New paper on fossil plant from the Neotropics

Yes, fossil plants! This is a first in this blog, which is otherwise, heavily biased towards marine tetrapods. However, that doesn't mean that when I do fieldwork I only focus on collecting fossil vertebrates. this of course has resulted in a number of publication on fossil invertebrates (e.g. Schweitzer et al., 2006), and now plants. The paper which was just published in the journal International Journal of Plant Sciences is a collaborative work led by former Florida Museum of Natural History colleague Fabiany Herrera, one of the few experts on fossil plants from the Neotropics, and his former advisors Steven R. Manchester and Carlos Jaramillo. The paper is a follows up on a previous paper published about four years ago in the same journal (Herrera et al., 2010), and that seeks to better understand the evolutionary and paleobiogeographic history of a group of plants called Humiriaceae that are found in the Neotropics, and western Africa. This group is mainly composed of large trees, most greater than 20 meters tall, and fruits have woody parts with very particular morphology, which results in a relatively high preservation and identification potential.

Map showing the distribution of fossil Humiriaceae endocarps (symbols) and extant genera (dashed lines) (modified from Herrera et al., 2010:fig. 1).

In the paper we describe fossilized endocarps (the inside part of the fruit) from the early Oligocene of Peru and Puerto Rico, and the late Miocene of Panama, and fossilized wood from the late Eocene of Panama (Herrera et al., 2014). The fossilized fruit from the Oligocene of Peru, which we dubbed Duckesia berryi, represents a new species of a tree that is nowadays only found in Amazonia, and is the oldest record of that genus. This not only shows that this particular taxon has an older history than previously thought, but it also shows that its former distribution was much more widespread. In addition to that, the fossil wood we describe, called Humiriaceoxylon ocuensis, shows that by the late Eocene parts of what is now Panama, was forested by large trees belonging to this particular group of plants. In addition, Fabiany had previously described a fossil Humiriaceae endocarp which he names Lacunofructus cuatrecasana from a locality near where the wood was found, and it may actually be that they represent the same tree (Herrera et al., 2012, 2014)*. This is a really cool find, as the region and where the fossils were collected, was not connected by land to neither North or South America, showing again, that overwater dispersal is not as much a problem for plants.
*Paleobotanists use different scientific names for the different parts of a plant as they are usually found separate, hence the endocarp has a name, and the wood another, even though they may be the same plant.

The fossil endocarp Duckesia berry (A-L) from the Oligocene of Peru, compared with the endocarp of the modern species D. verrucosa (M-O). (Modified from Herrara et al., 2014:fig. 1.)

The fossil from Puerto Rico consists of an endocarp of Sacoglottis tertiaria, otherwise known from the Neogene of Peru, Ecuador, Colombia and Panama (Herrera et al., 2010). Several species of the genus Sacoglottis are still found today, in the Amazonian region and west Africa. The fossil from Puerto Rico is from the early Oligocene San Sebastian Formation, one of my favorite formations where I've spent many hours searching for fossils. Actually, the locality where I found the endocarp is not far from where Aktiogavialis puertoricensis, Priscosiren atlantica, and a Caviomorph rodent tooth were collected (Velez-Juarbe et al., 2007; Velez-Juarbe and Domning, 2014; Velez-Juarbe et al., 2014).

The fossil endocarp Sacoglottis tertiaria from the early Oligocene San Sebastian Formation of Puerto Rico.

Fossil plants were previously described from the San Sebastian Fm. by previous workers, mainly, Arthur Hollick (1928) and Alan Graham and David Jarzen (1969). But none of the material they described indicated the presence of Humiriaceae in the island. In fact, they list many plant groups present in San Sebastian Fm. which are now absent from the flora of the island, now the Humiriaceae can be added to that list. As I recently said in a newspaper interview, Puerto Rico during the Oligocene was very different from nowadays, and there is still more to be discovered!

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

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

References

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

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

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

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

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

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

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

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

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

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.

Domningia and other Indian sirenians

Over the last 12 years a number of fossil sirenians have been described from Eocene through Miocene deposits from India. This has not stopped as, new taxa are been discovered and described; much of the effort is spearheaded by Sunil Bajpai of the Dept. of Earth Sciences, Indian Institute of Technology and J. G. M. Thewissen of the Dept. of Anatomy and Neurobiology, NEOUCOM (Thewissen Lab). So far all the fossil sirenians from India have been collected from outcrops in the Kachchh District near the western coast of the country; this area is better known for primitive cetaceans such as remingtonocetids (Kumar & Sahni, 1986).

Eocene

The oldest Indian sirenians come from the Middle Eocene Harudi Formation; a total of three species representing two families: Protosiren sp. (Protosirenidae), Eotheroides babiae, Eosiren sp. (Dugongidae, Halitheriinae) (Bajpai et al. 2006). Other species belonging to those genera are found in Eocene deposits in northern Africa (Domning, 1996) indicating a southern Tethyan influence, better referred to as the Indo-African Region of the Tethys Realm (Harzhauser et al. 2002). There should be more on Eocene sirenians from India, sometime this year.

Oligocene

So far only one Oligocene sirenian is known from this region, Bharatisiren indica (Dugongidae, Dugonginae) from the Maniyara Fort Formation of Late Oligocene age (Bajpai et al. 2006). This is the oldest dugongine found in this region, and it is as old as the dugongines, Crenatosiren olseni and Dioplotherium manigualti from the Western Atlantic (Domning, 1989; Domning, 1997) and a new taxon from Puerto Rico. Dugongines are a group that is thought to have evolved in the Western Atlantic and Caribbean Region (Domning, 2001), the occurrence of B. indica seems to point to a latest Eocene or Early Oligocene origin for the group with subsequent trans-Atlantic dispersal to the Indo-African Region. An alternative scenario, is the origin of dugongines in the Mediterranean region of the Tethys Realm with subsequent east and west dispersal, unfortunately no Early or Late Oligocene dugongines are known from this region (the names of the paleobiogeographic areas based on Harzhauser et al. 2002).

Lateral view of the skull of B. indica from Bajpai et al. (2006).

Miocene

During the Early Miocene there seems to have been a radiation of dugongines in this region. With at least two taxa known Bharatisiren kachchhense, Domningia sodhae* from Khari Nadi Formation (Bajpai & Domning, 1997; Thewissen & Bajpai, 2009), and another currently under study, it is an good example of multispecies communities like the ones present in the Western Atlantic and Caribbean Region (Domning, 2001). As you can see in the composite picture below, a notable difference between B. kachchhense and Domningia sodhae is the rostral deflection, which probably reflects different degrees of specializations for bottom feeding (Domning, 2001). They seem to have had similar shape of their tusks, large and more-or-less oval in cross-section, this morphology most likely aided in obtaining large seagrass rhizomes (Domning, 2001; Domning & Beatty, 2007), although probably at different levels due to the differences in rostral deflection. Interestingly, it seems that there were no post-Eocene halitheriines.

*Domningia sodhae was aptly named after Dr. Daryl P. Domning of Howard University, world renowned paleosirenologist, a well deserved recognition.

Lateral view of B. kachchhense (top; from Bajpai & Domning, 1997) and Domningia sodhae (bottom; image reversed, from Thewissen & Bajpai, 2009).

Where are the halitheriines?

So far no post-Eocene halitheriines are known from the Oligocene and Miocene of India, at least two possible explanations come to mind: (1) they were present but no fossils have been found yet, or; (2) they were totally absent. An explanation for the second alternative could be provided by the invertebrate fauna. Based on gastropod fauna it seems that, during the Oligocene, changes in the geological settings in the Tethys Realm led to changes in the ocean currents and what was previously known as the Indo-African Region was subsequently divided into the Mediterranean-Iranian and Western Indian-Eastern African provinces of the Western Tethyan Region (Harzhauser et al. 2002). Subsequently, during the Early Miocene, further fragmentation of the Western Indian-Eastern African Province led to an increase of South-East Asian influence and the formation of the Proto-Western Indian Ocean Province; this separation was further augmented by the closure of the Eastern Mediterranean seaway during the latest Early Miocene (Harzhauser et al. 2002). The progressive isolation of what would become the Proto-Western Indian Ocean Province from a Tethyan influence might have prevented halitheriines from entering the region, especially after the Late Oligocene, leading to specialization and radiation of dugongines in this part of the world.

 References

Bajpai, S. & D. P. Domning. 1997. A new dugongine sirenian from the Early Miocene of India. Journal of Vertebrate Paleontology 17(1):219-228.

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

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

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

Harzhauser, M., W. E. Piller & F. F. Steininger. 2002. Circum-Mediterranean Oligo-Miocene biogeographic evolution – the gastropods’ point of view. Palaeogeography, Palaeoclimatology, Palaeoecology 183:103-133.

Kumar, K. & A. Sahni. 1986. Remingtonocetus harudiensis, new combination, a Middle Eocene Archeocete (Mammalia, Cetacea) from western Kutch, India. Journal of Vertebrate Paleontology 6(4):326-349.

Thewissen, J. G. M. & S. Bajpai. 2009. A new Miocene sirenian from Kutch, India. Acta Palaeontologica Polonica 54(1):7-13.