Dominican Republic, part II

Hello there dear readers. Today I bring you the second installment of this series. Like I mentioned in the previous post we moved from the Albian to the Miocene.

The Miocene of the Dominican Republic is probably best known for the amber deposits. These have not only produced beautiful amber, which is used in jewelry, but also a number of fossil taxa have been described based on remains entombed in the amber. These fossils consist mostly of invertebrates, however, several vertebrates have been described as well, including anoles, frogs, and an insectivore. This gives us a glimpse into the smaller fauna that inhabited the island during the Miocene, in contrast to what is known from the same age in Puerto Rico and Cuba where most of the vertebrate fossils of that age consist of larger animals (sloths, rodents, primates, sirenians, crocodylians and turtles). A good summary of the vertebrates known from the Tertiary of the Greater Antilles can be found in table 1 of MacPhee et al. (2003) (free download here).

One of our first excursions into the Miocene was on a Wednesday afternoon. We spent the morning collecting the last samples from one of the outcrops of the Hatillo Limestone and some rudists from near the entrance of a cave in that same limestone unit. We then headed west towards the Monte Plata Province where we knew there was a new road cut exposing units of Miocene age.

The first outcrop of the Miocene that we visited. This is actually one of the best Miocene outcrops I've seen in a while.

By the time we got there it was near the middle of the afternoon, which meant we didn’t had much sunlight left. We stopped in the first large outcrop we saw (Picture above) and it was worth it! We quickly started finding turtle shell fragments, and occasionally we saw cross sections of turtle shells that went into the outcrop. I decided to collect one of these. It was not big, and as I found out after prep work, not too good (see pictures below). However, it was worth going there, as there is more material to be found and collected!

Top left: me overturning the jacket with the turtle shell remains; top right: the jacket back in the lab; bottom left: view of the jacket when it was first opened; bottom right: the fossil turtle remains. (Click on the image to see the larger version.)

Several days latter we had some free time, so we decided to go back to this area. This time we drove in from the north, driving through Los Haitises where we saw beautiful karst topography. Our time was limited in this occasion as well, but we made the most out of it.

The second outcrop we visited, smaller than the first one, but still interesting. Alvin Bonilla for scale.

We hit a couple of outcrops (Picture above) where we found the usual (at least what usual for the Tertiary of Puerto Rico and Cuba); croc teeth, turtle shells and sirenians (which was what I was really after). The sirenian material was in a concretion, half of it which had fallen off the outcrop so it made it easy to collect. The other part is still in the outcrop and I hope to go back sometime in the spring to collect it. Actually, if it wasn't for Alvin, who saw it, I almost stepped on the part that had fallen trying to get a better view of the part still in the outcrop. After some prep work on the sirenian material it turned out to be mostly ribs, and a few vertebrae. However, even if it is a little frustrating, it will nonetheless be an important contribution to the understating of sirenian distribution during the Miocene of the Caribbean and demonstrates that there is probably more, better material waiting to be collected!

Top left: me collecting the concretion with the sirenian remains; top right: view of the jacket in the lab; bottom left: the jacket, recently opened; bottom right: the concretion, as of now, still need some prep work to be done. Oysters (near the center of the concretion and to the right) used some of the ribs as hardground. (Click on the image to see the larger version.)

MacPhee, R. D. E., M. A. Iturralde-Vinent & E. S. Gaffney. 2003. Domo de Zaza, an Early Miocene vertebrate locality in south-central Cuba, with notes on the tectonic evolution of Puerto Rico and the Mona Passage. American Museum Novitates 3394:1-42.

A river runs through an Oligocene sea: parte III

So, keeping up with stuff I’ve collected at the Río G locality (for previous entries about this locality go here and here), I bring you some of the sirenian material that I collected last January. It is not much, but it adds to stuff I’ve collected previously (as you’ll see).

In tropical regions, a lot of times, good outcrops are along riverbanks. Puerto Rico is no exception and that is why the Rio G locality is so good, the exposure is kept “fresh” because of the nearly constant river erosion. Of course the drawback is that fossils are also lost if nobody visits these type of localities at least once a month or after big rainstorms (at least that’s what I used to do).

The picture above shows one such example. I spotted this bone fragment on the wall, very close to water level (a little more than half a meter). And, as you can see in the inset, the surface facing away from the rock looks freshly broken. I did collect the fossil, and unsuccessfully looked for additional fragments nearby.

Above is the picture of the fossil, in dorsal view (anterior towards the top), and the interpretative drawing. As it turns out the fossil was part of a sirenian skull. What was left of the fossil, is the anterior part of the frontals (Fr) and the nasals (N), the supraorbital processes of the frontals are missing. The convex frontal roof and shallow nasal incisure (the concave area between the frontals) are some characters that identify this fossil as belonging to a halitheriine dugongid. In fact, it is very similar to the same part of a much more complete skull that I collected from that locality several years ago (see below).

The figure above shows the more complete skull. A and B show a close-up dorsal view (anterior towards the top) of the area that was preserved in the fragmentary fossil. C is a dorsal view of the skull (anterior to the right) with the outline of the enlarged area in A. As you can see the frontal and its relationship with the nasals look much the same as the fragmentary fossil. It also displays a shallow nasal incisure at the posterior end of the mesorostral fossa (MRF) and convex frontal roof. In the more complete skull the premaxilla (Pmx) partially cover the nasals and the supraorbital processes (SOP) are preserved. (In A and C it is missing the left nasal process of the premaxilla which is loose and needs to be re-attached, but I was able to put it in the drawing).

I’m pretty certain that the fragmentary fossil belongs to the same species as the more complete specimen; it was a pity that part of it was lost previous to it being found. If you’re well acquainted with extinct sirenians, you can probably guess what genus this skull belongs to. So, go ahead and make a guess!

Chevron weirdness in a sirenian

Following up on a comment I made over at Updates from the Vertebrate Paleontology Lab, I here bring you pictures of an unusual sirenian chevron from the Late Oligocene of Puerto Rico.

These fossils were collected as part of a partial articulated postcranium which I mentioned here. Chevrons, also known as hemal arches, are (normally) V-shaped* bones, consisting of two rami that meet ventrally, hence the shape. They protect blood vessels.

*More like Y-shaped due to the length of the symphysis in some specimens.

In the picture above you can see left lateral views of two of the chevrons I collected with that specimen. The chevron on the left seems to represent fused chevron 2 + 3 (missing part of its ventral edge), and the other one is chevron 4, which is normal.

Here is a posterior view, again, the one on the left likely represents fused chevron 2 + 3, and on the right is chevron 4, which is missing the right ramus.

To me this seems to have been a developmental anomaly rather than occurring due to an injury. Apparently, this is a first, at least for Sirenia! As I mentioned above these are part of an articulated partial postcranium, belonging to a new dugongine taxon from the Late Oligocene of northern Puerto Rico (you can see the skull here).

So, leave a comment, let me know what you think about this unusual bone!

Nuevos sirénidos del Eoceno

Este año ha visto la publicación de dos artículos describiendo sirénidos del Eoceno (55.8-33.9 millones de años atrás [ma]). En adición a esos, otras dos especies más fueron descritos del Mioceno, los cuales mencioné en una entrada pasada. Creo que no está mal, cuatro nuevas especies de sirénidos fósiles en un año, especialmente para un grupo que hoy día solo cuenta con cuatro especies (tres de manatí y el dugong). El conocimiento sobre la diversidad de sirénidos en el pasado sigue aumentando a muy buen ritmo.

Durante el Eoceno existían al menos tres de las cuatro familias de sirénidos que se conocen, Prorastomidae (que incluyen los miembros más primitivos), Protosirenidae y Dugongidae. El registro fósil de los Trichechidae (la cuarta familia, que es la que incluye al manatí) es más escaso y hasta donde sabemos, estos se originaron durante el Oligoceno Tardío (28.4-23 ma). Los nuevos fósiles del Eoceno representan a los Prorastomidae y los Dugongidae.

Un protosirénido de India

En una entrada pasada mencioné que India es uno de los lugares más prolíficos en términos de hallazgos de fósiles de sirénidos. No debe sorprendernos que otra especie adicional ha sido descrita. Ashokia antiqua Bajpai et al. 2009 representa un nuevo género y especie de protosirénido del Eoceno Medio temprano (Lutetian: 48.6-40.4 ma). Esta nueva especie se distingue por una combinación de caracteres primitivos y derivados, morfológicamente se acerca más a otro protosirénido de Libya, aún sin describir (Bajpai et al., 2009). Ashokia se distingue de otros prorastómidos al tener una cresta sigmoidal prominente, la apertura auditiva externa más ancha, el puente zigomatico-orbital elevado, el borde del exoccipital más delgado y poseer un proceso zigomático del hueso temporal que disminuye en grosor gradualmente en dirección al rostro.

En la figura arriba Ashokia antiqua en vista lateral (foto e ilustración hechas por mi).

Este no es el primer prorastómido que se conoce de India. Bajpai et al. (2006) refirieron un cráneo incompleto al género Protosiren sp., este, al igual que Ashokia, provienen de la Formación Harudi. Otros sirénido del Eoceno indio son Eotheroides babiae y Eosiren sp. también de la misma formación, estos sin embargo son dugónguidos (Bajpai et al. 2006). Los protosirénidos se distinguen de todas otras especies conocidas de sirénidos por ciertas características craneales y por tener las epífisis* cartilaginosas, incluso en los adultos (Sickenberg, 1934; Zalmout et al., 2003; Bajpai et al. 2006). Aún no se han reportado protosirénidos en rocas del Oligoceno, por lo que parece que este grupo fue uno exclusivo del Eoceno.

*Los extremos de los huesos.

Un dugónguido de Madagascar

Poco se conoce de los animales que habitaron Madagascar en el pasado. No fue hasta en años recientes que se comenzó a descubrir fósiles de los antiguos pobladores de esta isla, especialmente durante el Mesozoico y Cenozoico. Uno de los descubrimientos más recientes es el cráneo de un dugóngido proveniente de estratas del Eoceno Medio. Estos han sido descritos por Samonds et al. (2009) quienes han identificado el fósil como una especie nueva del género Eotheroides. E. lambondrano Samonds et al. 2009 pertenece a un género de dugónguidos que están se han encontrado en rocas del Eoceno de Egipto e India (Domning, 1996; Bajpai et al., 2006), esta nueva especie es la única dentro de ese género que posee el rostro completo, permitiéndonos conocer su morfología en mayor detalle. Único entre otras especies de Eotheroides por la morfología distinta de los huesos nasales, tener procesos supraorbitales bien desarrollados y el puente zigomatico-orbital de la maxilla corto, también se distingue de otras especies de dugonguidos al poseer una fórmula dental primitiva.

En la figura arriba Eotheroides lambondrano en vista lateral, escala = 4 cm (compuesto de fig. 3A y 4A de Samonds et al., 2009).

El largo del cráneo de E. lambondrano es de 270 mm (~10 pulg.), haciendo de este uno de los dugones de menor dimensión que se conocen. Posiblemente se acercaba en tamaño a las especies de dugónguino Nanosiren los cuales tenian dimensiones similares y cual largo total de cuerpo se estima en alrededor de 2 metros (~6’6”) (Domning & Aguilera, 2008), lo cual es considerado pequeño para un sirénido adulto. Asi que según se ha escrito en otros lugares (como en este artículo de National Geographic) puede que la nueva especie de Madagascar sea el dugónguido más pequeño que conocemos.

La importancia de estos nuevos fósiles resta en que demuestran cuan diverso eran los sirénidos durante el Eoceno. El nuevo protosirénido añade otra especie a un interesante grupo de sirénidos, que incluso se ha postulado como el grupo que dio origen a los Trichechidae. El hallazgo de Eotheroides lambondrano en Madagascar nos muestra cuán amplia era la distribución de este género y de los dugónguidos durante el Eoceno; en adición nos puede ayudar a entender la evolución de estos en la región del Tethys. Y por supuesto, aquellos de nosotros que están al tanto de lo que ocurren en el mundo de la paleosirenología saben que aún quedan más por ser descritos, asi que pendientes!

Otras entradas sobre sirénidos

Prep work: update II and a note on sirenian periotics

Prep work: update

Sirenian diversity in the past

De la tierra al agua (English version here)

Domningia and other Indian sirenians

What’s wrong with the hands of Steller’s sea cow

Bajpai, S., D. P. Domning, D. P. Das & V. P. Mishra. 2009. A new middle Eocene sirenian (Mammalia, Protosirenidae) from India. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 252/3:257-267.

Bajpai, S., J. G. M. Thewissen, V. V. Kapur, B. N. Tewari & A. Sahni. 2006. Eocene and Oligocene sirenians (Mammalia) from Kachchh, India. Journal of Vertebrate Paleontology 26(2):400-410.

Domning, D. P. 1996. Bibliography and index of the Sirenia and Desmostylia. Smithsonian Contributions to Paleobiology 80:1-611.

Domning, D. P. & O. A. Aguilera. 2008. Fossil Sirenia of the West Atlantic and Caribbean region. VIII. Nanosiren garciae, gen. et sp. nov. and Nanosiren sanchezi, sp. nov. Journal of Vertebrate Paleontology 28(2):479-500.

Samonds, K. E., I. S. Zalmout, M. Irwin, D. W. Krause, R. R. Rogers & L. L. Raharivony. 2009. Eotheroides lambondrano, new Middle Eocene seacow (Mammalia, Sirenia) from the Mahajanga basin, northwestern Madagascar. Journal of Vertebrate Paleontology 29(4):1233-1243.

Prep work: update II and a note on sirenian periotics

This has been a long hiatus! I’ve been really busy doing some more prep work on the Puerto Rican Dioplotherium and the Yucatán skull (another new species of dugongine). On top of that I’ve been preparing a couple of manuscripts describing some sirenian remains from PR, which I hope to submit sometime next year.

The subject of this post is to show you more of the Puerto Rican Dioplotherium, which was featured on the previous post. I have done additional prep work on the left squamosal, which was detached from the skull.

The composite picture above shows the skull as it was back in 2006 (top picture) and an outline of the enlarged area below. All that was visible of the sqamosal were the zygomatic arch, post-tympanic process and the mastoid part of the periotic (bottom picture). It was exciting knowing that part of the ear bones were preserved, even if it was only the periotic. Fortunately I got more than I bargained for.

In this figure we see the squamosal, now free of matrix, in lateral (A) and medial (B) views. Notice that the tympanic bone was preserved as well as the periotic. And there is more!

Additional removal of matrix revealed the three auditory ossicles, in articulation! The picture below shows a posteroventral view into the middle ear (anterior is to the right, medial towards the top of the picture). This is really neat as these bones are easily lost in most fossils (they are, apparently, missing on the right side of the skull).

A little on sirenian periotics

The periotic can be divided into three parts tegmen tympani, pars mastoidea and pars petrosa. The latter can be subdivided into pars canalicularis and pars cochlearis (see picture above) (Robineau, 1969). In the pars cochlearis, the structure labeled perilymphatic foramen, is uniquely found in (most) sirenians, (most) proboscideans and Arsinoitherium (hinting at their tethytherian affinity?). The homologous structure in other mammals consists of two openings known as the fenestra cochleae (rotunda) and aqueductus cochleae (Court, 1994).

The occurrence of a perilymphatic foramen in some tethytheres (I’m not sure what is the condition in desmostylians) seems to indicate that it might be a unique derived character of the group. Nonetheless, when we look at the fossil record, primitive proboscideans (Phosphatherium escuilliei) and sirenians (Prorastomus sirenoides) do have fenestra cochleae and aqueductus cochleae (Gheerbrant et al., 2005; Court, 1990; Savage et al., 1994). Meaning that this condition is homoplasic in tethytheres (Court, 1994, Gheerbrant et al. 2005). Whether resulting from multiple origins or multiple reversals, I still think it is an interesting characteristic that is found in at least some tethytheres.

Previous post about sirenians:

Prep work: update

Sirenian diversity in the past

De la tierra al agua (English version here)

Domningia and other Indian sirenians

What's wrong with the hands of Steller's sea cow

Court, N. 1990. Perotic anatomy of Arsinoitherium (Mammalia, Embrithopoda) and its phylogenetic implications. Journal of Vertebrate Paleontology 10(2):170-182.

Court, N. 1994. The periotic of Moeritherium (Mammalia, Proboscidea): homology or homoplasy in the ear region of Tethytheria McKenna, 1975? Zoological Journal of the Linnean Society 112:13-28.

Gheerbrant, E., J. Sudre, P. Tassy, M. Amaghzaz, B. Bouya and M. Iarochène. 2005. Nouvelles données sur Phosphatherium escuilliei (Mammalia, Proboscidea) de ‘Éocène inférieur du Maroc, apports à la phylogénie des Proboscidea et des ongulés lophodontes. Geodiversitas 27(2):239-333.

Robineau, D. 1969. Morphologie externe du complexe osseux temporal chez les sireniens. Mémoires du Muséum National d’Histoire Naturelle, Série A, Zoologie 60(1)-1-32.

Savage, R. J. G., D. P. Domning and J. G. M. Thewissen. 1994. Fossil Sirenia of the west Atlantic and Caribbean region. V. The most primitive known sirenian, Prorastomus sirenoides Owen, 1855. Journal of Vertebrate Paleontology 14(3):427-449.

Prep Work: Update

Wow! It’s been a while since I posted something. Working with some of the material collected back in August when I went to Puerto Rico to do fieldwork with my advisor has kept me busy. In addition, I had no computer for a while, just after posting the previous post, my computer’s hard drive died! Luckily, not much was lost.

Back in June I wrote about some prep work I had been doing on a sirenian skull from Puerto Rico. Well I am glad to say that four years after I collected said skull (in 2005), it is nearly done! So, here are some pictures, from the time it was collected until now.

In the picture above (from 2005) I am in the outcrop with my hand next to where the fossil is. This is a Late Oligocene limestone unit from northern Puerto Rico.

Here is a dorsal and right lateral view of how the fossil looked around 2006 (and actually it looked like that for the last 2 years). You might notice that on the top picture there is a bone floating in the matrix next to the braincase, this is the left squamosal, which is disarticulated.

Here is how the fossil looks like now (2009), with most of the matrix gone and the left squamosal removed. Beautiful, don't you think?!

If you know something about sirenians, you might have noticed that this is a dugongine (large tusks [broken, unfortunately], thickened supraorbital process of frontal, etc.). It is actually quite similar to Dioplotherium manigaulti from the Early Miocene of South Carolina and Florida (Cope, 1883; Domning, 1989). Nonetheless, the Puerto Rican skull is older, Late Oligocene, and it also has some primitive characters that sets it apart from D. manigaulti. This skull along with another one from the same locality make up an important part of my thesis. Fortunately, some postcranial material that was collected this summer, from the same outcrop and same unit, is referable to this taxon. This material also displays differences from other known sirenian postcrania. Pretty cool stuff!!

Previous post about sirenians:

Sirenian diversity in the past

De la tierra al agua (English version here)

Domningia and other Indian sirenians

What's wrong with the hands of Steller's sea cow

Cope, E. D. 1883. On a new extinct genus of Sirenia from South Carolina. Proceedings, Academy of Natural Sciences of Philadelphia 1883:52-54.

Domning, D. P. 1989. Fossil Sirenia of the West Atlantic and Caribbean region. II. Dioplotherium manigualti Cope, 1883. Journal of Vertebrate Paleontology 9:415-428.

Sirenian diversity in the past

Its been quiet here for a while as I’ve been busy working on the preparation of two sirenians skull, as well as getting ready for the upcoming field season.

It’s also been a while since I wrote something about sirenians so, here it goes.

Living sirenians can be divided into two families, Trichechidae (manatees) and Dugongidae (dugongs). Most people are probably more familiar with the manatees, after all, there are three species, West Indian, Amazonian and African, whereas there is only one species of dugong. The geographic distribution of extant sirenians is such that there is mostly no overlap between the different species. As the only living herbivorous marine mammals, it might be that by living in separate regions it reduced the chances of competing for the same resources (i.e. seagrasses). But what about in the past, what does the fossil record of sirenian tells us about their paleoecology.

When we look at the fossil record, sirenians were much more speciose, including multispecies communities in some regions (Domning, 2001). Now lets look at one good example.

The Late Oligocene of Florida

The Late Oligocene sirenian fauna of Florida includes at least three species of dugongids*. The dugongines, Crenatosiren olseni and Dioplotherium manigaulti, and the halitheriine Metaxytherium sp. (Domning, 1989, 1997, 2001). (See illustration below).

*The family Dugongidae includes three subfamilies: Dugonginae, Halitheriinae & Hydrodamalinae.

Illustration of known Late Oligocene sirenians from Florida (all at the same scale). Top, Crenatosiren olseni (modified from Domning, 1997); middle, Dioplotherium manigaulti (from Domning, 1989); bottom, Metaxytherium sp. (this last drawing based on a very similar skull from Puerto Rico, tusks not preserved, but presumed to be small as in the Fl specimen). The numbers in the circles are the degrees of rostral deflection. Mandibles absent in the middle and bottom specimens.

These three species, as you can see, differ in size, and to a lesser degree in rostral deflection. Also different from each other is the size of their tusks, increasing in size from Metaxytherium - C. olseni - Dioplotherium manigaulti. Taken as a whole, these differences (specially tusks size) could be indicators of different feeding habits, with small-tusked sirenians feeding of small rhizomes* and large-tusked sirenians feeding on larger ones (Domning, 2001; Domning & Beatty, 2007). Dugongids most likely used their tusks as a tool to dig out the rhizomes, with the most extreme specialization observed in the dugongines, including very large blade-like tusks as well as cranial adaptations that seemed to have help withstand the forces exerted when digging (Domning & Beatty, 2007).

*Rhizomes = the nutrient-rich, underground stems of seagrasses.

Other examples of sirenian multispecies communities are found in the Early Oligocene of Puerto Rico and the Early Miocene of India, among others (more on this sometime in the future). In addition, in the Pacific, sirenians were not the only herbivorous marine mammals. In the northern Pacific region, sirenians seem to have shared their resources with the desmostylians (see picture below), an interesting (and bizarre) group of mammals that lived from the Oligocene to the Miocene and were presumably feeding and spending time in the marine realm (Domning et al., 1986; Inuzuka et al., 1994). Whereas, in the southeastern Pacific, fossils of aquatic sloths (Thalassocnus spp.) have been found in the same formations as sirenians (Muizon & McDonald, 1995; Canto et al., 2008; Muizon & Domning, 1985; Bianucci et al., 2006; Domning & Aguilera, 2008).

Mounted cast of Palaeoparadoxia tabatai taken at the AMNH.

So, why is it so different in modern times, why do we see such a reduced diversity of sirenians and/or lack of any other herbivorous marine mammals? There has been, apparently, little change in the marine seagrass communities since the Eocene, so what happened? The answers for these and other questions could be answered with more fossils and more research. For now, we can certainly say that, like their close relatives, the proboscideans (elephants), sirenians are the last remnants of a once much more diverse group of animals.

References

Bianucci, G., S. Sorbi, M. E. Suárez & W. Landini. 2006. The southernmost sirenian record in the eastern Pacific Ocean, from the Late Miocene of Chile. Comptes Rendus Palevol 5:945-952.

Canto, J., R. Salas-Gismondi, M. Cozzuol & J. Yáñez. 2008. The aquatic sloth Thalassocnus (Mammalia, Xenarthra) from the Late Miocene of north-central Chile: biogeographic and ecological implications. Journal of Vertebrate Paleontology 28(3):918-922.

Domning, D. P. 1989. Fossil Sirenia of the West Atlantic and Caribbean region. II. Dioplotherium manigaulti Cope, 1883. Journal of Vertebrate Paleontology 9:415-428.

Domning, D. P. 1997. Fossil Sirenia of the West Atlantic and Caribbean region. VI. Crenatosiren olseni (Reinhart, 1976). Journal of Vertebrate Paleontology 17:397-412.

Domning, D. P. 2001. Sirenians, seagrasses, and Cenozoic ecological change in the Caribbean. Palaeogeography, Palaeoclimatology, Palaeoecology 166:27-50.

Domning, D. P. & O. A. Aguilera. 2008. Fossil Sirenia of the West Atlantic and Caribbean region. VIII. Nanosiren garciae, gen. et sp. nov. and Nanosiren sanchezi, sp. nov. Journal of Vertebrate Paleontology 28:479-500.

Domning, D. P. & B. L. Beatty. 2007. Use of tusks in feeding by dugongid sirenians: observations and tests of hypotheses. Anatomical Record 290:523-538.

Domning, D. P., C. E. Ray & M. C. Mckenna. 1986. Two new Oligocene desmostylians and a discussion of Tethytherian systematics. Smithsonian Contributions to Paleobiology 59:1-56.

Inuzuka, N., D. P. Domning & C. E. Ray. 1994. Summary of taxa and morphological adaptations of the Desmostylia. Island Arc 3(4):522-537.

Muizon, C. de & D. P. Domning. 1985. The first records of fossil sirenians in the southeastern Pacific Ocean. Bulletin du Muséum National d’Histoire Naturelle (Paris) (4)7, Sect. C, no. 3:189-213.

Muizon, C. de & H. G. McDonald. 1995. An aquatic sloth from the Pliocene of Perú. Nature 375:224-227.

A day in the field, Tertiary

This time our field area is in northern Puerto Rico. We decided to visits a couple of outcrops of the Late Oligocene Lares Limestone. If the name of the formation sounds familiar you either know about the geology of Puerto Rico or, have read about it on a previous post.

One of these localities (see picture below), I have visited at least since 2000, and up until very recently, we thought that the only formations present there were the Early Oligocene San Sebastián Formation and the overlying Lares Limestone. Now, thanks to new information regarding the stratigraphy of the Tertiary limestones of the north coast of Puerto Rico (Ortega Ariza, 2009), we know that in this locality, overlying the Lares Ls, there are also units of the Montebello Limestone. The age of the Lares Limestone and Montebello Limestone were designated as Late Oligocene – lower Early Miocene and upper Early Miocene, respectively (Seiglie & Moussa, 1984). New data, using strontium isotopes obtained from tubes of the pelecypod Kuphus incrassatus, seems to indicate, instead, that both formations span the Late Oligocene (Johnson et al., 2006; Ramírez et al., 2006; Ortega Ariza, 2009). If this is correct (more samples need to be run, hint, hint!!) I will like this outcrop even more (sorry, can't hide my love for the Oligocene)!!

Here's the one of my favorite outcrops, where the Lares and Montebello limestones are exposed. The arrow points to a sirenian fossil that is yet to be collected.

Of course, what I’ve been mostly searching in these localities are sirenian remains, but like I mentioned on that previous post, other vertebrates have also been collected. Interestingly, the best sirenian remains have been collected from the upper Lares Limestone, with a total (so far) of two skulls, and a set of nine articulated vertebrae (see picture below). There are more fossils but those will be collected in due time. As for the sirenian skulls, well, they are an important part of my thesis work and I will discuss them at some point in the future.

Some articulated sirenian vertebrae, these have already been collected. This is an earlier picture, there were three more vertebrae behind the one labeled Ca1, the ones anterior to L3 were collected earlier.

References

Johnson, C. C., W. R. Ramírez, L. R. Mark, S. Y. Hernandez, E. A. Barrow, M. Hegewald & J. Velez. 2006. Oligocene reef deposits linked to OPD site 999 with strontium isotope stratigraphy. Geological Society of America Abstracts with Program 38:557.

Ortega Ariza, D. L. 2009. Establishing a high resolution sequence stratigraphy and sea-level curve for Tertiary limestones, Puerto Rico. M.S. thesis, University of Puerto Rico, Mayagüez, Puerto Rico, 132 pp.

Ramírez, W. R., C. C. Johnson, M. Martínez, M. C. Torres & V. Ortiz. 2006. Strontium isotope stratigraphy from Kuphus incrassatus, Cenozoic limestones, Puerto Rico. Geological Society of America Abstracts with Program 38:90.

Seiglie, G. A. & M. T. Moussa. 1984. Late Oligocene-Pliocene trangressive-regressive cycles of sedimentation in northwestern Puerto Rico. American Association of Petroleum Geologist Memoir 36:89-95.

From land to sea

Some of the living marine mammals, like cetaceans, sirenians and pinnipeds* are so well adapted to a life in water that it might be difficult for us to relate them to their closest terrestrial relatives. However, we do know that the oldest members of these groups were indeed terrestrial. Here I’ll discuss some of the evidence known so far.

*(cetaceans = whales & dolphins; sirenians = manatees & dugongs; pinnipeds = seals, walruses & sea lion).

Cetaceans

Modern whales can be divided into two groups, odontocetes and mysticetes. Odontocetes are characterized for having teeth and using echolocation; mysticetes are characterized for having baleen instead of teeth (there are other adaptations that I won’t discuss now). Some examples of odontocetes are orcas and bottlenose dolphins; mysticetes include blue whales and right whales.

Based on molecular evidence, whales evolved from artiodactyls – a group that includes pigs, hippopotamus, camels, cows, lambs, etc – (Graur & Higgins, 1994; Shimamura et al., 1997), whereas, for a long time, morphological studies used to indicate a close relationship with mesonychids – a group of extinct terrestrial carnivores – (Luo & Gingerich, 1999). In part, the reason for this disagreement about the origin of whales was that the oldest fossils of cetaceans consisted of forms that were already completely adapted to a life in the water, or were only known from crania. This has already been resolved.

In 2001, two groups of paleontologist published papers where they described primitive cetaceans, including parts of the postcranium that corroborated an artiodactyl relationship (Gingerich et al., 2001; Thewissen et al., 2001). Gingerich and his team found remains of Artiocetus clavis and Rhodocetus balochistanensis, whereas the Thewissen team described Ichthyolestes pinfoldi and Pakicetus attocki (illustration above of Pakicetus by Carl Buell, taken from the Thewissen Lab webpage); the remains included one of the ankle bones, the astragalus, which was key to determine that cetaceans evolved from artiodactyls. All these fossils were found Middle Eocene (49-41 million years ago) deposits. More recently, Thewissen et al. (2007) describe postcranial material of the primitive artiodactyl, Indohyus, and show that it was an animal with aquatic adaptations, providing additional evidence about the origin of cetaceans. (Go here for a magnificent reconstruction of Indohyus).

Sirenians

The closest living relatives of manatees and dugongs are elephants, this relationships is supported by both, molecular and morphological evidence (Seiffert, 2007; Tabuce et al. 2007). Sirenians originated in northern Africa about 54 million years ago, where they last shared a common ancestor with proboscideans (elephants). Interestingly, the most primitive sirenian fossils have been found in Jamaica, which demonstrate that, very early, they seem to have been well adapted for life in an aquatic environment. For a long time, the most primitive sirenian known was Prorastomus sirenoides found in Jamaica in deposits that are between 51-49 million years old (Owen, 1855; Savage et al., 1994). Unfortunately, the postcranium was and it is still mostly unknown.

Another sirenian from Jamaica, found in slightly younger deposits – 49-45 million years old – was described by Domning (2001). The remains of this new sirenian, named Pezosiren portelli (illustration above from Domning, 2001), include cranial and postcranial material. The postcranial material include fore and hind limbs, pelvis and most of the vertebral column; all together, these indicate that Pezosiren was able to support its own weight on land, but at the same time, it had aquatic adaptations such as pachyosteosclerotic (enlarged & dense) ribs and in the cranium, retracted external nares (Domning, 2001). The combination of characters imply that Pezosiren spent time, both, in and out of the water.

Pinnipeds

Morphological and molecular studies show that pinnipeds belong to a group of mammals called arctoids (Deméré et al., 2003), that, along with pinnipeds, includes bears (ursids), weasels (mustelids), raccoons (procyonids) and skunks (mephitids), among others. Nonetheless, the origin of pinnipeds from one of these arctoids is not clear, and different studies place pinnipeds as originating from a common ancestor with mustelids or with ursids (Deméré et al., 2003). Other experts in the field support a multiple origin for pinnipeds, with seals sharing a common ancestor with mustelids and sea lion and walruses with ursids (Uhen, 2007 and references therein). Anyways, whatever is the group from which pinnipeds originated – this can only be resolved by finding more fossils – it is well known that these originate from a terrestrial ancestor. Interestingly, pinniped fossils that show a transitional morphology had not been found until recently.

The discovery of Puijila darwini in lacustrine sediments deposited between 23-21 million years ago, gives us an idea about the morphology of the earliest pinnipeds (Rybczynski et al., 2009). Although fossils of yet even older pinnipeds, such as Enaliarctos tedfordi and E. barnesi, have been found in rocks that date between 28.5-23.8 million years in Oregon (Deméré et al., 2003), these already show full adaptations to a life in the marine realm like those observed in modern taxa; this means that pinnipeds must have evolved previous to that date. So, even if Puijila (illustration above from Rybczynski et al., 2009) comes from younger deposits, its importance rests in that morphologically it is the most primitive known pinnipeds, providing evidence about the evolutionary steps that were taken in the transition from land to sea in this group of mammals.

Puijila the official website

Puijila in National Geographic

Also, here is the Spanish version of this post.

References

Deméré, T. A., A. Berta & P. J. Adams. 2003. Pinnipedomorph evolutionary biogeography. Bulletin of the American Museum of Natural History 279:32-76.

Domning, D. P. 2001. The earliest known fully quadrupedal sirenian. Nature 413:625-627.

Gingerich, P. D., M. ul Haq, I. S. Zalmout, I. H. Khan & M. S. Malkani. 2001. Origin of whales from early artiodactyls: hands and feet of Eocene Protocetidae from Pakistan. Science 293:2239-2242.

Graur, D. & D. G. Higgins. 1994. Molecular evidence for the inclusion of cetaceans within the order Artiodactyla. Molecular Biology and Evolution 11(3):357-364.

Luo, Z. & P. D. Gingerich. 1999. Terrestrial Mesonychia to aquatic Cetacea: transformation of the basicranium and evolution of hearing in whales. University of Michigan Papers on Paleontology 31:1-98.

Owen, R. 1855. On the fossil skull of a mammal (Prorastomus sirenoides, Owen), from the island of Jamaica. Quarterly Journal of the Geological Society of London 11:541-543.

Rybczynski, N., M. R. Dawson & R. H. Tedford. 2009. A semi-aquatic Arctic mammalian carnivore from the Miocene epoch and origin of Pinnipedia. Nature 458:1021-1024.

Savage, R. J. G., D. P. Domning & J. G. M. Thewissen. 1994. Fossil Sirenia of the West Atlantic and Caribbean region. V. Prorastomus sirenoides Owen, 1855. Journal of Vertebrate Paleontology 14(3):427-449.

Seiffert, E. R. 2007. A new estimate of afrotherian phylogeny based on simultaneous analysis of genomic, morphological, and fossil evidence. BMC Ecolutionary Biology 7:224 Open access

Shimamura, M., H. Yasue, K. Ohshima, H. Abe, H. Kato, T. Kishiro, M. Goto, I. Munechika & N. Okada. 1997. Molecular evidence from retroposons that whales form a clade within even-toed ungulates. Nature 388:666-670.

Tabuce, R., L. Marivaux, M. Adaci, M. Bensalah, J.-L. Hartenberger, M. Mahboubi, F. Mebrouk, P. Tafforeau & J.-J. Jaeger. 2007. Early Tertiary mammals from North Africa reinforce the molecular Afrotheria clade. Proceedings of the Royal Society B 274:1159-1166.

Thewissen, J. G. M., E. M. Williams, L. J. Roe & S. T. Hussain. 2001. Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls. Nature 413:277-281.

Thewissen, J. G. M., L. N. Cooper, M. T. Clementz, S. Bajpai & B. N. Tiwari. 2007. Whales originated from aquatic artiodactyls in the Eocene epoch of India. Nature 450:1190-1195.

Uhen, M. D. 2007. Evolution of marine mammals: back to the sea after 300 million years. Anatomical Record 290:514-522.