Wednesday, September 3, 2014

New paper on fossil plant from the Neotropics

Yes, fossil plants! This is a first in this blog, which is otherwise, heavily biased towards marine tetrapods. However, that doesn't mean that when I do fieldwork I only focus on collecting fossil vertebrates. this of course has resulted in a number of publication on fossil invertebrates (e.g. Schweitzer et al., 2006), and now plants. The paper which was just published in the journal International Journal of Plant Sciences is a collaborative work led by former Florida Museum of Natural History colleague Fabiany Herrera, one of the few experts on fossil plants from the Neotropics, and his former advisors Steven R. Manchester and Carlos Jaramillo. The paper is a follows up on a previous paper published about four years ago in the same journal (Herrera et al., 2010), and that seeks to better understand the evolutionary and paleobiogeographic history of a group of plants called Humiriaceae that are found in the Neotropics, and western Africa. This group is mainly composed of large trees, most greater than 20 meters tall, and fruits have woody parts with very particular morphology, which results in a relatively high preservation and identification potential.
Map showing the distribution of fossil Humiriaceae endocarps (symbols) and extant genera (dashed lines) (modified from Herrera et al., 2010:fig. 1).
In the paper we describe fossilized endocarps (the inside part of the fruit) from the early Oligocene of Peru and Puerto Rico, and the late Miocene of Panama, and fossilized wood from the late Eocene of Panama (Herrera et al., 2014). The fossilized fruit from the Oligocene of Peru, which we dubbed Duckesia berryi, represents a new species of a tree that is nowadays only found in Amazonia, and is the oldest record of that genus. This not only shows that this particular taxon has an older history than previously thought, but it also shows that its former distribution was much more widespread. In addition to that, the fossil wood we describe, called Humiriaceoxylon ocuensis, shows that by the late Eocene parts of what is now Panama, was forested by large trees belonging to this particular group of plants. In addition, Fabiany had previously described a fossil Humiriaceae endocarp which he names Lacunofructus cuatrecasana from a locality near where the wood was found, and it may actually be that they represent the same tree (Herrera et al., 2012, 2014)*. This is a really cool find, as the region and where the fossils were collected, was not connected by land to neither North or South America, showing again, that overwater dispersal is not as much a problem for plants.
*Paleobotanists use different scientific names for the different parts of a plant as they are usually found separate, hence the endocarp has a name, and the wood another, even though they may be the same plant.
The fossil endocarp Duckesia berry (A-L) from the Oligocene of Peru, compared with the endocarp of the modern species D. verrucosa (M-O). (Modified from Herrara et al., 2014:fig. 1.)
The fossil from Puerto Rico consists of an endocarp of Sacoglottis tertiaria, otherwise known from the Neogene of Peru, Ecuador, Colombia and Panama (Herrera et al., 2010). Several species of the genus Sacoglottis are still found today, in the Amazonian region and west Africa. The fossil from Puerto Rico is from the early Oligocene San Sebastian Formation, one of my favorite formations where I've spent many hours searching for fossils. Actually, the locality where I found the endocarp is not far from where Aktiogavialis puertoricensis, Priscosiren atlantica, and a Caviomorph rodent tooth were collected (Velez-Juarbe et al., 2007; Velez-Juarbe and Domning, 2014; Velez-Juarbe et al., 2014).
The fossil endocarp Sacoglottis tertiaria from the early Oligocene San Sebastian Formation of Puerto Rico.
Fossil plants were previously described from the San Sebastian Fm. by previous workers, mainly, Arthur Hollick (1928) and Alan Graham and David Jarzen (1969). But none of the material they described indicated the presence of Humiriaceae in the island. In fact, they list many plant groups present in San Sebastian Fm. which are now absent from the flora of the island, now the Humiriaceae can be added to that list. As I recently said in a newspaper interview, Puerto Rico during the Oligocene was very different from nowadays, and there is still more to be discovered!

Assorted Musing
The fossil endocarp from Puerto Rico, was previously featured on this blog, it was the only thing I found in my two days of fieldwork in January 2009. I was a bit disappointed at first, but not any more!

I should also acknowledge my wife, it was because of her that I ended up visiting the Florida Museum of Natural History in the fall of 2012, which is where I met Fabiany, told him about the fossil endocarp, and I ended up being a co-author in his paper.

References

Graham, A., and D. M. Jarzen. 1969. Studies in Neotropical paleobotany. I. The Oligocene communities of Puerto Rico. Annals of the Missouri Botanical Garden 56:308-357.

Herrera, F., S. R. Manchester, and C. Jaramillo. 2012. Permineralized fruits from the late Eocene of Panama give clues of the composition of forests established early in the uplift of Central America.

Herrera, F., S. R. Manchester, J. Velez-Juarbe, and C. Jaramillo. 2014. Phytogeographic history of the Humiriaceae (Part 2). International Journal of Plant Sciences 175:828-840.

Herrera, F., S. R. Manchester, C. Jaramillo, B. MacFaddem, S. A. da Silva-Caminha. 2010. Phytogeographic history and phylogeny of the Humiriaceae. International Journal of Plant Sciences 171:392-408.

Hollick, A. 1928. Paleobotany of Porto Rico. Scientific Survey of Porto Rico and the Virgin Islands 7(3):177-393.

Schweitzer, C. E., M. Iturralde-Vinent, J. L. Hetler, and J. Velez-Juarbe. 2006. Oligocene and Miocene decapods (Thalassinidea and Brachyura) from the Caribbean. Annals of Carnegie Museum 75:111-136.

Velez-Juarbe, J., and D. P. Domning. 2014. Fossil Sirenia of the West Atlantic and Caribbean region: X. Priscosiren atlantica, gen. et sp. nov. Journal of Vertebrate Paleontology 34:951-964.

Velez-Juarbe, J., C. A. Brochu, and H. Santos. 2007. A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile. Proceedings of the Royal Society B 274:1245-1254.

Velez-Juarbe, J. T. Martin, R. D. E. MacPhee, and D. Ortega-Ariza. 2014. The earliest Caribbean rodents: Oligocene caviomorphs from Puerto Rico. Journal of Vertebrate Paleontology 34:157-163.

Wednesday, July 9, 2014

Its the 10th installment of Fossil Sirenia of the West Atlantic and Caribbean Region!

Today came out the most recent issue of the Journal of Vertebrate Paleontology. Amongst many other interesting papers, there is one by yours truly and former PhD advisor Daryl Domning. In our paper we describe a new sirenian taxon from early Oligocene deposits in Puerto Rico and South Carolina, its our second new species this year, as some months ago we published the description of Metaxytherium albifontanum Velez-Juarbe and Domning, 2014 (read more about it here). The fossil from Puerto Rico, which is fairly complete, comes from the same overall locality as some other fossils I've mentioned in previous posts, like Aktiogavialis puertoricensis Velez-Juarbe et al., 2007, and the oldest West Indian rodent (Velez-Juarbe et al., 2014).  This paper also marks the Tenth (!!!!) installment of the series on Fossil Sirenia of the West Atlantic and Caribbean Region, which Daryl started in 1988 (Domning, 1988)! Such long-lasting series are very uncommon!

The last time a new species of sirenian was described from Puerto Rico was 1959, when Roy H. Reinhart in his monumental work on Sirenia and Desmostylia, described Caribosiren turneri (see picture below) from the San Sebastian Formation in the northwestern part of the island. Caribosiren is a weird dugongid, it has a rostral deflection (downturning of the snout) of nearly 90º, and apparently no tusks!!
Caribosiren turneri Reinhart, 1959, from the San Sebastian Formation of Puerto Rico. Notice the extremely downturned snout  which always reminds me of Gonzo! The tip of the snout, although not preserved very well, hints at a lack of tusks.
(Photo courtesy of N.D. Pyenson) (Click on image to see larger version.)
The new fossil is from the same formation as Caribosiren. We named our new species Priscosiren atlantica, in reference to its ancestral relationship to other fossil dugongids (prisco means ancient, former) and its occurrence in the Western Atlantic region.
Priscosiren atlantica is known from multiple elements of two individuals, one from the Puerto Rico (USNM 542417) the other from South Carolina (SC 89.254). Its one of the most complete early Oligocene sirenians known. (Outline of skeleton modified from Cope, 1890).
(Click on image to see larger version.)

Slides from a talk that Daryl and I gave at the 2013 SVP annual meeting. Here we point out to several of the characters that diagnose Priscosiren atlantica as well as its relationship to other dugongids.
(Click on image to see larger version.)
 Priscosiren is represented by at least two individuals (an adult and a subadult), with associated cranial and postcranial material, making it one of the best known early Oligocene sirenians. This species occupies a special place amongst other dugongids as it seems to be ancestral (hence its name) to a clade that includes Metaxytherium spp. + Hydrodamalinae, and Dugonginae (see above). More interestingly, is that Priscosiren is found in the same formation as Caribosiren in Puerto Rico, and Crenatosiren olseni in South Carolina, and hints at the presence of sirenian multispecies communities (Velez-Juarbe et al., 2012) during the early Oligocene.

The day we found the holotype specimen of Priscosiren (USNM 542417) was the same day we found the holotype of Aktiogavialis puertoricensis, which we were able to collect that same day. In contrast, collecting Priscosiren was an ordeal, it is a long story, of discovery, failed attempts at collecting it, loss of parts, and final recovery. So stay tuned for an upcoming post about that story!

References

Domning, D. P. 1988. Fossil Sirenia of the West Atlantic and Caribbean Region. I. Metaxytherium floridanum Hay, 1922. Journal of Vertebrate Paleontology 8:395-426.

Reinhart, R. H. 1959. A review of the Sirenia and Desmostylia. University of California Publications in Geological Sciences 36(1):1-146.

Velez-Juarbe, J., C. A. Brochu, and H. Santos. 2007. A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a nonmarine reptile. Proceedings of the Royal Society B 274:1245-1254.

Velez-Juarbe, J., and D. P. Domning. 2014. Fossil Sirenia of the West Atlantic and Caribbean Region. IX. Metaxytherium albifontanum. Journal of Vertebrate Paleontology 34:444-464.

Velez-Juarbe, J., and D. P. Domning. 2014. Fossil Sirenia of the West Atlantic and Caribbean Region. X. Priscosiren atlantica gen. et sp. nov. Journal of Vertebrate Paleontology 34:951-964.

Velez-Juarbe, J., D. P. Domning, and N. D. Pyenson. 2012. Iterative evolution of sympatric seacow (Dugongidae, Sirenia) assemblages during the past ~26 million years. PLoS ONE 7:e31294.

Velez-Juarbe, J., T. Martin, R. D. E. MacPhee, and D. Ortega-Ariza. 2014. The earliest Caribbean rodents: Oligocene caviomorphs from Puerto Rico. Journal of Vertebrate Paleontology 34:157-163.

Tuesday, June 24, 2014

Time for a brief update!

Dear readers, things have been a bit quiet here, mostly because lots of exciting things have been happening! To begin, earlier this year Caribbean Paleobiology moved to the West Coast, after spending most of the last 6 and a half years between Washington DC, Florida and Panama. The reason for the move is that I am now an NSF Postdoctoral Fellow based with Jim Parham and at the John D. Cooper Archaeological and Paleontological Center, Department of Geological Sciences, California State University-Fullerton, and also, starting next month, I'll be the new curator of Marine Mammals (living and extinct) at the Natural History Museum of Los Angeles County!!
My postdoc project will be sort of a follow-up on a paper I published on sirenian multispecies assemblages a couple of years ago, now it will be taxonomically broader and focused on a different part of the world. By taxonomically broader I mean sirenians, desmostylians and aquatic sloths! All which were, or are considered, marine mammal herbivores.

Since arriving in California I've been busy visiting some sites like Sharktooth Hill, and museum collections, including the San Diego Natural History Museum, the museum at the Universidad Autónoma de Baja California, in Ensenada, and a return trip to Washington DC to attend the 2014 Secondary Adaptations meeting and of course the collections at the National Museum of Natural History in DC.
Sharktooth Hill National Natural Landmark in Bakersfield, California. The famous bone bed is about 10 meters or so below the car.
Tania (my wife) hold the first neck vertebra of Hydrodamalis cuestae, which was the largest species of sea cow, ever! It reached more than 8 meters in length, large for a sea cow, but small when compared to some whales.
One of the highlights of the 2014 SecAd meeting. We got a brief lecture on beaked whales from Jim Mead curator emeritus of marine mammals at the Smithsonian and one of the Worlds expert on that group of whales.

The other thing that's has kept me busy is that back in May, I was in Baja California Sur with several colleagues from Howard University (HU), New York Institute of Technology (NYIT), Universidad Autónoma de Baja California (UABC), and Universidad Autónoma de Baja California Sur (UABCS), most which you can see in the picture below. This trip was a continuation of work we did back in 2012 (see previous post about that trip). The trip was fun, and we found many interesting fossils.
Part of the BCS Paleo Project 2014 reopening the main quarry. From left to right, Arely Cedillo (UABCS), Gerardo González (UABCS), Ehecatl Hernández (UABCS), Brian Beatty (NYIT), Daryl Domning (HU), and Lizeth González (UABCS). Missing from the picture Azucena Solis (UABCS) and Fernando Salinas (UABC). Click on the picture to see the larger version.
So, stay tune as new papers will be coming out soon and I'll get around to post some more about the Baja trip!!

Wednesday, March 5, 2014

Florida gets a new species of fossil seacow!

Yesterday saw the publication (online) of the second issue of 2014 of Journal of Vertebrate Paleontology. Published in this issue is the description of the first new species of seacow from the Western Atlantic that I get to name. In collaboration with Daryl P. Domning, this is the latest installment in the series titled "Fossil Sirenia of the West Atlantic and Caribbean Region" which Daryl started in 1988 (Domning, 1988). Our new species, named Metaxytherium albifontanum is known from late Oligocene deposits in Florida and South Carolina. The generic name albifontanum translates into white springs (albus = white; fontanus = spring or fountain). But why did we choose that name? and what is Metaxytherium? Keep reading and you'll find out why and more. 

Scientific Names
The scientific name of organisms consist of two parts: the genus and the species. The genus is a more inclusive rank, whereas the species is more unique. In a way, you can think of the genus name as an equivalent to your last name, where there will be more members (e.g. siblings and/or parents) with that same last name, and the species name as your first name; the two, together, will form a unique combination which applies only to you. We use scientific names in order to infer relationships amongst organisms, and these are usually latinized so that they can be understood by anyone, anywhere, as a common language, instead of using the common name which changes by country and language. Now, when describing a new species and giving it a scientific name, you can choose whichever name you think appropriate, as long as its not your own (Linnaeus was the one exception; there are other rules for naming, which you can find here). You can name a species after a musician who was an inspiration, the country where it was found, or in honor of a fellow researcher, just to name a few examples.
Renowned paleontologist George Gaylord Simpson named several fossil sirenians from Florida (Simpson, 1932). Simpson had a thing for using cleverly latinized versions of formation or locality names for his new species. For example, he described some fossils from the Bone Valley district in central Florida and gave them the scientific name Felsinotherium ossivallense*, (ossivallense = Bone Valley), while another one he named Hesperosiren crataegensis*, which takes its name from Crataegus, the genus name of a plant commonly known as hawthorn, which in turn is also the name of the sedimentary unit, the Hawthorn Group, where Simpson's specimen was found. So, as a homage to G. G. Simpson and his work on the fossil sirenians from Florida we decided to use a latinized version of the name of the town of White Springs, FL, which is close to where the holotype (= name-bearing specimen) of our new species was collected; resulting in the combination Metaxytherium albifontanum.
*both Felsinotherium and Hesperosiren were later synonymized with Metaxytherium


Metaxytherium albifontanum is known from multiple elements of several individuals (each color identifies elements represented by one or more specimens; white = unknown). This makes it one of the most complete fossil sirenians known. (Outline of skeleton modified from Cope, 1890). (Click on the image to see larger version.)
What is Metaxytherium
Metaxytherium is a widespread and relatively well-known genus of fossil dugongid. There are now a total of eight species under this genus, it has a wide temporal distribution, ranging from the late Oligocene through early Pliocene, and a broad geographical distribution, with species known from Europe, northern Africa, and the Americas. Most of the species known were described and named between 1822 and the first half of the 1900's, so, unexpectedly, there was a bit of a taxonomic mess (this happens more often than we'd like). Fortunately, since 1987, there have been several papers providing us with detailed descriptions of some of the known species, as well as phylogenetic analyses (e.g. Domning and Thomas, 1987; Domning, 1988; Aranda-Manteca et al., 1994; Domning and Pervesler, 2001; Sorbi, 2008; Sorbi et al., 2012). These works have help clarify some of the taxonomic confusion surrounding some of the old names, and even a new species was described, Metaxyterium arctodites Aranda-Manteca et al., 1994, from Baja California and California. That makes M. albifontanum the first species of Metaxytherium named in 20 years!! Meaning that we are not done learning about the diversity of this group, and more may still be waiting to be described.


Slides from a talk Daryl and I gave at the Society of Vertebrate Paleontology 2013 Annual Meeting. Here we use M. albifontanum to illustrate some of the features that characterize the genus Metaxytherium (top two and bottom left). The phylogenetic tree on the bottom right shows the relationship between Metaxytherium spp. and other sirenians (modified from figure 15 of our paper). (Click on the image to see larger version.) 
Our new species differed from all other known species in the group. Not only that, it is the geologically oldest species of Metaxytherium. Previous assumptions on the origins of Metaxytherium had hypothesized an European origin for the group, our discovery changes that and seems to indicate a Western Atlantic origin for the genus.

Relationships with Other Species
One of the relevant results of our paper is that we got to properly define Metaxytherium. Our phylogenetic analysis (see tree above, bottom right) was consistent with previous work (e.g. Domning, 1994), showing a close relationship between Metaxytherium spp. and hydrodamalines (the group that include Steller's seacow). We also got some interesting results regarding the relationships amongst the different species of Metaxytherium. Our results indicate that the split between Metaxytherium albifontanum and the geologically younger M. krahuletzi (from the early Miocene of Europe), occurred before the late Oligocene, as the latter occupies a more basal position within the tree. The relationships within the group also seem to point to multiple dispersals across the Atlantic and/or a high degree in morphological convergence.

Paleoecology
Metaxytherium albifontanum was part of a sirenian multi-species assemblage in the late Oligocene of Florida, together with Dioplotherium manigaulti and Crenatosiren olseni. As part of that assemblage, we hypothesize M. albifontanum as a consumer of small-sized seagrasses such as eelgrass, while the other species likely fed on larger species. If this sounds familiar is because I wrote about this subject in a previous post. In fact, M. albifontanum was one of the species that inspired the iterative evolution project with Daryl and Nick Pyenson, which resulted in our open access publication in PLoS ONE (Velez-Juarbe et al., 2012). 

Assorted Random Musing: 
  • I visited the Florida Museum of Natural History in 2011 to study one of the specimens (UF 49051), little did I know at that time that I would end up as a Postdoc here!
  • You can see the name-bearing specimen, UF 49051, in the Florida Fossils: Evolution of Life and Land exhibit at the Florida Museum of Natural History.
  • It so happened that I wrote this post from a desk at the Simpson Library of Paleontology, its filled with books and reprints donated by him, and...
  • There are a lot of pictures of Simpson in this library, in some, he kind of looks like the long lost brother of Colonel Sanders...
Stay tuned as more new fossils seacows will be showing up here later this year!!

References

Aranda-Manteca, F. J., D. P. Domning, and L. G. Barnes. 1994. A new Middle Miocene sirenian of the genus Metaxytherium from Baja California: relationships and paleobiogeographic implications. Proceedings of the San Diego Society of Natural History 29:191-204.

Cope, E. D. 1890. The extinct Sirenia. American Naturalist 24:697-702.

Domning, D. P. 1988. Fossil Sirenia of the West Atlantic and Caribbean Region. I. Metaxytherium floridanum Hay, 1922. Journal of Vertebrate Paleontology 8:395-426.

Domning, D. P. 1994. A phylogenetic analysis of the Sirenia. Proceedings of the San Diego Society of Natural History 29:177-189.

Domning, D. P., and P. Pervesler. 2001. The osteology and relationships of Metaxytherium krahuletzi Depéret, 1895 (Mammalia: Sirenia). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 553:1-89.

Domning, D. P., and H. Thomas. 1987. Metaxytherium serressii (Mammalia: Sirenia) from the early Pliocene of Libya and France: a reevaluation of its morphology, phyletic position, and biostratigraphic and paleoecological significance; pp. 205-232 in N. Boaz, A. El-Arnauti, A. W. Gaziry, J. de Heinzelin, and D. D. Boaz (eds.), Neogene Paleontology and Geology of Sahabi. New York (Liss).

Simpson, G. G. 1932. Fossil Sirenia of Florida and the evolution of the Sirenia. Bulletin of the American Museum of Natural History 59:419-503.

Sorbi, S. 2008. New record of Metaxytherium (Mammalia, Sirenia) form the lower Miocene of Manosque (Provence, France). Geodiversitas 30:433-444.

Sorbi, S., D. P. Domning, S. C. Vaiani, and G. Bianucci. 2012. Metaxytherium subapenninum (Bruno, 1839) (Mammalia, Dugongidae), the latest sirenian of the Mediterranean Basin. Journal of Vertebrate Paleontology 32:686-707.

Velez-Juarbe, J., and D. P. Domning. 2014. Fossil Sirenia of the West Atlantic and Caribbean Region. IX. Metaxytherium albifontanum sp. nov. Journal of Vertebrate Paleontology 34:444-464.

Velez-Juarbe, J., D. P. Domning, and N. D. Pyenson. 2012. Iterative evolution of sympatric seacow (Dugongidae, Sirenia) assemblages during the past ~26 million years. PLoS ONE 7:e31294.

Tuesday, February 25, 2014

¡Muerte Súbita en el Océano!

Hoy sale publicado en la revista científica Proceedings of the Royal Society B el trabajo titulado Repeated mass strandings of Miocene marine mammals from Atacama Region of Chile point to sudden death at sea (lo pueden bajar GRATIS). El mismo, es el resultado de un proyecto colaborativo entre investigadores nacionales e internacionales y donde se combina el uso de herramientas tradicionales de la paleontología con equipos y técnicas innovadoras para obtener datos tales como digitalización 3D y fotogrametría. En este trabajo tomamos un enfoque multidisciplinario para descubrir que le ocurrió a un grupo de vertebrados marinos que murieron al mismo tiempo y posteriormente fueron depositados y preservados en lo que fuese una planicie de marea en la costa del norte de Chile a finales del periodo Mioceno (entre 6-9 millones de años atrás). La localidad donde se hizo el estudio se le conoce como Cerro Ballena, la cual está localizada en la Región de Atacama (cerca del desierto más seco del mundo), y al norte del pueblo de Caldera (lugar que inspiró al autor de Condorito y donde se venden las mejores empanadas que he comido).
Vista desde el extremo sur de Cerro Ballena, desde aquí se puede observar el pueblo de Caldera y océano Pacífico. La Carretera que se ve es parte de la Autopista Panamericana.
Gracias a las fuerzas tectónicas que a través del tiempo han cambiado la costa occidental de América del Sur, Cerro Ballena se encuentra hoy día alejado de la costa y sobre el nivel del mar. Si "Cerro Ballena, Chile y cetáceos fósiles" les suena familiar en este blog, es porque hace casi dos años atrás fui parte de una expedición científica donde recolectamos datos para este trabajo (visiten las entradas anteriores sobre ese viaje aquí, aquí, aquí, aquí, y aquí).

El Descubrimiento
Previo a nuestro trabajo, ya se conocía sobre la existencia de fósiles en Cerro Ballena, por ende el nombre. Sin embargo, los fósiles que se encontraban allí no estaban completamente expuestos, por lo cual colectarlos para estudiarlos hubiera sido muy laborioso y costoso. Afortunadamente, durante los trabajos de ensanchamiento de la Autopista Panamericana, la cual pasa por la localidad, muchos de estos fósiles quedaron expuestos y se notó que estos sólo se encontraban en ciertas capas sedimentarias, todas pertenecientes a una formación geológica llamada Formación Bahía Inglesa. Al ser tan grande la cantidad de fósiles que quedaron expuestos tuvieron que detener el trabajo de ensanchamiento de la autopista para rescatar los mismos. Estos esfuerzos fueron liderados por un equipo de paleontólogos del Museo Paleontológico de Caldera, el Museo Nacional de Historia Natural de Chile y la Universidad de Chile, los cuales se encargaron de mapear la posición de cada fósil en su respectivo horizonte. Más tarde se unieron a ese esfuerzo los integrantes de la oficina de digitación 3D del Smithsonian quienes hicieron scans de los fósiles in situ para poder salvaguardar esa información ya que pasará mucho tiempo para que algunos de estos sean preparados y estudiados en detalle.

Algunas de las ballenas siendo preparadas para ser colectadas. Cada carpa negra indica la localización de una ballena. A la derecha Ana M. Valenzuela-Toro, una de las coautoras del trabajo junto al espécimen B33.
El equipo de digitación 3D del Smithsonian. Adam Metallo (izquierda) y Vincent Rossi (derecha) utilizan varias herramientas para escanear a la ballena B33. Gracias al scan se pueden hacer imágenes como la que utilicé arriba para el banner de este blog al igual que imprimir copias 3D del mismo.
Aquí las ballenas ya colectadas y guardadas en el Museo Paleontológico de Caldera. 
No es inusual encontrar fósiles de vertebrados marinos en esta parte del mundo, de hecho, más al norte en Perú se encuentra la Formación Pisco, la cual es similar en edad a la Formación Bahía Inglesa (ronda entre los 10-4 millones de edad) y es una de las formaciones geológicas más estudiadas y más conocidas por su diversidad de mamíferos marinos extintos. Sin embargo, Cerro Ballena es diferente a lo que se ha encontrado en la Pisco y estas diferencias son lo que hacen Cerro Ballena un lugar especial.

Miembros del Team Ballena tomando datos de los horizontes fosilíferos en Cerro Ballena.
El Misterio de Cerro Ballena
En Cerro Ballena logramos identificar cuatro horizontes fosilíferos. Cada horizonte contenía un conjunto de vertebrados marinos, siendo la atracción principal la presencia de esqueletos completos o casi completos de ballenas barbadas, algunas hasta de 8 metros (26 pies) de largo. Nuestras observaciones indicaban que lo que pasó en Cerro Ballena fueron varios eventos de varamiento masivo. Quizás ya hayan escuchado sobre varamiento masivos de mamíferos marinos, ya que es un fenómeno que generalmente capta la atención de lo medios noticiosos. De hecho, muchos de los los varamientos masivos que ocurren hoy día generalmente incluyen a los cetáceos dentados (odontocetos: delfines, marsopas, cachalotes, etc). Un causante principal de este tipo de varamiento resulta como efecto secundario del uso de sonar por las marinas de guerra. Sin embargo los eventos que ocurrieron en Cerro Ballena tomaron lugar millones de años atrás, cuando nuestros ancestros eran muy primitivos y ni siquiera habían salido de África, así que podemos descartar esa causa para lo que ocurrió allí. Lo peculiar de Cerro Ballena, es que junto con las ballenas barbadas también encontramos otros vertebrados marinos, incluyendo cachalotes, delfín-morsa, focas, perezosos acuáticos y marlines. Esta ocurrencia multi-taxonómica señala a algún otro tipo de mecanismo que afectó por igual a todos estos vertebrados marinos y causó su varamiento masivo. Afortunadamente (bueno, quizás no para los animales) varamientos masivos que incluyen distintos grupos de vertebrados marinos ocurren hoy día. Estudios detallados de este tipo de varamientos han determinado que los mismos ocurren por intoxicación causada por floración de algas nocivas (también conocidos como marea roja). Las mareas rojas son comunes en zonas de surgencia, y gracias a estudios previos, sabemos que la costas de Chile y Perú son zonas de surgencia desde hace millones de años. Esta peculiar localidad no solo nos permite una ventana al pasado y descubrir un ecosistema marino con organismos distintos a los que vemos hoy día, si no que también nos permitió descubrir que eventos de marea roja, muy similares a los que todavía observamos hoy día y que continúan afectando a los vertebrados marinos han estado ocurriendo durante millones de años. Incluso, podemos predecir que otros depósitos fosilíferos ricos en vertebrados marinos en zonas de turgencia, como por ejemplo la Formación Pisco en Perú, puede que hayan preservado eventos como los que descubrimos en Cerro Ballena.
Nuestro trabajo en Cerro Ballena aún no termina, todavía nos queda describir en detalle muchos de los organismos que vivieron, y murieron en la costa chilena hace millones de años atrás.

Para mayor información sobre nuestro proyecto visiten la página oficial de Cerro Ballena.

En la página de Smithsonian X 3D pueden ver parte los fósiles que fueron escaneados e incluso si tienes acceso a una impresora 3D, imprimir tu copia personal de los fósiles.

También visiten la página del Pyenson Lab donde encontrarán más fotos y entradas sobre las distintas expediciones a Chile.

Visiten también:
Not Exactly Rocket Science
Smithsonian Science
Smithsonian Magazine
Science Magazine
PhysOrg
BBC News
Washington Post

Referencia:

Pyenson, N. D., C. S. Gutstein, J. F. Parham, J. P. Le Roux, C. Carreño Chavarría, H. Little, A. Metallo, V. Rossi, A. M. Valenzuela-Toro, J. Velez-Juarbe, C. M. Santelli, D. Rubilar Rogers, M. A. Cozzuol, and M. E. Suárez. 2014. Repeated mass strandings of Miocene marine mammals from Atacama Region of Chile point to sudden death at sea. Proceedings of the Royal Society B [in press].

Monday, January 27, 2014

Los roedores más antiguos del Caribe

Ya estamos en enero, el primer mes del nuevo año y con el llega la primera edición del año de la revista científica Journal of Vertebrate Paleontology. En este ejemplar se han publicado muchos trabajos interesantes, incluyendo varios sobre mamíferos marinos, lo cual es un tema recurrente en este blog. Sin embargo, en esta entrada hablaremos de otros fósiles. En esa misma edición ha salidos publicado un trabajo donde mis colegas y yo describimos fósiles de roedores de Puerto Rico (Vélez-Juarbe et al., 2014). Estos fósiles son únicos ya que son la evidencia más antigua de roedores caviomorfos en las Antillas Mayores, y nos ayudan a entender el cuando llegaron los vertebrados terrestres a la región, lo cual ha sido tema de debate durante varias décadas.

Roedores fósiles de Puerto Rico
No es la primera vez que se encuentran fósiles de roedores en Puerto Rico. De hecho, es bien conocido que durante el Pleistoceno* en Puerto Rico, La Española, Cuba, Jamaica, y algunas de las Antillas Menores, habitaban distintos grupos de roedores endémicos, perezosos terrestres, monos, y musarañas (Woods y Sergile, 2001). Muchos de estos fósiles de mamíferos terrestres del Pleistoceno fueron descritos a principios del siglo pasado (e.g. Anthony, 1918), aunque han habido descubrimientos más recientes (Turvey et al., 2006). Algunos de los roedores fósiles que han sido descritos de estas islas eran gigantes, como por ejemplo, Amblyrhiza inundata de la isla de Anguila, la cual tenía una masa corporal similar a la de un oso negro americano (Biknevicious et al., 1993)!
*periodo geológico que comenzó hace 2.6 millones de años y duró hasta hace 10,000 años

Siendo más específico, durante el Pleistoceno, en Puerto Rico habitaban al menos tres especies de ratas espinosas (Heteropsominae), y posiblemente dos especies de hutías gigantes (Heptaxodontidae) (información adicional aquí). Un tercer grupo de roedores endémicos que existió durante el mismo tiempo, en incluso todavía habita algunas de las otras Antillas, eran las hutías capromíidas, pero hasta donde sabemos estas nunca habitaron Puerto Rico. Todos estos grupos de roedores pertenecer a un grupo taxonómico más grande llamados caviomorfos. Los caviomorfos son endémicos de Sur América; sus ancestros llegaron de África alrededor de 54 millones de años atrás (Antoine et al., 2012). Una vez llegan a Sur América, los caviomorfos se diversifican y se dispersan a través de todo el continente y el Caribe. Algo muy similar también ocurrió con los primates suramericanos (platirrinos) (Kay, 2014), los cuales tienen una historia casi tan antigua como los roedores caviomorfos.
Foto tomada a principios de enero cuando visité de nuevo la localidad de las Calizas Lares donde encontré el diente incisor.
Los nuevos fósiles de Puerto Rico consisten de un par de dientes incisores, uno proveniente de la Formación San Sebastián (en esta localidad), y el otro de las Calizas Lares (de esta localidad; ver foto arriba). Respectivamente, estas formaciones se depositaron durante las épocas geológicas conocidas como Oligoceno temprano y Oligoceno tardío. Cuando encontré los fósiles allá para el 2005, me comuniqué con Ross MacPhee, curador de mamíferos del Museo Americano de Historia Natural en Nueva York, y quien durante años ha tenido interés en los orígenes de la fauna terrestre de las Antillas Mayores. Basado en lo que ya conocíamos sobre el registro fósil de Puerto Rico, Ross y yo sospechábamos que estos dientes incisores pertenecían a un roedor caviomorfo. Sin embargo, teníamos un problema, como pueden observar en la foto abajo a la derecha, los incisores son muy sencillos, ya que carecen de las crestas, cúspides y valles que uno vería en un molar, esto nos hacía difícil el poder identificarlos usando solo sus características externas. Lo siguiente que se nos ocurrió fue tratar de ver la estructura microscópica (microestructura) del esmalte, esta si preserva características que nos podrían ayudar a identificar el diente y saber a que grupo de roedores pertenecía. Estudiar la estructura microscópica de los dientes toma varios pasos: hay que hacer un corte transversal del diente, pulir la superficie y finalmente, utilizando un microscopio electrónico de barrido, ver la microestructura del diente. Para llevar a cabo esta tarea decidimos contactar a Thomas Martin, paleontólogo de la Universidad de Bonn, en Alemania y quien es uno de los expertos en el estudio de microestructura de los dientes. Luego convencer a Thomas, le enviamos los fósiles, y luego de varios meses, nos envió un correo electrónico con detalles de los resultados.
A la izquierda una representación de como son los roedores caviomorfos mostrando la posición del diente incisor. A la derecha el diente incisor fósil de la Formación San Sebastián. Arriba, imagen de microscopio electrónico de barrido mostrando la estructura microscópicas del incisor (píquenle a la imagen para que la vean más grande).  
El correo que nos envió Thomas fue muy alentador. Después de todo, resultó que nuestros fósiles si pertenecían a roedores caviomorfos, confirmando lo que ya sospechábamos. Ahora que sabíamos a que tipo de roedor pertenecían los dientes, y su importancia respecto a los orígenes de la fauna antillana, deseábamos obtener edades más precisas para las localidades. Para esto me comuniqué con mi amiga y colega Diana Ortega-Ariza, candidata doctoral de la Universidad de Kansas. Diana hizo su maestría en el Departamento de Geología en la Universidad de Puerto Rico (mi alma mater) donde estudió las distintas unidades calizas del norte de la isla, incluyendo las Calizas Lares. Como parte de su estudio ella obtuvo edades radiométricas utilizando isótopos de estroncio* preservados en los tubos calcíticos que servían de hogar a el bivalvo Kuphus incrassatus, el cual es uno de los fósiles más comunes en rocas del Oligoceno y Mioceno. Los resultados de ese estudio isotópico revelaron que las Calizas Lares se depositaron entre 27-24 millones de años atrás durante una era geológica llamada Chattiano la cual es la subdivisión más joven del Oligoceno**. Esa edad la podíamos entonces utilizar para estimar que la Formación San Sebastián, la cual se encuentra debajo de la Lares, tiene una edad mayor de 27 millones de años, en otras palabras, que esa formación se depositó durante el Rupeliano, la cual es la subdivisión más antigua del Oligoceno.
*Este tipo de análisis funciona de forma similar al Carbono 14, con la ventaja que nos sirve para fechar fósiles más viejos. Las edades máximas que se pueden obtener con Carbono 14 son de 50,000 años, mientras que el estroncio nos sirve para calcular fechas de hasta cientos de millones de años.
**Todo el periodo Oligoceno duró entre 33.9-23.0 millones de años atrás y tiene dos subdivisiones Chattiano y Rupeliano.

Orígenes de los vertebrados terrestres de las Antillas Mayores
Charles Darwin nunca estuvo en las Antillas. Aún así, la fauna de la región no pasó desapercibida y este incluso escribió en uno de sus libros más populares Voyage of the Beagle que "El caracter suramericano de los mamíferos antillanos parece indicar que este archipiélago estuvo en algún momento unido al continente sureño, y que subsiguientemente ha sido un área de subsistencia." Darwin, al igual que muchos otros antes y después de el, tenía razón respecto al lugar de origen de los mamíferos terrestres de las Antillas. Sin embargo, una de las preguntas más debatidas no ha sido el donde, sino el cuando llegaron los ancestros de los vertebrados terrestres de las Antillas.

Por años, el debate se ha centrado en dos hipótesis: una propone que estos llegaron a la región mediante múltiples eventos de dispersión a lo largo de los últimos 60 millones de años (e.g. Hedges et al., 1992), la otra, que llegaron alrededor del mismo tiempo durante un solo evento de dispersión (e.g. MacPhee e Iturralde-Vinent, 1995). La hipótesis de GAARlandia* propone que entre el Eoceno tardío y Oligoceno temprano las islas de Cuba, Española, y Puerto Rico junto con la Cresta de Aves formaron un puente terrestre casi continuo que los unió con el norte de Sur América (vean la figura abajo). Este puente terrestre, aunque de poca duración - probablemente estuvo formado entre 37.8-28.1 millones de años atrás - sirvió como un corredor para la dispersión de vertebrados terrestres hacia las masas terrestres que más tarde pasarían a ser las Antillas Mayores (para un resumen detallado de la evidencia geológica vean Iturralde-Vinent y MacPhee [1999]). Esto significa que, idealmente, deberíamos encontrar fósiles de mamíferos terrestres en depósitos del Eoceno tardío-Oligoceno temprano que representen a los mismos grupos que conocemos del Pleistoceno.
*Greater Antillean Aves Ridge land (MacPhee e Iturralde-Vinent, 1995)
Figura 1 de nuestra publicación. Aquí mostramos las distintas localidades (A) en Sur América donde se encuentran roedores fósiles del Eoceno y Oligoceno junto con la localidad de Domo de Zaza (DZ) en Cuba donde se han encontrado roedores del Mioceno temprano. En la figura B vemos las localidades de Puerto Rico, Río Guatemala (RG) y Calizas Lares (LL)C muestra la reconstrucción paleogeográfica de la región del Caribe durante el Eoceno tardío-Oligoceno temprano. Durante este periodo Cuba, La Española y Puerto Rico estaban unidos a la Cresta de Aves formando un puente terrestre conectando con el norte de Sur América.
El registro fósil de la región no nos ha fallado y nos ha dado algunos fósiles que apoyan esta segunda hipótesis. Uno de los primero que se descubrieron fue parte de un fémur (hueso del muslo) de un perezoso terrestre (megalonichido) en la Formación Juana Díaz, al suroeste de Puerto Rico, la cual se depositó alrededor de 31 millones de años atrás (MacPhee e Iturralde-Vinent, 1995). Los roedores que describimos en nuestro trabajo son similarmente antiguos, y juntos representan la evidencia más temprana de la presencia de perezosos terrestres y roedores caviomorfos en la región.

A lo largo del Mioceno se encuentran otros fósiles de vertebrados terrestres en la región de la Antillas Mayores, estos al igual que los del Oligoceno pertenecen a grupos que existieron durante el Pleistoceno. El Mioceno es el periodo geológico que le sigue al Oligoceno y que duró entre 23 a 5.3 millones de años. Estos fósiles del Mioceno incluyen perezosos terrestres, primates platirrinos, y hutías del Mioceno temprano de Cuba (MacPhee et al., 2003), una iguana y una boa del Mioceno temprano de Puerto Rico (MacPhee y Wyss, 1990), al igual que varias especies de ranas, guecos y lagartijas del Mioceno medio de La Española (e.g. De Queiroz et al., 1998; Daza y Bauer, 2012). Como podrán notar la lista de fauna del Mioceno es más larga y diversa que la del Oligoceno. La ausencia de algunos de estos en los depósitos más antiguos se podría atribuir al registro fósil, el cual es imperfecto, ó podría ser una ausencia real, implicando que estos organismos llegaron más tarde durante otros eventos de dispersión luego de la fragmentación de GAARlandia (Dávalos, 2004). Por ejemplo, basándose en el registro fósil y relojes moleculares, se ha estimado que los primates platirrinos llegaron a la región durante el Mioceno temprano (Cooke et al., 2011; Kay, 2014). En comparación, el grupo de sapos endémicos de las Antillas Mayores, los cuales se les conoce con el nombre científico de Peltophryne (y es el grupo que incluye el sapo concho), tienen un registro fósil que solo se extiende al Cuatenario (Pregill, 1981). Sin embargo utilizando relojes moleculares se ha estimado que los sapos Peltophryne llegaron a la región caribeña mucho antes, durante el Eoceno tardío-Oligoceno temprano lo cual sería consistente con la hipótesis de GAARlandia (Alonso et al., 2012).

Así que como pueden ver, la evidencia que tenemos hasta ahora apunta a un origen más complicado de los vertebrados terrestres de las Antillas Mayores y parece que ambas hipótesis han jugado un rol en el origen de los mismos. Además, como mencionamos brevemente en nuestro trabajo, no todo tiene que haberse dispersado de Sur América a las Antillas. Por ejemplo, los gaviales gryposuquinos pudieron utilizar ese puente terrestre para dispersarse del Caribe a Sur América. Para lograr entender mejor esta complicada historia se necesita hacer más trabajo y más descubrimientos en la región. Trabajar en los trópicos no es fácil por la extensa vegetación y limitada exposición de las rocas, sin embargo, la recompensa es grande cuando se hacen descubrimientos como los que acabamos de publicar.


Referencias

Ali, J. R. 2012. Colonizing the Caribbean: is the GAARlandia land-bridge hypothesis gaining a foothold? Journal of Biogeography 39:431-433.

Alonso, R., A. J. Crawford, and E. Bermingham. 2012. Molecular phylogeny of an endemic radiation of Cuban toad (Bufonidae: Peltoprhyne based on mitochondrial and nuclear genes. Journal of Biogeography 39:434-451.

Anthony, H. E. 1918. The indigenous land mammals of Porto Rico, living and extinct. American Museum of Natural History, Memoirs 1(2):324-435.

Antoine, P.-O., L. Marivaux, D. A. Croft, G. Billet, M. Ganerod, C. Jaramillo, T. Martin, M. J. Orliac, J. Tejada, A. J. Altamirano, F. Duranthon, G. Fanjat, S. Rousse, and R. Salas Gismondi. 2012. Middle Eocene rodents from Peruvian Amazonia reveal the pattern and timing of caviomorph origins and biogeography. Proceedings of the Royal Society B 279:1319-1326.

Biknevicious, A. R., D. A. McFarlane, and R. D. E. MacPhee. 1993. Body size in Amblyrhiza inundata (Rodentia: Caviomorpha), an extinct megafaunal rodent from the Anguilla Bank, West Indies: estimates and implications. American Museum Novitates 3079:1-25.

Dávalos, L. M. 2004. Phylogeny and biogeography of Caribbean mammals. Biological Journal of the Linnean Society 81:373-394.

Daza, J. D., and A. M. Bauer. 2012. A new amber-embedded sphaerodactyl gecko from Hispaniola, with comments on morphological synapomorphies of the Sphaerodactylidae. Breviora 529:1-29.

De Queiroz, K., Ling-Ru Chu, and J. B. Losos. 1998. A second Anolis in Dominican amber and the systematics and ecological morphology of Dominican amber anoles. American Museum Novitates 3249:1-23.

Hedges, S. B., C. A. Hass, and L. R. Maxson. 1992. Caribbean biogeography: molecular evidence for dispersal in West Indian terrestrial vertebrates. Proceedings of the National Academy of Sciences of the United States of America 89:1909-1913.

Iturralde-Vinent, M. A., and R. D. E. MacPhee. 1999. Paleogeography of the Caribbean region: implications for Cenozoic biogeography. Bulletin of the American Museum of Natural History 238:1-95.

Kay, R. F. 2014. Biogeography in deep time – what do phylogenetics, geology, and paleoclimate tell us about early platyrrhine evolution? Molecular Phylogenetics and Evolution In press.

MacPhee, R. D. E., and M. A. Iturralde-Vinent. 1995. Origins of the Greater Antillean land mammal fauna, 1: new Tertiary fossils from Cuba and Puerto Rico. American Museum Novitates 3141:1-31.

MacPhee, R. D. E., and A. R. Wyss. 1990. Oligo-Miocene vertebrates from Puerto Rico, with a catalog of localities. American Museum Novitates 2965:1-45. 

Pregill, G. 1981. Late Pleistocene herpetofaunas from Puerto Rico. University of Kansas Museum of Natural History Miscellaneous Publication 71:1-72.

Turvey, S. T., F. V. Grady, and P. Rye. 2006. A new genus and species of ‘giant hutia’ (Tainotherium valei) from the Quaternary of Puerto Rico: an extinct arboreal quadruped? Journal of Zoology 270:585-594.

Velez-Juarbe, J., T. Martin, R. D. E. MacPhee, and D. Ortega-Ariza. 2014. The earliest Caribbean rodents: Oligocene caviomorphs from Puerto Rico. Journal of Vertebrate Paleontology 34:157-163.


Woods, C. A., and F. E. Sergile (eds.). 2001. Biogeography of the West Indies: Patterns and Perspectives, second edition. CRC Press, Boca Raton, Florida, 608 pp.

Tuesday, January 14, 2014

The oldest Caribbean rodents

Its January 2014, and with it comes the first issue of the Journal of Vertebrate Paleontology. There are many interesting papers, including several ones on fossil marine mammals, which are a frequent subject in this blog. However this post will be about another paper in that same issue, where several of my colleagues and myself describe fossil rodents from Puerto Rico (Velez-Juarbe et al., 2014). These are not just any fossil rodents, they are the oldest evidence of caviomorph rodents in the Greater Antilles, and help us further understand the timing of arrival of terrestrial vertebrates to the region, an issue which has been the matter of debate for several decades now (keep reading).

An account of fossil rodents from Puerto Rico
It’s not the first time that fossil rodents have been found in Puerto Rico. In fact, it is well known that during the Pleistocene, Puerto Rico, Hispaniola, Cuba, Jamaica, and several of the Lesser Antilles, were home to endemic groups of rodents and other land mammals (Woods and Sergile, 2001). Many of these Pleistocene mammals were described in the early 1900’s (e.g. Anthony, 1918), although there have been some more recent discoveries as well (Turvey et al., 2006). Some of the rodents included giant forms, with some, like Amblyrhiza inundata, had body masses similar to those of the American black bear (Biknevicious et al., 1993)! 
During the Pleistocene Puerto Rico was home to about  two or three species of heteropsomine spiny rats (Echymyidae), and possibly two species of plate-tooth (Heptaxodontids). A third group that is still present in the region, are the Capromyids, but these, apparently, never reached Puerto Rico. All of these groups of rodents, are part of a larger, more inclusive group of rodents known as caviomorphs. caviomorphs are endemic to South America, their ancestors arriving from Africa about 54 million years ago (Antoine et al., 2012). Once in South America caviomorphs underwent an explosive radiation, spreading throughout the continent and as we now know, the Caribbean, early on in their evolutionary history. A similarly fast radiation also happened with South American primates (Platyrhines) (Kay, 2014), which share a comparable history to that of the caviomorphs.
This picture is from last week, when I revisited the Lares limestone site where i found the rodent incisor.
The new fossils from Puerto Rico consist of a couple of isolated incisors, one from the early Oligocene San Sebastian Formation (from this locality), and the other from the late Oligocene Lares Limestone (from this locality; also see picture above). When I found the first fossils back in 2005, I contacted Ross MacPhee, curator of mammals at the American Museum of Natural History, and who has had a long interest in the origins of the Greater Antillean land mammal fauna. Based on what we know about the fossil record of Puerto Rico, Ross and I suspected these incisors were from a caviomorph. However, we had a problem, as you can see in the picture below, isolated incisors are really hard to identify based only on external features, as they are very simple and lack the cusps, valleys and ridges that characterize their molars (this is true for nearly all mammals). We then thought about looking at the enamel microstructure, as it does preserved features that can be extremely useful in identifying isolated finds like ours. Studying the enamel microstructure usually involves making cross-sections of the teeth, the cut surfaces are polished, etched in a light acid, and observed with the aid of scanning electron microscope (SEM's). To do this, we decided to contact Thomas Martin at Universität Bonn, an authority on enamel microstructure. After recruiting Thomas, we sent him the fossils, and within several months, he emailed us back with the description and interpretation of the fossils. 
Lateral (left) and anterior (right) views of the San Sebastian caviomorph incisor. Here the fossil was still partially surrounded by rock and had not been sectioned to study the enamel microstructure. 
Figure 2 from our paper. Here we show the cross section outline and the enamel microstructure of the San Sebastian caviomorph (left column) and Lares caviomorph (right column).
Our fossils did belong to a caviomorph, thus confirming our initial suspicions and excitement about the discovery. Because of the importance of these fossils, we wanted to get as precise dates on the localities as possible, so I contacted my friend and colleague Diana Ortega-Ariza, a PhD candidate at the University of Kansas. For her masters at the University of Puerto Rico, Diana studied several of the limestone units in Puerto Rico, including the Lares Limestone. As part of her study, she obtained radiometric dates using isotopic signals preserved in the calcitic tubes that serve as home to the bivalve Kuphus incrassatus, which is commonly found in Oligocene through Miocene marine deposits. Based on the results she obtained, we are now able to say that the Lares Limestone was deposited between 27-24 million years ago during a geologic age called Chattian which is the youngest sub-division of the Oligocene. We could also used that timeframe to place deposition of the underlying San Sebastian Formation as older than 27 million years ago, or during the age known as Rupelian, the oldest subdivision of the Oligocene (the whole Oligocene period lasted between 33.9-23.0 million years ago).

Origins of the Greater Antillean land vertebrate fauna
“The South American character of the West Indian mammals seems to indicate that this archipelago was formerly united to the southern continent, and that it has subsequently been an area of subsidence.” Charles R. Darwin The Voyage of the Beagle

That is one of my favorite sentences in Voyage of the Beagle. Charles Darwin never visited the Greater Antilles, but he was right about where did the mammals of the region originated. However, one of the long-standing questions regarding the origins of Greater Antillean land vertebrates is not where, but when did they arrived. For years the debate has been whether they arrived to the region at different intervals throughout the Cenozoic (e.g. Hedges et al., 1992) or in tandem during a single dispersal event (e.g. MacPhee and Iturralde-Vinent, 1995). The GAARlandia* hypothesis postulates that during the late Eocene-early Oligocene the islands of Cuba, Hispaniola and Puerto Rico together with the Aves Ridge formed a continuous, or nearly continuous landspan connected to northern South America (see figure below). This landspan, even though short-lived - it probably lasted between about 37.8 to 28.1 million years ago, or less - would have served as a corridor for the dispersal of land vertebrates into what eventually became the Greater Antilles (see Iturralde-Vinent and MacPhee [1999] for a very detailed overview of the geologic evidence). Ideally, we should be finding fossils representing the different groups of Pleistocene and recent Greater Antillean land vertebrates in late Eocene-early Oligocene deposits, and we do, at least in part. 
*Greater Antillean Aves Ridge land (MacPhee and Iturralde-Vinent, 1995)
Figure 1 from our paper. Here we show the various localities (A) in South America where Eocene and Oligocene rodents are known and Domo de Zaza (DZ) in Cuba, where early Miocene rodents are known. In are the localities in Puerto Rico, Río Guatemala (RG), and Lares Limestone (LL). is the paleogeographic reconstruction of the Caribbean region during the late Eocene-early Oligocene. During this time Cuba, Hispaniola and Puerto Rico were joined with the Aves Ridge (C) forming a nearly continuous landspan connected to northern South America.
The fossil record of the region has so far provided a few clues supporting this hypothesis as well. One of the first ones found was a femur (leg bone) of a ground sloth (megalonychid) in the early Oligocene Juana Diaz Formation in southwestern Puerto Rico which was deposited about 31 million years ago (MacPhee and Iturralde-Vinent, 1995). The rodents described in our work are just as old, and together they are the earliest evidence of these two groups in the region.
Throughout the Miocene (the period between 23-5.3 million years ago) in the Greater Antilles there are other occurrences of terrestrial vertebrates that were present during the Pleistocene and some which are even still around today. There are early Miocene sloths, rodents and primates from Cuba (MacPhee et al., 2003), a boa and an iguana from the early Miocene of Puerto Rico (MacPhee & Wyss, 1990), as well as a number of frogs, geckos and anoles from the middle Miocene of Hispaniola (e.g. De Queiroz et al., 1998; Daza and Bauer, 2012). The absence of some of these groups in Oligocene rocks in the Greater Antilles could be due to the scarcity of the fossil record, or could indeed be real, implying that some of these groups arrived by random overwater dispersal after fragmentation of GAARlandia (Dávalos, 2004). For example, based on fossils and molecular data, primates have an estimated time of arrival to the region during the early Miocene (Cooke et al., 2011; Kay, 2014). On the other hand, toads of the genus Peltophryne which is endemic to the region, are known only from Pleistocene deposits (Pregill, 1981) but molecular estimates place their split from their closest relatives during the late Eocene-early Oligocene (Alonso et al., 2012).
So, as you can see, the evidence seems to point to a more complex origin of the Greater Antillean land vertebrates and does not seem to favor one over the other. Also, not everything had to have come from South American to the Greater Antilles. As we mentioned briefly in our paper, organisms that were present in the Antilles, such as gryposuchine gavials, could have used that same corridor to disperse to the southern continent. Further unraveling of this complex history can only be achieved by more fieldwork and discoveries in the Greater Antilles. 

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