Saturday, September 3, 2011
Bad weather, ancient oceans and modern seagrasses… a brief visit to Puerto Rico
Wednesday, June 15, 2011
The hand of Steller’s sea cow, revisited
Ok, so now to the meaty part of this post. Back in 2008 I wrote about the hand of Steller’s sea cow (Hydrodamalis gigas) or more likely the lack of one. Back then I mentioned that G. W. Steller was one of the few people who saw H. gigas alive, and in his account he mentions the lack of fingers in this species. However, so far no hand bones have been found associated with Hydrodamalis gigas or even H. cuestae (see reconstruction below) (the presumed direct ancestor of the former), this lead to skepticism about his observations. So, how was that Steller’s account about the hand of H. gigas was corroborated?
The discovery of Dusisiren dewana in the Late Miocene (10-8 million years ago [mya]) of Japan, provided the key elements to support Steller’s account. Dusisiren is a genus of sea cow that include two other, earlier species: D. reinharti from the Early Miocene (20-16 mya) and D. jordani from the Middle-Late Miocene (12-10 mya) (see picture of D. jordani below). The species of Dusisiren together with Hydrodamalis cuestae (see picture above), H. spissa and H. gigas are members of a subfamily of dugongids (which means they are more closely related to dugongs than to manatees) known as the Hydrodamalinae (Domning, 1994; Domning & Furusawa, 1994). Hydrodamalines, as they are also known, ranged throughout coastal waters of the northern Pacific, from Baja California to Japan, with one species surviving until historical times (Domning & Furusawa, 1994). Some distinctive features of this group include, large body size, reduction or loss of dentition, and reduction of the digits, which brings us back to D. dewana.
Dusisiren jordani, anterior view (specimen at exhibit at the Natural History Museum of Los Angeles). Notice the "normal" hand in this species.
Described by Takahashi et al. in 1986, the remains of Dusisiren dewana were found in Late Miocene deposits in Japan and together with other elements of the skeleton there was also a nearly complete forelimb, including the hand. These were crucial in providing an insight into what the hand of later hydrodamalines was like. Dusisiren dewana shows reduction of the hand bones, in this respect differing from D. jordani, its predecessor, whose hands showed no reduction of elements (i.e. long metacarpals and long fingers composed of multiple elements) (see picture above). As for D. dewana, the carpals (wrist bones) are quite similar to those of other sirenians, however, the metacarpals (the bones between the wrist and fingers) and the phalanges (finger bones) are reduced, actually extremely reduced in the case of the phalanges (see picture below of Hydrodamalis cuestae). D. dewana as the sister taxon (closest relative/ancestor) of Hydrodamalis spp. is best evidence at hand (unintended pun) supporting Steller’s account on the hand of the sea cow that bears his name and allowing for reconstructions like the one seen here for H. cuestae.
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. & H. Furusawa. 1994. Summary of taxa and distribution of Sirenia in the North Pacific Ocean. Island Arc 3:506-512.
Takahashi, D., D. P. Domning & T. Saito. 1986. Dusisiren dewana, n. sp. (Mammalia: Sirenia), a new ancestor of Steller’s sea cow from the upper Miocene of Yamagata Prefecture, northeastern Japan. Transactions and Proceedings of the Paleontological Society of Japan, New Series 141:296-321.
Saturday, March 5, 2011
When you're strange, or how different are the teeth of extant sirenians
*T. manatus includes two subspecies: the West Indian manatee (T. manatus manatus) and the Florida manatee (T. manatus latirostris).
Although overall very similar to their extinct relatives, some characteristics of these extant sirenians are quite different and unique, not really representative of each particular group. One of those characteristics, and the one I will discuss here, is the dentition of Dugong dugon. This species, which lives in the Indo-Pacific region, is characterized by having molars that are best described as nearly enamel-less (a thin enamel cap is quickly worn off), open-rooted, ever-growing pegs (see pictures below), resembling more those of sloths, than those of manatees!!
To understand how they differ let me give you a little background info on sirenian dentition. Primitively, the dental formula of sirenians consisted of three incisors, one canine, five premolars* and three molars (Domning, 1994; Savage et al., 1994). Throughout sirenian evolution, this formula has been reduced, first by loss of the fifth permanent premolar in the earliest dugongids (see figure below), then by subsequent loss of second and third incisors, canine and the other permanent premolars. In terms of general features, the molars of most fossil sirenians can be described as bilophodont, with enameled crowns that are clearly differentiated from the root (these being closed and multiple [with three in upper molars and two in the lowers]). These kind of molars somewhat resemble those of the most primitive proboscideans (Sanders et al., 2010), which, by the way, are the closest relatives of sirenians.
*Presence of a fifth premolar is reminiscent of the primitive eutherian condition and, among Cenozoic placental mammals, a pretty unique condition only seen in sirenians (Rose, 2006).
Partial left maxilla of a late Eocene (37.2-33.9 Ma) dugongid, (A) lateral view (reversed); (B) occlusal view (anterior to the right). Notice that the deciduous premolar is molarized and multi-rooted, whereas the permanent premolars are more conical and single rooted. dP = deciduous premolar; M = molar; P premolar (permanent).
Beginning about the early Oligocene (33.9-28.4 Ma) the dental formula of most dugongids (including Dugong) stayed the same; it consisted of the first upper incisor, upper and lower deciduous premolars 3-5 and permanent molars 1-3 (see A-B in figure below)* (Domning & Pervesler, 2001). Up until the appearance of Dugong dugon the morphology of the molars was, in general, much the same as that described above. The peculiar, somewhat aberrant, dentition of dugong likely evolved quite recently (geologically speaking). Unfortunately there are no fossils of D. dugon to really know when. The best evidence comes from a very Dugong-like sirenian skull from the Pleistocene of Florida (Domning, 2001) (sadly, it is in a private collection). This skull is from sediments that were deposited about two million years ago and is nearly identical to Dugong dugon; one of the few differences, actually the most obvious one, is that its teeth are much like those of earlier dugongids, bilophodont, with enamel and multiple, closed roots. Therefore, if this fossil is taken as the closest ancestor of extant dugong, it would imply that its bizarre dentition is a quite recent (less than 2 million years) evolutionary novelty.
*Or even further reduced from that condition (as was the case of Steller’s sea cow and its closest relatives).
Occusal view of various sirenians (A-C). A. Left upper toothrow of a late Oligocene (28.4-23 Ma) dugongid from Puerto Rico. B. Right lower toothrow of Dugong dugon, notice that the lack of enamel, leaves a nearly featureless surface; the black rim is just stain on the portion of the tooth that was above the gum line. C. Left upper toothrow of Trichechus inunguis.
Manatees (Trichechus spp.) also have unusual dentition when compared to dugongids and some of their ancestors (for starters they lack incisors). If you look at the figure above (C) you’ll notice that there seems to be more molars than there should be. And that is correct; manatees have supernumerary molars that are constantly being replaced throughout their entire life (not to be confused with the delayed eruption that characterizes elephants). The molars in Trichechus are also proportionately small, that way more can fit in the toothrow at a given time (sometimes up to eight! [Domning & Hayek, 1984]). The earliest evidence for supernumerary molars in trichechids is seen in Ribodon limbatus, a fossil manatee from the late Miocene (11.6-5.3 Ma) or early Pliocene (5.3-3.6 Ma) of Argentina (Ameghino, 1883; Pascual, 1953; Domning, 1982). Ribodon does differ in that its molars were not reduced, so there were less per toothrow. Other fossil trichechids have the more “normal” type of molars described above.
Anyways there you have it; extant seacows have quite different dentition from their extinct relatives. Of course, there remains the questions of why evolve ever-growing molars, or ever-replacing supernumerary molars. But I’ll leave those for some other occasion.
Ameghino, F. 1883. Sobre una colección de mamíferos fósiles del piso mesopotámico de la formación patagónica, recogidos en las barrancas del Paraná por el professor Pedro Scalabrini. Boletín de la Academia Nacional de las Ciencias de Córdoba 5:101-116.
Domning, D. P. 1982. Evolution of manatees: a speculative history. Journal of Paleontology 56:599-619.
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. 2001. Sirenians, seagrasses, and Cenozoic ecological change in the Caribbean. Palaeogeography, Palaeoclimatology, Palaeoecology 166:27-50.
Domning, D. P. & L.-A. C. Hayek. 1984. Horizontal tooth replacement in the Amazonian manatee (Trichechus inunguis). Mammalia 48:107-127.
Domning, D. P. & P. Pervesler. 2001. The osteology and relationships of Metaxytherium krahuletzi Depéret, 1895 (Mammalia: Sirenia). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 553:1-89.
Pascual, R. 1953. Sobre nuevos restos de sirénidos del mesopotamiense. Revista de la Asociación Geológica de Argentina 8:163-181.
Rose, K. D. 2006. The Beginning of the Age of Mammals. John Hopkins University Press, Baltimore, Maryland, 428 pp.
Sanders, W. J., E. Gheerbrant, J. M. Harris, H. Saegusa & C. Delmer. 2010. Proboscidea; pp. 161-251 in L. Werdelin & W. J. Sanders (eds.), Cenozoic mammals of Africa. University of California Press, Berkeley, California.
Savage, R. J. G., D. P. Domning & J. G. M. Thewissen. 1994. Fossil Sirenia of the West Atlantic and Caribbean region. V. The most primitive known sirenian, Prorastomus sirenoides Owe, 1855. Journal of Vertebrate Paleontology 14:427-449.
Thursday, December 2, 2010
Dominican Republic, part II
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!
Tuesday, November 2, 2010
Dominican Republic, part I

To begin with, I had never been to the Dominican Republic before; it was actually my first time in another Caribbean island other than my own, Puerto Rico. I did not know what to expect, but I ended up with a very good impression of the island and its people! There were many things I liked, for example, there seems to be a lot of land dedicated to agriculture (which I think is awesome), the food was delicious and of course, the geology is very interesting!!
The focus of the trip to the DR was to collect rock samples and fossils from Early Cretaceous limestones as part of the thesis project of my friend and colleague Alvin Bonilla-Rodriguez from the Geology Department at the University of Kansas. Our collecting efforts were mainly focused in the central part of the country where Albian-age limestones occur. We stayed most of the time near the towns of Bonao in the Monseñor Nouel Province and in Hato Mayor del Rey in Hato Mayor Province. Other people that joined us during our time there were: Luis Gonzalez from the Department of Geology at the University of Kansas (and Alvin’s advisor) and Wilson Ramírez and Hernán Santos from the Department of Geology at the University of Puerto Rico-Mayagüez. The Servicio Geológico Nacional and its director Santiago Muñoz Tapia were also very helpful in terms of logistics and advise.
At first it was hard finding good outcrops, some were heavily overgrown and in other occasions our compact car was obviously ill-fitted for some of the roads we needed to go through. But, by about the middle of our first week we finally found some outcrops, not the best, but it was a start!
Our luck got better when we moved east towards Hato Mayor. There we found a very nice outcrop (with lots of fossils) of the Hatillo Limestone! The fossils were very important, specially the rudist, for determining the age of the rocks. And yes, we were in the Albian!
Well, this is it for this first post. Remember there are more to come, so stick around! On the next post we’ll move from the Albian, to the Miocene!
Sunday, July 18, 2010
Convergencia: el caso de Odobenocetops ó el delfín que parecía morsa
*Información adicional sobre desmostilios, incluyendo reconstrucciones, aquí, aquí y aquí.
En organismos vivos es relativamente fácil determinar si estructuras parecidas son convergentes o no, ya que se pueden hacer observaciones directas sobre el comportamiento o función de la estructuras o morfología que se consideran convergentes. Sin embargo, con los fósiles es distinto ya que generalmente no podemos hacer observaciones directas sobre el comportamiento de un organismo extinto. Es por esto que nos toca entonces interpretar la funcionalidad de esas estructuras comparando con lo que conocemos de organismos vivos. A continuación les traigo un interesante ejemplo de convergencia entre un fósil y un organismo moderno
Odobenocetops
En el 1993, Christian de Muizon describió un fósil de delfínido (el grupo de ballenas que incluye a los delfines) que representa uno de los mejores ejemplos de convergencia que conozco. El fósil consistía en parte de un cráneo (ver la imágen abajo) colectado en el sur de Perú. Los sedimentos donde se encontró el fósil son parte de una secuencia de rocas sedimentarias conocida como la Formación Pisco, de edad Pliocénico Temprano (5.3-3.6 millones de años). El fósil no solo representaba una nueva especie, sino también un nuevo grupo de delfínidos al cual llamó Odobenocetopsidae.
El primer espécimen descrito lo llamaron Odobenocetops peruvianus Muizon, 1993 (ver imagen abajo). Los rasgos morfológicos más distintivos de esta especie consisten en tener el rostro corto (el área frente a los ojos), procesos alveolares de la premaxilla (los espacios para los dientes) agrandados y orientados hacia abajo; los procesos alveolares era donde tenían unos colmillos también agrandados. La premaxilla también tiene varios forámenes y áreas de inserción de músculos bien marcadas que indican la presencia de labios grandes y quizás hasta bigotes. Otras características del cráneo incluye tener las órbitas de los ojos posicionadas dorsalmente y un paladar profundo. Estas características que tanto distinguen a los odobenocetópsidos son reminiscentes a las morsas* más que a otros delfinoideos.
*Vean las imágenes de un cráneo de morsa y comparen.
Odobenocetops peruvianus, arriba, foto del cráneo en vista lateral (anterior hacia la derecha); abajo, ilustración del cráneo mostrando los diferentes huesos que lo componen y el área de la órbita (cículo azul), modificada de Muizon (1993). Abreviaciones: Al = aliesfenoide; Bo = basioccipital; Fr = frontal; Mx = maxila; Na = nasal; Oc = occipital; Or = órbita; Pa = parietal; Pal = palatino; Pmx = premaxila; Ppo = proceso postorbital; Ppr = proceso preorbital; Pt = terigoide; Z = zigomático.
Otra especie adicional fue descrita unos años más tarde (Muizon et al., 1999). Odobenocetops leptodon también fue encontrado en la Formación Pisco, sin embargo, proviene de un horizonte alrededor de un millón de años más joven que donde se encontró O. peruvianus. Algunas de las características que distinguen a esta nueva especie es un rostro más grande y redondeado, mayor tamaño del paladar y la presencia de fosas (depresiones) para los sacos premaxilares. La ocurrencia de esta última estructura que menciono, se correlaciona con la presencia del órgano del melón (Mead, 1975), la cual es una estructura utilizada para ecolocalización. O. peruvianus aparentemente carecía de esta estructura ya que carece de fosas para los sacos premaxilares, sin embargo, la posición de las órbitas y la forma de los procesos supraorbitales le permitía tener visión binocular; en O. leptodon los procesos supraorbitales eran distintos y sus orbitas estaban posicionadas más hacia el lado, careciendo de visión binocular, pero al menos auxiliado por la presencia del órgano del melón y la capacidad de hacer ecolocalización. Así que al parecer, O. peruvianus dependía de la visión para localizar su alimento, mientras que O. leptodon probablemente dependía mayormente de ecolocalización (como es el caso de muchos otros delfinoideos).
Los odobenocetópsidos están relacionados al único otro grupo de cetáceos que tienen colmillos agrandados, los narvales. Estos también tienen un colmillo agrandado (y en ocasiones los dos), pero estos están orientados hacia adelante. En los odobenocetópsidos la función de estos colmillos parece haber sido social ya que presentan dimorfismo sexual; los machos poseen un colmillo (el de la derecha) más agrandado que el otro (algunos hasta midiendo más de un metro), mientras que en las hembras ambos eran más pequeños y similares en tamaño (Muizon et al., 1999; Muizon & Domning, 2002).
Reconstrucción de Odobenocetops ilustración de Mary Parrish (National Museum of Natural History, Washingon, DC), tomado de Muizon et al. (1999).
¿Que pudo llevar a un delfínido a parecer una morsa?
El parecido de los odobenocetópsidos a las morsas (convergencia) llevó a los paleontólogos que los han estudiado a inferir que su modo de alimentación era muy similar. O sea, que posiblemente, al igual que las morsas, los odobenocetópsidos se alimentaban de bivalvos bentónicos (que viven en el suelo marino), localizándolos con asistencia de los bigotes, aguantando el bivalvo con los labios y utilizando la lengua como un pistón, creando un vacío dentro de la cavidad bucal para lograr succionar el organismo fuera de su concha.
La distribución geográfica de las morsas, tanto vivas como las fósiles, está limitada al hemisferio norte (Deméré et al., 2003). Asumiendo que las especies extintas de morsas tenían las mismas preferencias alimenticias que la especie moderna (y al parecer así era) y al tener una distribución limitada, ese espacio ecológico, ocupado por las morsas en el hemisferio norte, estuvo “vacío” en el sur. Es así que posiblemente surgió este caso de convergencia, donde un delfinoideo termina pareciendo una morsa, ocupando ese espacio ecológico y utilizando los recursos disponibles.
Este es uno de los casos en paleontología donde el presente es la llave del pasado. Si no conocieramos las morsas habría sido muy difícil interpretar la funcionalidad morfológica de estos extraños delfinoideos.
Deméré, T. A., A. Berta & P. J. Adam. 2003. Pinnipedimorph evolutionary biogeography. Bulletin of the American Museum of Natural History 13 (279):32-76.
Mead, J. G. 1975. Anatomy of the external nasal pasajes and facial complex in the Delphinidae (Mammalia, Cetacea). Smithsonian Contributions to Zoology 207:1-72.
Muizon, C. de. 1993. Walrus-like feeding adaptation in a new cetacean from the Pliocene of Peru. Nature 365:745-748.
Muizon, C. de & D. P. Domning. 2002. The anatomy of Odobenocetops (Delphinoidea, Mammalia), the walrus-like dolphin from the Pliocene of Peru and its paleobiological implications. Zoological Journal of the Linnean Society 134:423-452.
Muizon, C. de, D. P. Domning & M. Parrish. 1999. Dimorphic tusas and adaptive strategies in a new species of walrus-like dolphin (Odobenocetopsidae) from the Pliocene of Peru. Comptes-rendus de l’Academie des Sciences, Paris, Sciences de la Terre et des Planetes 329:449-455.Sunday, July 11, 2010
Better late than never: GSA 2009
Anyways, if you missed my poster presentation at the 2009 annual meeting of the Geological Society of America, here is your chance to see it. Below is the abstract (you can also see it here) as well as the poster itself.
Evolution and diversification of Dugonginae (Sirenia: Dugongidae) in the West Atlantic and Caribbean region
Jorge Velez-Juarbe & Daryl P. Domning
Sirenian evolution presumably started in the southern Tethys realm of the Old World. However, the most primitive sirenians have been found in the Caribbean region in rocks that date back to the Early or Middle Eocene, evidence that they took on an aquatic lifestyle early in their history and were able to disperse around the Atlantic. Since then, the West Atlantic and Caribbean (WAC) region has been important for the evolution of this group of mammals, with all four of the known families (Prorastomidae, Protosirenidae, Dugongidae, Trichechidae) occurring in the region. From the Oligocene through Pliocene, the predominant groups in the WAC region were members of the dugongid subfamilies Dugonginae and Halitheriinae, with the former being the more speciose of the two. In some parts of the WAC, dugongines and halitheriines are found in the same deposits, indicating that some form of niche partitioning was occurring among three or even more sympatric species of sirenians, something that is not observed among the living species.
Niche partitioning among sympatric sirenian species is most obviously explained by differences in: (1) tusk morphology; (2) rostral deflection; and (3) body size. The most primitive members of the Dugonginae are found in the WAC region. Their distinctive cranial morphology seems to have evolved as a specialization for harvesting larger seagrass rhizomes that were out of reach of the less specialized halitheriines. New fossils from Puerto Rico, Florida, and Yucatán help to reinforce our ideas about niche partitioning among sympatric sirenian species as well as establish the WAC region as the center of known dugongine diversity.
This was my first poster presentation about one of the topics that I’m working on as part of my thesis. The poster was a success! I was awarded runner-up of the 2009 Paleontological Society Poster Award. Congratulations to the first place and to the other runner-up too! It was a great meeting with very interesting talks as well as getting to spend some time with good friends!!
Enjoy the poster and feel free to ask question or make comments!