Eocene Antarctica:a window into the earliest history of modern whales
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摘要
The Eocene–Oligocene Southern Ocean is thought to have played a major role in cetacean evolution.Yet,fossils from its heart—Antarctica—are rare,and come almost exclusively from the Eocene La Meseta and Submeseta formations of Marambio(Seymour)Island.Here,we provide a summary and update of this crucial fossil assemblage,and discuss its relevance in the broader context of cetacean evolution.To date,Eocene specimens from Antarctica include basilosaurids,a group of archaic stem cetaceans that had already fully adapted to life in water;and the archaic toothed mysticete Llanocetus,the second oldest crown cetacean on record(ca.34 Ma).This Eocene co-occurrence of stem and crown cetaceans is highly unusual,and otherwise only observed in Peru.Though related,at least some of the Antarctic species appear to be different from,and notably larger than their Peruvian counterparts,suggesting an early differentiation of the high latitude cetacean fauna.
The Eocene–Oligocene Southern Ocean is thought to have played a major role in cetacean evolution.Yet,fossils from its heart—Antarctica—are rare,and come almost exclusively from the Eocene La Meseta and Submeseta formations of Marambio(Seymour)Island.Here,we provide a summary and update of this crucial fossil assemblage,and discuss its relevance in the broader context of cetacean evolution.To date,Eocene specimens from Antarctica include basilosaurids,a group of archaic stem cetaceans that had already fully adapted to life in water;and the archaic toothed mysticete Llanocetus,the second oldest crown cetacean on record(ca.34 Ma).This Eocene co-occurrence of stem and crown cetaceans is highly unusual,and otherwise only observed in Peru.Though related,at least some of the Antarctic species appear to be different from,and notably larger than their Peruvian counterparts,suggesting an early differentiation of the high latitude cetacean fauna.
The Eocene–Oligocene Southern Ocean is thought to have played a major role in cetacean evolution.Yet,fossils from its heart—Antarctica—are rare,and come almost exclusively from the Eocene La Meseta and Submeseta formations of Marambio(Seymour)Island.Here,we provide a summary and update of this crucial fossil assemblage,and discuss its relevance in the broader context of cetacean evolution.To date,Eocene specimens from Antarctica include basilosaurids,a group of archaic stem cetaceans that had already fully adapted to life in water;and the archaic toothed mysticete Llanocetus,the second oldest crown cetacean on record(ca.34 Ma).This Eocene co-occurrence of stem and crown cetaceans is highly unusual,and otherwise only observed in Peru.Though related,at least some of the Antarctic species appear to be different from,and notably larger than their Peruvian counterparts,suggesting an early differentiation of the high latitude cetacean fauna.
引文
Bebej R M,Zalmout I S,El-Aziz A A A,et al.2015.First remingtonocetid archaeocete(Mammalia,Cetacea)from the middle Eocene of Egypt with implications for biogeography and locomotion in early cetacean evolution.J Paleontol,89:882-893.
Berger W H.2007.Cenozoic cooling,Antarctic nutrient pump,and the evolution of whales.Deep-Sea Res Part II:Top Stud Oceanogr,54:2399-2421.
Borsuk-Bia?ynicka M.1988.New remains of Archaeoceti from the Paleogene of Antarctica.Pol Polar Res,9:437-445.
Buono M R,Fernández M S,Reguero M A,et al.2016.Eocene basilosaurid whales from the La Meseta Formation,Marambio(Seymour)Island,Antarctica.Ameghiniana,53:296-315.
Clementz M T,Fordyce R E,Peek S L,et al.2014.Ancient marine isoscapes and isotopic evidence of bulk-feeding by Oligocene cetaceans.Palaeogeogr Palaeocl,400:28-40.
Concheyro A,Amenábar C,Santillana S,et al.2016.Primer registro de nanofósiles calcáreos en la Formación La Meseta(Eoceno medio),Isla Marambio,Antártida.Proceedings of the 11th Congreso de la Asociación Paleontológica,Argentina:Abstract 126.
DeméréT A,McGowen M R,Berta A,et al.2008.Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales.Syst Biol,57:15-37.
Dingle,R V,Lavelle M.1998.Antarctic Peninsular cryosphere:Early Oligocene(c.30 Ma)initiation and a revised glacial chronology.JGeol Soc,155:433-437.
Douglas P M J,Affek H P,Ivany L C,et al.2014.Pronounced zonal heterogeneity in Eocene southern high-latitude sea surface temperatures.Proc Natl Acad Sci USA,111:6582-6587.
Ekdale E G,DeméréT A,Berta A.2015.Vascularization of the gray whale palate(Cetacea,Mysticeti,Eschrichtius robustus):soft tissue evidence for an alveolar source of blood to baleen.Anat Rec,298:691-702.
Elliot D H,Rinaldi C,Zinsmeister W J,et al.1975.Geological investigations on Seymour Island.Antarc J U S,10:182-186.
Elliot D H,Trautman T A.1982.Lower Tertiary strata on Seymour Island,Antarctic Peninsula//Craddock C.Antarctic Geoscience.Madison:University of Wisconin Press,287-297.
élouard P.1966.Découverte d'un archéocète dans les environs de Kaolack.Notes Afr,109:8-10.
Fordyce R E.1977.The development of the circum-Antarctic current and the evolution of the Mysticeti(mammalia:Cetacea).Palaeogeogr Palaeocl,21:265-271.
Fordyce R E.1980.Whale evolution and Oligocene Southern Ocean environments.Palaeogeogr Palaeoclimatol Palaeoecol,31:319-336.
Fordyce R E.1989.Origins and evolution of Antarctic marine mammals.Geol Soc Spec Pub,47:269-281.
Fordyce R E.2003.Cetacean evolution and Eocene-Oligocene oceans revisited//Prothero D R,Ivany L C,Nesbitt E A.From Greenhouse to Icehouse-The Marine Eocene-Oligocene Transition.New York:Columbia University Press,154-170.
Fordyce R E,Marx F G.2018.Gigantism precedes filter feeding in baleen whale evolution.Curr Biol,28:1670-1676.e1672.http://dx.doi.org/10.1016/j.cub.2018.04.027.
Fostowicz-Frelik L.2003.An enigmatic whale tooth from the upper Eocene of Seymour Island,Antarctica.Pol Polar Res,24:13-28.
Geisler J H,Sanders A E,Luo Z-X.2005.A new protocetid whale(Cetacea:Archaeoceti)from the Late Middle Eocene of South Carolina.Am Mus Novit,3480:1-65.
Geisler J H,Boessenecker R W,Brown M,et al.2017.The origin of filter feeding in whales.Curr Biol,27:2036-2042.e2032.http://dx.doi.org/10.1016/j.cub.2017.06.003.
Gingerich P D,Smith B H,Simons E L.1990.Hind limbs of Eocene Basilosaurus:evidence of feet in whales.Science,249:154-157.
Gingerich P D,Haq M U,Zalmout I S,et al.2001.Origin of whales from early artiodactyls:hands and feet of Eocene Protocetidae from Pakistan.Science,293:2239-2242.
Gingerich P D,Cappetta H.2014.A new archaeocete and other marine mammals(Cetacea and Sirenia)from lower Middle Eocene phosphate deposits of Togo.J Paleontol,88:109-129.
Gingerich P D,Zouhri S.2015.New fauna of archaeocete whales(Mammalia,Cetacea)from the Bartonian middle Eocene of southern Morocco.J Afr Earth Sci,111:273-286.
Gol’din P,Zvonok E.2013.Basilotritus uheni,a new cetacean(Cetacea,Basilosauridae)from the late Middle Eocene of Eastern Europe.JPaleontol,87:254-268.
Hernández Cisneros A E,Tsai C-H.2016.A possible enigmatic kekenodontid(Cetacea,Kekenodontidae)from the Oligocene of Mexico.Paleontol Mex,5:147-155.
Kellogg R.1936.A review of the Archaeoceti.Carnegie Inst Wash Publ,482:1-366.
K?hler R,Fordyce R E.1997.An archaeocete whale(Cetacea:Archaeoceti)from the Eocene Waihao Greensand,New Zealand.J Vertebr Paleontol,17:574-583.
Lambert O,Martínez-Cáceres M,Bianucci G,et al.2017.Earliest mysticete from the Late Eocene of Peru sheds new light on the origin of baleen whales.Curr Biol,27:1535-1541.e1532.
Lambert O,Bianucci G,Salas-Gismondi R,et al.2019.An amphibious whale from the Middle Eocene of Peru reveals early South Pacific dispersal of quadrupedal cetaceans.Curr Biol.http://dx.doi.org/10 .1016/j.cub.2019.02.050(published online).
Loch C,Buono M R,Kalthoff D C,et al.2019.Enamel microstructure in Eocene cetaceans from Antarctica(Archaeoceti and Mysticeti).JMamm Evol.https://doi.org/10.1007/s10914-018-09456-3(published online).
Marenssi S A,Santillana S N,Rinaldi C A.1998.Stratigraphy of the La Meseta Formation(Eocene),Marambio(Seymour)Island,Antarctica.Asoc Paleontol Argent Publ Espec,5:137-146.
Martínez Cáceres M,Lambert O,Muizon C d.2017.The anatomy and phylogenetic affinities of Cynthiacetus peruvianus,a large Dorudonlike basilosaurid(Cetacea,Mammalia)from the late Eocene of Peru.Geodiversitas,39:7-163.
Marx F G,Fordyce R E.2015.Baleen boom and bust:a synthesis of mysticete phylogeny,diversity and disparity.R Soc Open Sci,2:140434.
Marx F G,Lambert O,Uhen M D.2016a.Cetacean Paleobiology.Chichester:Wiley Blackwell.
Marx F G,Hocking D P,Park T,et al.2016b.Suction feeding preceded filtering in baleen whale evolution.Mem Mus Vic,75:71-82.
Marx F G,Buono M R,Evans A R,et al.2019.Gigantic mysticete predators roamed the Eocene Southern Ocean.Antarc Sci,31:98-104.
Mitchell E D.1989.A new cetacean from the Late Eocene La Meseta Formation,Seymour Island,Antarctic Peninsula.Can J Fish Aquat Sci,46:2219-2235.
Montes M,Nozal F,Santillana S,et al.2013.Mapa Geológico de la isla Marambio(Seymour)Escala 1:20.000 Primera Edición.Serie Cartográfica Geocientífica Antártica.Madrid/Buenos Aires:Instituto Geológico y Minero de Espa?a and Instituto Antártico Argentino.
Pivorunas A.1979.The feeding mechanisms of baleen whales.Am Sci,67:432-440.
Sadler P M.1988.Geometry and stratification of uppermost Cretaceous and Paleogene units on Seymour Island,northern Antarctic Peninsula.Geol Soc Am Mem,169:303-320.
Sarmiento J L,Gruber N,Brzezinski M A,et al.2004.High-latitude controls of thermocline nutrients and low latitude biological productivity.Nature,427:56-60.
Slater G J,Goldbogen J A,Pyenson N D.2017.Independent evolution of baleen whale gigantism linked to Plio-Pleistocene ocean dynamics.Proc R Soc B,284:20170546.
Thewissen J G M,Hussain S T,Arif M.1994.Fossil evidence for the origin of aquatic locomotion in archaeocete whales.Science,263:210-212.
Thewissen J G M,Williams E M,Roe L J,et al.2001.Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls.Nature,413:277-281.
Uhen M D.1998.Middle to Late Eocene basilosaurines and dorudontines//Thewissen J G M.The emergence of whales.New York:Plenum Press,29-61.
Uhen M D.1999.New species of protocetid archaeocete whale,Eocetus wardii(Mammalia:Cetacea)from the Middle Eocene of North Carolina.J Paleontol,73:512-528.
Uhen M D,Pyenson N D.2007.Diversity estimates,biases and historiographic effects:resolving cetacean diversity in the Tertiary.Palaeontol Electron,10:10.12.10A.http://palaeo-electronica.org/paleo/2007_2/00123/index.html
Uhen M D.2008.New protocetid whales from Alabama and Mississippi,and a new cetacean clade,Pelagiceti.J Vertebr Paleontol,28:589-593.
Uhen M D,Pyenson N D,Devries T J,et al.2011.New Middle Eocene whales from the Pisco Basin of Peru.J Paleontol,85:955-969.
Uhen M D.2013.A review of North American Basilosauridae.Bull Alabama Mus Nat Hist,31:1-45.
Wiman C.1905.über die altterti?ren Vertebraten der Seymourinsel.Wissenschaftliche Ergebnisse der Schwedischen Südpolar-Expedition1901-1903,Vol 3.Stockholm:Lithographisches institut des Generalstabs,1-37.
Woodburne M O,Zinsmeister W J.1984.The first land mammal from Antarctica and its biogeographic implications.J Paleontol,58:913-948.
Wrenn J H,Hart G F.1988.Paleogene dinoflagellate cyst biostratigraphy of Seymour Island,Antarctica.Geol Soc Am Mem,169:321-447.
UCR:University of California Riverside;now housed at the Museum of Paleontology of the University of California,Berkeley,USA
USNM:United States National Museum of Natural History,Washington DC,USA
Berger W H.2007.Cenozoic cooling,Antarctic nutrient pump,and the evolution of whales.Deep-Sea Res Part II:Top Stud Oceanogr,54:2399-2421.
Borsuk-Bia?ynicka M.1988.New remains of Archaeoceti from the Paleogene of Antarctica.Pol Polar Res,9:437-445.
Buono M R,Fernández M S,Reguero M A,et al.2016.Eocene basilosaurid whales from the La Meseta Formation,Marambio(Seymour)Island,Antarctica.Ameghiniana,53:296-315.
Clementz M T,Fordyce R E,Peek S L,et al.2014.Ancient marine isoscapes and isotopic evidence of bulk-feeding by Oligocene cetaceans.Palaeogeogr Palaeocl,400:28-40.
Concheyro A,Amenábar C,Santillana S,et al.2016.Primer registro de nanofósiles calcáreos en la Formación La Meseta(Eoceno medio),Isla Marambio,Antártida.Proceedings of the 11th Congreso de la Asociación Paleontológica,Argentina:Abstract 126.
DeméréT A,McGowen M R,Berta A,et al.2008.Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales.Syst Biol,57:15-37.
Dingle,R V,Lavelle M.1998.Antarctic Peninsular cryosphere:Early Oligocene(c.30 Ma)initiation and a revised glacial chronology.JGeol Soc,155:433-437.
Douglas P M J,Affek H P,Ivany L C,et al.2014.Pronounced zonal heterogeneity in Eocene southern high-latitude sea surface temperatures.Proc Natl Acad Sci USA,111:6582-6587.
Ekdale E G,DeméréT A,Berta A.2015.Vascularization of the gray whale palate(Cetacea,Mysticeti,Eschrichtius robustus):soft tissue evidence for an alveolar source of blood to baleen.Anat Rec,298:691-702.
Elliot D H,Rinaldi C,Zinsmeister W J,et al.1975.Geological investigations on Seymour Island.Antarc J U S,10:182-186.
Elliot D H,Trautman T A.1982.Lower Tertiary strata on Seymour Island,Antarctic Peninsula//Craddock C.Antarctic Geoscience.Madison:University of Wisconin Press,287-297.
élouard P.1966.Découverte d'un archéocète dans les environs de Kaolack.Notes Afr,109:8-10.
Fordyce R E.1977.The development of the circum-Antarctic current and the evolution of the Mysticeti(mammalia:Cetacea).Palaeogeogr Palaeocl,21:265-271.
Fordyce R E.1980.Whale evolution and Oligocene Southern Ocean environments.Palaeogeogr Palaeoclimatol Palaeoecol,31:319-336.
Fordyce R E.1989.Origins and evolution of Antarctic marine mammals.Geol Soc Spec Pub,47:269-281.
Fordyce R E.2003.Cetacean evolution and Eocene-Oligocene oceans revisited//Prothero D R,Ivany L C,Nesbitt E A.From Greenhouse to Icehouse-The Marine Eocene-Oligocene Transition.New York:Columbia University Press,154-170.
Fordyce R E,Marx F G.2018.Gigantism precedes filter feeding in baleen whale evolution.Curr Biol,28:1670-1676.e1672.http://dx.doi.org/10.1016/j.cub.2018.04.027.
Fostowicz-Frelik L.2003.An enigmatic whale tooth from the upper Eocene of Seymour Island,Antarctica.Pol Polar Res,24:13-28.
Geisler J H,Sanders A E,Luo Z-X.2005.A new protocetid whale(Cetacea:Archaeoceti)from the Late Middle Eocene of South Carolina.Am Mus Novit,3480:1-65.
Geisler J H,Boessenecker R W,Brown M,et al.2017.The origin of filter feeding in whales.Curr Biol,27:2036-2042.e2032.http://dx.doi.org/10.1016/j.cub.2017.06.003.
Gingerich P D,Smith B H,Simons E L.1990.Hind limbs of Eocene Basilosaurus:evidence of feet in whales.Science,249:154-157.
Gingerich P D,Haq M U,Zalmout I S,et al.2001.Origin of whales from early artiodactyls:hands and feet of Eocene Protocetidae from Pakistan.Science,293:2239-2242.
Gingerich P D,Cappetta H.2014.A new archaeocete and other marine mammals(Cetacea and Sirenia)from lower Middle Eocene phosphate deposits of Togo.J Paleontol,88:109-129.
Gingerich P D,Zouhri S.2015.New fauna of archaeocete whales(Mammalia,Cetacea)from the Bartonian middle Eocene of southern Morocco.J Afr Earth Sci,111:273-286.
Gol’din P,Zvonok E.2013.Basilotritus uheni,a new cetacean(Cetacea,Basilosauridae)from the late Middle Eocene of Eastern Europe.JPaleontol,87:254-268.
Hernández Cisneros A E,Tsai C-H.2016.A possible enigmatic kekenodontid(Cetacea,Kekenodontidae)from the Oligocene of Mexico.Paleontol Mex,5:147-155.
Kellogg R.1936.A review of the Archaeoceti.Carnegie Inst Wash Publ,482:1-366.
K?hler R,Fordyce R E.1997.An archaeocete whale(Cetacea:Archaeoceti)from the Eocene Waihao Greensand,New Zealand.J Vertebr Paleontol,17:574-583.
Lambert O,Martínez-Cáceres M,Bianucci G,et al.2017.Earliest mysticete from the Late Eocene of Peru sheds new light on the origin of baleen whales.Curr Biol,27:1535-1541.e1532.
Lambert O,Bianucci G,Salas-Gismondi R,et al.2019.An amphibious whale from the Middle Eocene of Peru reveals early South Pacific dispersal of quadrupedal cetaceans.Curr Biol.http://dx.doi.org/10 .1016/j.cub.2019.02.050(published online).
Loch C,Buono M R,Kalthoff D C,et al.2019.Enamel microstructure in Eocene cetaceans from Antarctica(Archaeoceti and Mysticeti).JMamm Evol.https://doi.org/10.1007/s10914-018-09456-3(published online).
Marenssi S A,Santillana S N,Rinaldi C A.1998.Stratigraphy of the La Meseta Formation(Eocene),Marambio(Seymour)Island,Antarctica.Asoc Paleontol Argent Publ Espec,5:137-146.
Martínez Cáceres M,Lambert O,Muizon C d.2017.The anatomy and phylogenetic affinities of Cynthiacetus peruvianus,a large Dorudonlike basilosaurid(Cetacea,Mammalia)from the late Eocene of Peru.Geodiversitas,39:7-163.
Marx F G,Fordyce R E.2015.Baleen boom and bust:a synthesis of mysticete phylogeny,diversity and disparity.R Soc Open Sci,2:140434.
Marx F G,Lambert O,Uhen M D.2016a.Cetacean Paleobiology.Chichester:Wiley Blackwell.
Marx F G,Hocking D P,Park T,et al.2016b.Suction feeding preceded filtering in baleen whale evolution.Mem Mus Vic,75:71-82.
Marx F G,Buono M R,Evans A R,et al.2019.Gigantic mysticete predators roamed the Eocene Southern Ocean.Antarc Sci,31:98-104.
Mitchell E D.1989.A new cetacean from the Late Eocene La Meseta Formation,Seymour Island,Antarctic Peninsula.Can J Fish Aquat Sci,46:2219-2235.
Montes M,Nozal F,Santillana S,et al.2013.Mapa Geológico de la isla Marambio(Seymour)Escala 1:20.000 Primera Edición.Serie Cartográfica Geocientífica Antártica.Madrid/Buenos Aires:Instituto Geológico y Minero de Espa?a and Instituto Antártico Argentino.
Pivorunas A.1979.The feeding mechanisms of baleen whales.Am Sci,67:432-440.
Sadler P M.1988.Geometry and stratification of uppermost Cretaceous and Paleogene units on Seymour Island,northern Antarctic Peninsula.Geol Soc Am Mem,169:303-320.
Sarmiento J L,Gruber N,Brzezinski M A,et al.2004.High-latitude controls of thermocline nutrients and low latitude biological productivity.Nature,427:56-60.
Slater G J,Goldbogen J A,Pyenson N D.2017.Independent evolution of baleen whale gigantism linked to Plio-Pleistocene ocean dynamics.Proc R Soc B,284:20170546.
Thewissen J G M,Hussain S T,Arif M.1994.Fossil evidence for the origin of aquatic locomotion in archaeocete whales.Science,263:210-212.
Thewissen J G M,Williams E M,Roe L J,et al.2001.Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls.Nature,413:277-281.
Uhen M D.1998.Middle to Late Eocene basilosaurines and dorudontines//Thewissen J G M.The emergence of whales.New York:Plenum Press,29-61.
Uhen M D.1999.New species of protocetid archaeocete whale,Eocetus wardii(Mammalia:Cetacea)from the Middle Eocene of North Carolina.J Paleontol,73:512-528.
Uhen M D,Pyenson N D.2007.Diversity estimates,biases and historiographic effects:resolving cetacean diversity in the Tertiary.Palaeontol Electron,10:10.12.10A.http://palaeo-electronica.org/paleo/2007_2/00123/index.html
Uhen M D.2008.New protocetid whales from Alabama and Mississippi,and a new cetacean clade,Pelagiceti.J Vertebr Paleontol,28:589-593.
Uhen M D,Pyenson N D,Devries T J,et al.2011.New Middle Eocene whales from the Pisco Basin of Peru.J Paleontol,85:955-969.
Uhen M D.2013.A review of North American Basilosauridae.Bull Alabama Mus Nat Hist,31:1-45.
Wiman C.1905.über die altterti?ren Vertebraten der Seymourinsel.Wissenschaftliche Ergebnisse der Schwedischen Südpolar-Expedition1901-1903,Vol 3.Stockholm:Lithographisches institut des Generalstabs,1-37.
Woodburne M O,Zinsmeister W J.1984.The first land mammal from Antarctica and its biogeographic implications.J Paleontol,58:913-948.
Wrenn J H,Hart G F.1988.Paleogene dinoflagellate cyst biostratigraphy of Seymour Island,Antarctica.Geol Soc Am Mem,169:321-447.
UCR:University of California Riverside;now housed at the Museum of Paleontology of the University of California,Berkeley,USA
USNM:United States National Museum of Natural History,Washington DC,USA