus). Thus, we infer that Dasypus differentiated in the late Pliocene at low latitudes in the northern South America. It leads to two alternative hypotheses of dispersal: (a) some early Dasypus remained cryptically in South America until the Pleistocene, whereas others dispersed to North America between 2.2 and 2.7 Ma, or (b) they dispersed to North America subsequently to the emersion of the Panamanian isthmus and D. bellus differentiated there; later, during the Pleistocene, D. bellus entered South America and experienced speciation. The same process of re-ingression has been proposed to other xenarthrans, breaking with the traditional assumption that the GABI was unidirectional.     

Direct evidence of trophic interactions among apex predators in the Late Triassic of western North America

Hypotheses of feeding behaviors and community structure are testable with rare direct evidence of trophic interactions in the fossil record (e.g., bite marks). We present evidence of four predation, scavenging, and/or interspecific fighting events involving two large paracrocodylomorphs (=‘rauisuchians’) from the Upper Triassic Chinle Formation (～220–210 Ma). The larger femur preserves a rare history of interactions with multiple actors prior to and after death of this ～8–9-m individual. A large embedded tooth crown and punctures, all of which display reaction tissue formed through healing, record evidence of a failed attack on this individual. The second paracrocodylomorph femur exhibits four unhealed bite marks, indicating the animal either did not survive the attack or was scavenged soon after death. The combination of character states observed (e.g., morphology of the embedded tooth, ‘D’-shaped punctures, evidence of bicarination of the marking teeth, spacing of potentially serial marks) indicates that large phytosaurs were actors in both cases. Our analysis of these specimens demonstrates phytosaurs targeted large paracrocodylomorphs in these Late Triassic ecosystems. Previous distinctions between ‘aquatic’ and ‘terrestrial’ Late Triassic trophic structures were overly simplistic and built upon mistaken paleoecological assumptions; we show they were intimately connected at the highest trophic levels. Our data also support that size cannot be the sole factor in determining trophic status. Furthermore, these marks provide an opportunity to start exploring the seemingly unbalanced terrestrial ecosystems from the Late Triassic of North America, in which large carnivores far outnumber herbivores in terms of both abundance and diversity.     

Out of Africa: Fossils shed light on the origin of the hoatzin,an iconic Neotropic bird

We describe the earliest fossils of the enigmatic avian taxon Opisthocomiformes (hoatzins) from the Oligo-Miocene (22–24 mya) of Brazil. The bones, a humerus, scapula and coracoid, closely resemble those of the extant hoatzin, Opisthocomus hoazin. The very similar osteology of the pectoral girdle in the new Brazilian fossil compared to the extant O. hoazin, in which it reflects peculiar feeding adaptations, may indicate that hoatzins had already evolved their highly specialized feeding behavior by the mid-Cenozoic. We further show that Namibiavis senutae from the early Miocene of Namibia is another, previously misclassified representative of Opisthocomiformes, which documents that the extant Neotropic distribution of hoatzins is relictual. Because of the weak flight capabilities of hoatzins, their occurrence on both sides of the South Atlantic is of particular biogeographic interest. We detail that this distribution pattern is best explained by dispersal from Africa to South America, and that Opisthocomiformes provide the first example of transatlantic rafting among birds.     

Skeletal remains of a diminutive primate from the Paleocene of Germany

Most living mammal orders, including our own, started their career during the first 10 million years of the Cenozoic, the Age of Mammals. The fossil record documents that early Paleogene adaptive radiations of various clades included tiny species of the size of living shrews. Remains of particularly diminutive limb bones are described from the late Paleocene site of Walbeck, Sachsen-Anhalt. Discovered in 1939, it has remained the only known Paleocene mammal-bearing locality from Germany. The remains are referred to the family Adapisoriculidae, which is considered on the basis of the present postcranial evidence to represent plesiadapiform primates rather than alleged lipotyphlan insectivores as previously proposed. The Walbeck fossils compete with the Early Eocene species Toliapina vinealis from Europe and Picromomys petersonorum from North America for the status of the smallest known primate, fossil and living. Their estimated body weights are as small as 10 g. The limb bones show features related to enhanced flexion at the elbow and hip joint, suggesting arboreal habits and environments such as terminal branches. The diminutive size and tooth morphology suggest feeding on small insects and other invertebrates. Postcranials are important to assess early radiations, such tiny specimens as the present ones are extremely scarce in the fossil record, however.     

Comparisons of dental morphology in river stingrays (Chondrichthyes: Potamotrygonidae) with new fossils from the middle Eocene of Peruvian Amazonia rekindle debate on their evolution

Endemic South American river stingrays (Potamotrygonidae), which include the most diversified living freshwater chondrichthyans, were conspicuously absent from pre-Neogene deposits in South America despite the fact that recent phylogenetic analyses strongly suggest an older origination for this clade. To date, the rare representatives of this family were mostly represented by ambiguous isolated remains. Here, we report 67 isolated fossil teeth of a new obligate freshwater dasyatoid (Potamotrygon ucayalensis nov. sp) from the fossiliferous level CTA-27 (Yahuarango Formation), near Contamana, in the Peruvian Amazonia. We assigned this sample to a new representative of Potamotrygon by comparison with numerous fresh jaws of living specimens of Potamotrygonidae, thus providing the first detailed review of dental morphology for this poorly understood clade. These new fossils fill a long stratigraphic gap by extending the family range down to the middle Eocene (～41 Mya). Moreover, the relative modernity and diversity in tooth morphology among Eocene freshwater stingrays (including Potamotrygon ucayalensis nov. sp. and coeval North American dasyatoids) indicate that the hypothetically marine ancestor of potamotrygonids probably invaded the rivers earlier than in the middle Eocene. The first potamotrygonids and affiliates were possibly more generalized and less endemic than now, which is consistent with an opportunistic filling of vacated ecospace.     

A long-bodied centriscoid fish from the basal Eocene of Kabardino-Balkaria, northern Caucasus, Russia

The Paleocene–Eocene transition is of crucial interest for interpreting the Cenozoic evolutionary radiation of vertebrates. A substantial increase of the number of vertebrate families occurred between the Late Paleocene and Early Eocene, with the appearance of most of the representatives of extant lineages. Basal Eocene marine fish diversity is currently poorly known, exclusively restricted to two assemblages from Denmark and Turkmenistan, respectively. Exceptionally well-preserved articulated skeletal remains of fishes have recently been discovered from a basal Eocene sapropelitic layer exposed along the Kheu River in the Republic of Kabardino-Balkaria, northern Caucasus, Russia. Here, we report on Gerpegezhus paviai gen. et sp. nov., a new peculiar syngnathoid fish from this new Ciscaucasian locality. The morphological structure of the single available specimen suggests that it is the first long-bodied member of the superfamily Centriscoidea, representing the sole member of the new family Gerpegezhidae, which forms a sister pair with the extant family Centriscidae.     

The first Gondwanan borioteiioid lizard and the mid-Cretaceous dispersal event between North America and Africa

Borioteiioidea are an enigmatic group of Cretaceous lizards widely distributed in northern continents (Laurasia). Here, we describe the first borioteiioid lizard from Gondwana, represented by a new species of the polyglyphanodontine genus Bicuspidon, B. hogreli sp. nov., from the Cenomanian Kem Kem beds of Morocco. The discovery of Bicuspidon hogreli sp. nov., which is one of the oldest known member of Polyglyphanodontini, challenges previous assumptions on the center of origin and dispersal of the group. In addition, the known distribution of Bicuspidon (Cenomanian of Utah and Morocco, and Santonian–Maastrichtian of eastern Europe) suggests a complex palaeobiogeographical history for the genus. The existence of a terrestrial dispersal route persisting during the whole Early Cretaceous between North America and Africa is hypothesized to explain some similarities observed between the Cenomanian squamate assemblages of these two continents. Alternatively, dispersal between the two landmasses may have occurred by transatlantic rafting. During the Cenomanian–Santonian interval, Bicuspidon colonized the European archipelago probably from North Africa, like many “Eurogondwanan” taxa.     

Diversity in the later Paleogene proboscidean radiation: a small barytheriid from the Oligocene of Dhofar Governorate,Sultanate of Oman

Despite significant recent improvements to our understanding of the early evolution of the Order Proboscidea (elephants and their extinct relatives), geographic sampling of the group’s Paleogene fossil record remains strongly biased, with the first ~30 million years of proboscidean evolution documented solely in near-coastal deposits of northern Africa. The considerable morphological disparity that is observable among the late Eocene and early Oligocene proboscideans of northern Africa suggests that other, as yet unsampled, parts of Afro-Arabia might have served as important centers for the early diversification of major proboscidean clades. Here we describe the oldest taxonomically diagnostic remains of a fossil proboscidean from the Arabian Peninsula, a partial mandible of Omanitherium dhofarensis (new genus and species), from near the base of the early Oligocene Shizar Member of the Ashawq Formation, in the Dhofar Governorate of the Sultanate of Oman. The molars and premolars of Omanitherium are morphologically intermediate between those of Arcanotherium and Barytherium from northern Africa, but its specialized lower incisors are unlike those of other known Paleogene proboscideans in being greatly enlarged, high-crowned, conical, and tusk-like. Omanitherium is consistently placed close to late Eocene Barytherium in our phylogenetic analyses, and we place the new genus in the Family Barytheriidae. Some features of Omanitherium, such as tusk-like lower second incisors, the possible loss of the lower central incisors, an enlarged anterior mental foramen, and inferred elongate mandibular symphysis and diminutive P₂, suggest a possible phylogenetic link with Deinotheriidae, an extinct family of proboscideans whose origins have long been mysterious.     

A new phylogeny for basal Trechnotheria and Cladotheria and affinities of South American endemic Late Cretaceous mammals

The endemic South American mammals Meridiolestida, considered previously as dryolestoid cladotherians, are found to be non-cladotherian trechnotherians related to spalacotheriid symmetrodontans based on a parsimony analysis of 137 morphological characters among 44 taxa. Spalacotheriidae is the sister taxon to Meridiolestida, and the latter clade is derived from a primitive spalacolestine that migrated to South America from North America at the beginning of the Late Cretaceous. Meridiolestida survived until the early Paleocene (Peligrotherium) and early Miocene (Necrolestes) in South America, and their extinction is probably linked to the increasing competition with metatherian and eutherian tribosphenic mammals. The clade Meridiolestida plus Spalacotheriidae is the sister taxon to Cladotheria and forms a new clade Alethinotheria. Alethinotheria and its sister taxon Zhangheotheria, new clade (Zhangheotheriidae plus basal taxa), comprise Trechnotheria. Cladotheria is divided into Zatheria (plus stem taxa, including Amphitherium) and Dryolestida, including Dryolestidae and a paraphyletic array of basal dryolestidans (formerly classified as “Paurodontidae”). The South American Vincelestes and Groebertherium are basal dryolestidans.     

Roost type influences torpor use by Australian owlet-nightjars

Australian owlet-nightjars (Aegotheles cristatus; ～50?g) are one of only a few avian species that roost in cavities year-round and regularly enter torpor. Cavity roosts act as thermal buffers, and roost type likely affects energy expenditure of small birds. We used radiotelemetry to locate diurnal winter roost sites of owlet-nightjars in central Australia and to measure body (T (b)) and skin (T (skin)) temperature. We also recorded ambient temperature inside (T (IN)) and outside roosts. Individual owlet-nightjars used one to seven different roosts (tracking time 3-10?weeks), selecting either rock crevices (four birds) or tree hollows (four birds), or switching between the two roost types (seven birds). Rock crevices (T (IN) +9°C to +33°C) were warmer and thermally more stable than tree hollows (T (IN) -4.0°C to +37°C). Torpor, often expressed by a reduction of T (skin)/T (b) by >10°C for 3-4?h at dawn, was influenced by roost selection; torpor use in tree hollows was almost twice that in rock crevices. Despite the potential energy savings accrued from roosting in well-insulated cavities, owlet-nightjars roosted in tree hollows more often (65% bird days, n?=?398) than in rock crevices (35% bird days, n?=?211). Lower costs of arousal from torpor via passive rewarming and basking and decreased risk of predation are two possible explanations for the preference to roost in tree hollows. We provide the first evidence for the influence of cavity roost selection on torpor use in a free-ranging bird and show that roost selection and thermal biology are strongly interrelated in determining energy expenditure.     

Cladistic biogeographic analysis suggests an early Caribbean diversification in Mexico

The Great American Biotic Interchange has been the predominant paradigm for explaining biotic diversification in the Nearctic/Neotropical overlap or Mexican Transition Zone, which is commonly explained by the collision of the North and South American continental plates, which began in the Oligocene and fused both landmasses. In the most far-reaching cladistic biogeographical analysis of the area to date, evidence has been found supporting the existence of a remnant Caribbean region extending from eastern Mexico to southeastern USA, a hypothesis that challenges current views of the Great American Biotic Interchange and the Mexican Transition Zone. We show herein that an older terrane, which has drifted to the present day positions of Yucatan and Cuba, may be biogeographically linked to an early ‘Gondwanan’ biota of the Paleocene (ca. 60 Ma). The evidence indicates an east–west biotic divide in Mexico, existing before the collision and formation of Central America. The south–north division of the country, previously recognized by several authors as associated with the Great American Biotic Interchange and the Mexican Transition Zone, is of a younger age. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.