Origen e Historia Evolutiva de los Manatíes, Parte 2

Hoy les traigo el descubrimiento más reciente sobre la historia evolutiva de los manatíes. Y es que en esta nueva publicación científica junto con los colegas Catalina Suarez, Javier N. Gelfo y Jorge W. Moreno-Bernal, describimos los restos más antiguos de manatíes de las Américas. En la entrada previa de esta serie ya les había adelantado lo que sabíamos hasta el momento (2015) sobre la historia evolutiva de los manatíes, incluyendo el origen europeo del grupo. En este nuevo trabajo describimos una porción de un maxilar izquierdo con las dos muelas más posteriores (denominadas como M2 y M3), proveniente de la Formación Barzalosa, la cual se depositó entre 16.5-17.7 millones de años, lo cual nos ayuda a entender mejor los orígenes de las especies actuales de manatí (Suarez et al., 2021). Al estudiar y examinar el fósil de la Formación Barzalosa llegamos a la conclusión que guarda mucha similitud con Potamosiren magdalenensis, una especie de manatí extinto que vivía en lo que hoy día es Colombia entre 11.6-13.8 millones de años atrás (Reinhart, 1951; Kellogg, 1966; Domning, 1997). Por su parecido, designamos a este nuevo fósil como Potamosiren cf. P. magdalenensis, y pasa a ser el registro más antiguo de esta especie. Algo que distingue a Potamosiren de otros sirenios, especialmente de otros manatíes, son dos características particulares de sus dientes. La primera es que todavía retienen tres molares por cada cuadrante y la segunda, es que el esmalte de las muelas es inusualmente grueso.

A-C. Maxilar izquierdo de Potamosiren cf. P. magdalenensis de la Formación Barzalosa. 
D. Maxilar izquierdo de Potamosiren magdalenensis del Grupo Honda. (Modificado de Suarez et al., 2021).

El Potamosiren de la Fm. Barzalosa, junto a una especie poco conocida de manatí del Mioceno temprano de Perú (Antoine et al., 2016), representan los restos más antiguos de manatíes en las Américas. Ambos, coinciden geográficamente y temporalmente con el inicio de la formación del Mega-Humedal de Pebas, el cual abarcó una enorme zona entre 23-10 millones de años, e incluía partes de Perú, Colombia y Brazil (Hoorn et al., 2010). Este mega-humedal estaba conectado con el mar Caribe hacia el norte, lo cual funcionó como vía de entrada para distintos organismos marinos. Los depósitos sedimentarios que se formaron durante la duración de el Mega-Humedal de Pebas están principalmente compuestos de estratos que representan ambientes terrestres y de agua dulce, con algunas incursiones marinas. Mientras que el manatí de Pebas viene de sedimentos que representan ambientes acuáticos mixtos, los del Potamosiren de Barzalosa son de agua dulce, ambos entonces representan los primeros registros de sirenios del grupo terminal (crown group) fuera de ecosistemas marinos. (En este contexto, Sirenia es el crown group, que es el grupo más exclusivo compuesto de las especies actuales y sus parientes más cercanos, y Pan-Sirenia es el grupo total que incluye las especies extintas basales y las actuales; ver imagen abajo).

En este árbol filogenético se puede apreciar las relaciones entre los distintos grupos de sirenios, con énfasis en los manatíes (Trichechidae). En gris está representado la duración del Mega-Humedal de Pebas y los distintos colores representan los ambientes donde se encuentran las distintas especies. (Modificado de Suarez et al., 2021).

Como se puede observar en el árbol (ver arriba) entre los sirenios, solamente los manatíes (Trichechinae) incursionan en ambientes de agua dulce. Esto contrasta con otros grupos de organismos marinos. Por ejemplo, los odontocetos han incursionado en ambientes de agua dulce en múltiples ocasiones y en diversas partes del mundo (Fordyce, 1983; Cassens et al., 2000; Hulbert & Whitmore, 2006; Geisler et al., 2011; Boessenecker & Poust, 2015; Bianucci et al., 2013; Pyenson et al., 2015; Boersma & Pyenson, 2016; Benites-Palomino et al., 2020). El Mega-Humedal de Pebas no fue la excepción ya que se han encontrado fósiles de iniidos y platanistidos en distintas localidades (Bianucci et al., 2013; Benites-Palomino et al., 2020).

América del Sur y Central alrededor de 12 millones de años atrás, mostrando las localidades donde se han encontrado iniidos y platanistidos en ambientes marinos (azul) y de agua dulce (amarillo). (Modificado de Benites-Palomino et al., 2020).

En adición a los mamíferos marinos, el Mega-Humedal de Pebas fue un lugar excepcional para que distintos grupos de organismos de origen marino incursionaran y se adaptaran a ambientes de agua dulce y pantanosos, como por ejemplo las mantarrayas de agua dulce (Fontenelle et al., 2021) y algunos otros grupos de peces (Bloom & Lovejoy, 2017). Incluso algunos grupos se diversificaron para ocupar distintos espacios ecológicos y proliferaron durante la duración de este mega-humedal (Salas-Gismondi et al., 2015). No es la primera vez que se propone que la formación del Mega-Humedal de Pebas haya sido clave para la incursión de manatíes en cuerpos de agua dulce. Esta hipótesis se propuso por primera vez en los 1980's (Domning, 1982) y más recientemente en un estudio genético de las especies actuales (Silva de Zouza et al., 2021). El Potamosiren de Barzalosa es evidencia fósil que le da aún más veracidad a esta hipótesis (Suarez et al., 2021).

Las muelas de Potamosiren
Diferentes grupos de mamíferos herbívoros han evolucionado distintas adaptaciones como resultado de tener una dieta que incluye vegetación abrasiva o fibrosa. Por ejemplo, los caballos tienen dientes hipsodontes (dientes con coronas muy altas) que los hacen más resistentes o duraderos al desgaste. Otra posible alternativa para contrarrestar el desgaste es que el esmalte del diente sea más grueso de lo usual, haciéndolo más resistente. Mientras que una tercera estrategia es tener dientes que pueden ser reemplazados a medida que se van gastando.
Como mencione previamente, el esmalte inusualmente grueso de las muelas de Potamosiren es único entre los sirenios. Sin embargo, no es la primera vez que se observa este fenómeno en mamíferos marinos herbívoros. Y es que el esmalte engrosado nos recuerda a las muelas de Desmostylus, un mamífero marino herbívoro que habitaba a lo largo de las costas del Pacífico Norte hasta hace unos 10-12 millones de años. Desmostylus pertenece a un grupo completamente extinto llamado Desmostylia, y sus cráneos y esqueletos son tan particulares que todavía queda mucho por descifrar sobre su paleoecología e incluso sobre su postura en tierra y su relación con otras especies. Sin embargo, gracias a estudios isotópicos usando los dientes de Desmostylus conocemos un poco sobre su dieta (Clementz et al., 2003). Los resultados de ese estudio sugieren que los Desmostylus consumían distintos tipos de vegetación acuática, algunas, como los pastos marinos, cuales inevitablemente debía incluir interacción con el substrato e ingestión accidental de sedimento; en fin, una dieta relativamente variada y muy parecida a la del manatí de Florida (Trichechus manatus latirostris), los cuales tienen una dentadura muy particular como veremos más adelante. Como nota al calce, hay que mencionar que la dieta de los manatíes actuales es una excepción, ya que los sirenios han sido casi exclusivamente consumidores de pastos marinos durante la mayor parte de su historia evolutiva (Clementz, et al., 2003; MacFadden et al., 2004; Clementz & Sewall, 2011), y su morfología dental ha sido generalmente muy conservadora. Así que cuando observamos cualquier cambio significativo en la dentadura de una especie de sirenio inmediatamente despierta la curiosidad por conocer mejor sus hábitos alimenticios.

Parte de la mandíbula de un Desmostylus, incluyendo el cuarto premolar y las primeras dos muelas. Las muelas están compuestas de cúspides modificadas en columnas con esmalte muy grueso.

Los distintos ecosistemas que formaban parte del Mega-Humedal de Pebas permitieron que los manatíes tuviesen acceso a una cantidad mayor de vegetación, en adición a un incremento en la interacción con vegetación acuática cerca del substrato e ingestión accidental de sedimento. Esto claramente debió resultar en un cambio significativo en la dieta de estos sirenios (Domning, 1982; Beatty et al., 2012), que como antes mencionado, eran principalmente consumidores de pastos marinos. Tomando esto en consideración, en nuestro trabajo proponemos que el esmalte engrosado de las muelas de Potamosiren es una adaptación a una dieta más variada, incluyendo mayor cantidad de vegetación abrasiva y fibrosa. Y como verán, las especies de manatí que vinieron después de Potamosiren se adaptaron a esta dieta de una forma aún más particular.

Molares superiores (izquierda) y mandíbula (derecha) de Potamosiren magdalenensis. Aquí se puede apreciar la presencia de solamente tres molares por cuadrante.

Posterior a la extinción de Potamosiren aparece otra especie de manatí conocido como Ribodon limbatus el cual también se encuentra en depósitos de agua dulce del Mioceno tardío al Plioceno. Ribodon se diferencia de Potamosiren al tener más de tres molares por cuadrante (usualmente entre 4-5), los cuales son muy similares entre sí y son relativamente pequeños, tampoco tienen esmalte engrosado. Ribodon también se caracteriza porque sus muelas eran reemplazadas gradualmente. En otras palabras, a lo largo de la vida de un individuo las muelas se movían lentamente hacia al frente mientras que en la parte posterior iban saliendo muelas nuevas, una y otra vez! Este tipo de reemplazo horizontal de los dientes es una adaptación que se observa por primera vez en Ribodon y que todavía observamos en las tres especies de manatí actuales.

Detalles de la dentición de Ribodon (arriba) y Trichechus (abajo). Noten la presencia de multiples muelas, de similar morfología y tamaño, al igual que los dientes nuevos hacia la parte posterior de la fila dental.

Aún con esta nueva información, queda mucho por conocer sobre la historia evolutiva de los manatíes. Por ejemplo, todavía no conocemos bien la identidad del manatí de Pebas y tampoco conocemos bien la morfología craneal de Potamosiren y Ribodon. Similarmente, faltan estudios isotópicos que nos puedan dar una mejor idea de la dieta de estas especies de manatíes ancestrales. Esperemos que tengamos contestaciones a estas incógnitas en un futuro no tan lejano.

Los manatíes actuales pertenecen al género Trichechus, e incluyen tres especies - T. manatus (manatí antillano), T. inunguis (manatí amazónico) y T. senegalensis (manatí africano). En adición a estas tres especies, se conoce una subespecie extinta del manatí antillano del Pleistoceno de Norteamérica (T. manatus bakerorum) y una especie del Pleistoceno amazónico (T. hesperamazonicus) (Domning, 2005; Perini et al., 2020). Todas las especies de Trichechus (actuales y extintas) reemplazan sus muelas horizontalmente, y estas son aún más pequeñas que las de Ribodon, teniendo en algunos casos hasta siete muelas por cuadrante! Esto resulta en un sistema relativamente eficiente donde se reemplazan los dientes continuamente para contrarrestar una dieta variada que puede incluir vegetación fibrosa e ingestión accidental de sedimento. Sin embargo, no es perfecto, y la composición y tipo de sedimento también juega un papel importante. En lugares como el estado de Florida donde la arena tienen una mayor composición de cuarzo, el desgaste en los dientes de los manatíes es mayor que el observado en los que viven en el Caribe, donde las arenas están principalmente compuestas de carbonato de calcio (Domning & Hayek, 1984).

Literatura

Antoine, P.-O., M. A. Abello, S. Adnet, A. J. Altamirano Sierra, P. Baby, G. Billet, M. Boivin, Y. Calderón, A. Candela, J. Chabain, F. Corfu, D. A. Croft, M. Ganerod, C. Jaramillo, S. Klaus, L. Marivaux, R. E. Bavarrete, M. J. Orliac, F. Parra, M. E. Pérez, F. Pujos, J. C. Rage, A. Ravel, C. Robinet, M. Roddaz, J. V. Tejada-Lara, J. Velez-Juarbe, F. P. Wesselingh, and R. Salas-Gismondi. 2016. A 60-million-year Cenozoic history of western Amazonian ecosystems in Contamana, eastern Peru. Gondwana Research 31:30–59.

Beatty, B. L., T. Vitkovski, O. Lamnert, and T. E. Macrini. 2012. Osteological associations with unique tooth development in manatees (Trichechidae, Sirenia): a detailed look at modern Trichechus and a review of the fossil record. Anatomical Record 295:1504–1512.

Benites-Palomino, A., G. Aguirre-Fernandez, J. W. Moreno-Bernal, A. Vanegas, and C. Jaramillo. 2020. Miocene freshwater dolphins from La Venta, Huila, Colombia suggest independent invasions of riverine environments in tropical South America. Journal of Vertebrate Paleontology 40:e1812078.

Bianucci, G., O. Lambert, R. Salas-Gismondi, J. Tejada, F. Pujos, M. Urbina, and P.-O. Antoine. 2013. A Miocene relative of the Ganges river dolphin (Odontoceti, Platanistidae) from the Amazonian basin. Journal of Vertebrate Paleontology 33:741–745.

Bloom, D. D., and N. R. Lovejoy. 2017. On the origins of marine-derived freshwater fishes in South America. Journal of Biogeography 44:1927–1938.

Boersma, A. T., and N. D. Pyenson. 2016. Arktocara yakataga, a new fossil odontocete (Mammalia, Cetacea) from the Oligocene of Alaska and the antiquity of Platanistoidea. PeerJ 4:e2321.

Boessenecker, R. W., and A. W. Poust. 2015. Freshwater occurrence of the extinct dolphin Parapontoporia (Cetacea: Lipotidae) from the upper Pliocene nonmarine Tulare Formation of California. Palaeontology 58:489–496.

Cassens, I., S. Vicario, V. G. Waddell, H. Balchowsky, D. Van Belle, W. Ding, C. Fan, R. S. Lal Mohan, P. C. Simoes-Lopes, R. Bastida, A. Meyer, M. J. Stanhope, and M. C. Milinkovitch. 2000. Independent adaptation to riverine habitats that allowed survival of ancient cetacean lineages. Proceeding of the National Academy of Sciences U.S.A. 97:11343–11347.

Clementz, M. T., and J. O. Sewall. 2011. Latitudinal gradients in greenhouse seawater d18O: evidence from Eocene sirenian tooth enamel. Science 332:455–458.

Clementz, M. T., K. A. Hoppe, and P. L. Koch. 2003. A paleoecological paradox: the habitat and dietary preferences of the extinct tethythere Desmostylus inferred from stable isotope analysis. Paleobiology 29:506–519.

Domning, D. P. 1982. Evolution of manatees: a speculative history. Journal of Paleontology 56:599–619.

Domning, D. P. 1997. Sirenia. In: R. F. Kay, R. H. Madden, R. L. Cifelli, and J. J. Flynn (Eds.), Vertebrate Paleontology in the Neotropics: the Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press, Washington and London, pp. 383–391.

Domning, D. P., and L.-A. C. Hayek. 1984. Horizontal tooth replacement in the Amazonian manatee (Trichechus inunguis). Mammalia 48:105–127.

Fontenelle, J. P., F. P. Luna Marques, M. A. Kolmann, and N. R. Lovejoy. 2021. Biogeography of the neotropical freshwater stingrays (Myliiobatiformes: Potamotrygoninae) reveals effects of continent-scale paleogeographic change and drainage evolution. Journal of Biogeography 00:1–14.

Fordyce, R. E. 1983. Rhabdosteid dolphins (Mammalia: Cetacea) from the middle Miocene, lake Frome area, South Australia. Alcheringa 7:27–40.

Geisler, J. H., M. R. McGowen, G. Yang, and J. Gatesy. 2011. A supermatrix analysis of genomic, morphological, and paleontological data from crown Cetacea. BMC Evolutionary Biology 11:1–33.

Hoorn, C., F. P. Wesselingh, H. T. Steege, M. A. Bermudez, A. Mora, J. Sevink, I. Sanmartín, A. Sanchez-Meseguer, C. L. Anderson, J. P. Figuerido, C. Jaramillo, D. Riff, F. R. Negri, H. Hooghiemstra, J. Lundberg, T. Stadler, T. Särkinen, and A. Antonelli. 2010. Amazonia through time: Andean uplift, climate change, landscape evolution, and biodiversity. Science 330:927–931.

Hulbert, R. C., and F. C. Whitmore. 2006. Late Miocene mammals from the Mauvilla local fauna, Alabama. Bulletin of the Florida Museum of Natural History. 46:1–28.

Kellogg, R. 1966. Fossil marine mammals from the Miocene Calvert Formation of Maryland and vVirginia, parts 3 and 4. Bulletin of the U. S. National Museum 247:65–101.

MacFadden, B. J., P. Higgins, M. T. Clementz, and D. S. Jones. 2004. Diets, habitat preferences, and niche differentiation of Cenozoic sirenians from Florida: evidence from stable isotopes. Paleobiology 30:297–324.

Pyenson, N. D, J. Velez-Juarbe, C. S. Gutstein, H. Little, D. Vigil, and A. O'Dea. 2015. Isthminia panamensis, a new fossil inioid (Mammalia, Cetacea) from the Chagres Formation of Panama and the evolution of 'river dolphins' in the Americas. PeerJ 2015: e1227. 

Reinhart, R. H. 1951. A new genus of sea cow from the Miocene of Colombia. Bulletin of the Department of Geological Sciences 28:203–214.

Salas-Gismondi, R., J. J. Flynn, P. Baby, J. V. Tejada-Lara, F. P. Wesselingh, and P.-O. Antoine. 2015. A Miocene hyperdiverse crocodylian community reveals peculiar trophic dynamics in proto-Amazonian mega-wetlands. Proceedings of the Royal Society B 282:20142490.

Silva de Souza, E. M., L. Freitas, E. K. da Silva Ramos, G. Selleghin-Veiga, M. C. Rachid-Ribeiro, F. A. Silva, M. Marmontel, F. Rodrigues dos Santos, A. Laudisoit, E. Verheyen, D. P. Domning, and M. Freitas Nery. 2021. The evolutionary history of manatees told by their mitogenomes. Scientific Reports 11:3564.

Suarez, C., J. N. Gelfo, J. W. Moreno-Bernal and J. Velez-Juarbe. 2021. An early Miocene manatee from Colombia and the initial Sirenian invasion of freshwater ecosystems. Journal of South American Earth Sciences 109:103277

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.

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.

The Southernmost Atlantic Seacows

Its been a while since I posted news on fossil sirenians. I've been very busy with fieldwork, manuscripts, among other things. The Spring interns have now gone back home. So, while I wait for the arrival of the next round of interns, here's the latest on fossil sirenians.


Where are sirenians found

With the exception of the now extinct Steller's seacow (Hydrodamalis gigas), all extant sirenians have tropical to subtropical distribution, with some species having a notably broad latitudinal and longitudinal distribution (Marsh et al., 2011). But, when we look at the fossil record of sirenians, we see a slightly different pattern of distribution, mostly tied to tectonic and/or climatic events. For example, during parts of the Cenozoic global temperatures were higher than today (Zachos et al., 2001), so you find fossils of sirenians far off their modern range (e.g. Belgium). These climatic variations amongst other physical drivers have played a prominent role in the distribution of seagrasses and seacows (expect more on this in the nearby future).

Nowadays, in the Western Atlantic and Caribbean (WAC) region, the most common and widespread sirenian is the West Indian Manatee (Trichechus manatus) whose range extends from as far north as the Carolina's (with some individuals reaching New England) to northeastern Brazil; another species found in the region is the Amazonian manatee (Trichechus inunguis) which lives in the Amazon basin (see map below). But, it hasn't always been like this. Throughout most of the Cenozoic, dugongids, a group of sirenians are now restricted to the Indo-Pacific region, were the predominant seacow group in the WAC, including multispecies communities in the region (Domning, 2001; Velez-Juarbe et al., 2012a; see previous post on this subject). Fossil of dugongids in the WAC are found in deposits as far north as Maryland, and as far south as Argentina. However, these southernmost dugongids, are poorly known, and have had a somewhat rocky taxonomic history.


From Metaxytherium to Dioplotherium a case of mistaken identity

The most common, and temporally and geographically widespread seacow genus known is the Halitheriine dugongid Metaxytherium. Species of this genus are known from late Oligocene through Pliocene deposits, and are found from the Eastern Pacific, Caribbean, Western and Northern Atlantic, and Western Tethys regions (e.g. Domning, 1988; Sorbi et al., 2012). Therefore it shouldn't have been much of a surprise when Roy H. Reinhart (1976) described a molar from the late Miocene Paraná Formation of Entre Ríos, Argentina as that of Metaxytherium. The importance of this find, lies in that prior to its discovery, the youngest species of Metaxytherium known from the WAC was the middle Miocene M. floridanum, which is not known outside of Florida (Domning, 1988). The Paraná molar was then, the youngest and southernmost record of the genus from the Western Atlantic.

However, species of Metaxytherium display a generally conservative morphology, and because of this, it has had a long, somewhat convoluted, taxonomic history. This is, fortunately, slowly being resolved as most species of Metaxytherium have been re-described (e.g. Domning, 1988; Domning & Pervesler, 2001; Sorbi et al., 2012) and studied in detail within the last 25 years, giving us, paleosirenologist a better idea of the valid species within the genus and variation within each species. Since Reinhart's description, several workers (Cozzuol, 1996; Cione et al., 2000; Domning, 2001) have disagreed with his interpretation regarding the affinities of the Argentinian molar. All of them referring the Paraná molar to Dioplotherium, still a dugongid, but one that belongs to the Dugonginae, a group very different from that to which Metaxytherium belongs. And indeed, the overall morphology of the tooth conforms well with what we know about Dioplotherium, it is in fact, very similar to those of Dioplotherium cf. D. allisoni from the early Miocene of Brazil (Toledo & Domning, 1991). This meant that Metaxytherium may have gone extinct in the WAC at the end of the middle Miocene (Domning, 1988), and that the genus only reached as far south as northeastern Brazil (Toledo & Domning, 1991), or did it?

Left: Map showing the distribution of Miocene seacows throughout the Americas. (ER = Entre Ríos).
Right: Map showing the distribution of extant sirenian in the Americas.
(Click on the map to view larger version.)

New fossils from the Paraná Formation

A couple of years ago I received an email from an Argentinian colleague, Jorge Noriega from CONICET in Diamante, informing me of a new discovery from the Paraná Formation in Entre Ríos. The new fossils consisted of left and right partial maxillae and most of the molars of a single individual (see figure below). At this point I was close to finishing my PhD, which meant that I had look at a lot of specimens and was well acquainted with the morphology of most, if not all Oligocene through Pliocene sirenians. Once I looked at the pictures of the new material, I quickly recognize these as most likely representing a species of Metaxytherium.

Molars of Metaxytherium from the late Miocene Paraná Formation. 1-2) left maxilla and M1-3 in occlusal view. 3-4) right maxilla and M3 (modified from Velez-Juarbe et al., 2012b)

Now, I must admit that dugongid teeth are not the most diagnostic, so figuring out if these actually belonged to Metaxytherium was not an easy and quick task. After a considerable amount of reading, and detailed observations of material from various species of Metaxytherium as well as other dugongids I was confident they belonged to that genus. And so, working together with Jorge and Brenda Ferrero (also from CONICET in Diamante) we took on the task of formally re-designating the fossil described by Reinhart (1976) as well as describing the new material which actually represented a species of Metaxytherium (Velez-Juarbe et al., 2012b). The new Parana molars are quite similar to those of the middle Miocene Metaxytherium floridanum, but, their dimensions are below the range exhibited by M. floridanum and may represents a different species. One of the positive outcomes resulting from this work, was realizing that teeth of dugongids can sometimes be of taxonomic usefulness. We noticed, that the molars of some of the more derived species of Metaxytherium often have additional cusp and/or cuspules, a derived character which is not observed in Dioplotherium or any of its kin (i.e. Dugongines). The contemporaneous presence of both, Dioplotherium and Metaxytherium is not something unheard of. This same duet, occurs in the late Oligocene of Florida, early Miocene of Brazil and possibly in the middle Miocene of California and Baja California (Domning, 2001; Velez-Juarbe et al., 2012a). This again shows that multispecies communities and niche partitioning seems to have been the norm, not the exception, throughout sirenian history.


References

Cione, A. L., M. M. Azpelicueta, M. Bond, A. Carlini, J. Casciotta, M. A. Cozzuol, M. de la Fuente, Z. Gasparini, F. Goin, J. Noriega, G. Scilato-Yané, L. Soibelzon, E. Tonni, D. Verzi, and M. G. Vucetich. 2000. Miocene vertebrates from Entre Ríos Province, Argentina. INSUGEO, Serie Correlación Geológica 14:191-238.

Cozzuol, M. A. 1996. The record of the aquatic mammals in southern South America. Münchner Geowissenschaftliche Abhandlungen A30:321-342.

Domning, D. P. 1988. Fossil Sirenia of the West Atlantic and Caribbean region. I. Metaxytherium floridanum Hay, 1922. Journal of Vertebrate Paleontology 8:295-426.

Domning, D. P. 2001. Sirenians, seagrasses, and Cenozoic ecological change in the Caribbean. Palaeogeography, Palaeoclimatology, Palaeoecology 1:27-50.

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

Marsh, H. D., T. J. O'Shea, and J. E. REynolds, III. 2011. Ecology and conservation of the Sirenia: dugongs and manatees. Cambridge University Press, 521p.

Reinhart, R. H. 1976. Fossil sirenians and desmostylids from Florida and elsewhere. Bulletin of the Florida State Museum, Biological Sciences 20:187-300.

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

Toledo, P. M., and D. P. Domning. 1991. Fossil Sirenia (Mammalia: Dugongidae) from the Pirabas Formation (Early Miocene), northern Brazil. Boletim do Museu Paraense Emílio Goeldi, Série Ciencias da Terra 1:119-146.

Velez-Juarbe, J., D. P. Domning, and N. D. Pyenson. 2012a. Iterative evolution of sympatric seacow (Dugongidae, Sirenia) assemblages during the past ~26 million years. PLoS ONE 7(2):e31294.

Velez-Juarbe, J., J. I. Noriega, and B. S. Ferrero. 2012b. Fossil Dugongidae (Mammalia, Sirenia) from the Paraná Formation (late Miocene) of Entre Ríos Province, Argentina. Ameghiniana 49:585-593.

Zachos, J., M. Pagani, L. Sloan, E. Thomas, and K. Billups. 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292:686-693.