{{Short description|Extinct genus of mammals}} {{Automatic taxobox | fossil_range = Pleistocene to Holocene {{fossil range|1.9|0.0034}} | image = Información del camélido extinto, Paleolama - A740640.jpg | image_caption = ''Palaeolama'' fossils | display_parents = 2 | taxon = Palaeolama | authority = Gervais, 1869 | type_species = {{Extinct}}''Palaeolama weddeli'' | type_species_authority = Gervais, 1855 | subdivision_ranks = Species | subdivision = {{Extinct}}''P. aequatorialis'' <small>(Hoffstetter, 1952)</small><br/> {{Extinct}}''P. brevirostris'' <small>(Rusconi, 1930)</small><br/> {{Extinct}}''P. crassa'' <small>(Hoffstetter, 1952)</small><br/> {{Extinct}}''P. crequii'' <small>(Boule & Thévenin, 1920)</small><br/> {{Extinct}}''P. leptognata'' <small>(Ameghino, 1889)</small><br/> {{Extinct}}''P. major'' <small>(Liais, 1872)</small><br/> {{Extinct}}''P. mirifica'' <small>(Simpson, 1929)</small><br/> {{Extinct}}''P. niedae'' <small>(Guérin & Faure, 1999)</small><br/> {{Extinct}}''P. paradoxa'' <small>(Cabrera, 1935)</small><br/> {{Extinct}}''P. promesolithica'' <small>(Ameghino, 1889)</small><br/> {{Extinct}}''P. reissi'' <small>(Branco, 1883)</small><br/> {{Extinct}}''P. weddelli'' <small>(Gervais, 1855)</small><ref name="Fossilworks">{{cite web | title=Fossilworks: Palaeolama | website=Fossilworks | url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=42539 | access-date=2020-06-05}}</ref> }} '''''Palaeolama''''' ({{Lit|ancient llama}}) is an extinct genus of lamine camelids that existed from the Pleistocene to the Holocene ({{Ma|1.9|0.0034|Ma}}). Their range extended from North America to the intertropical region of South America.
== Description == ''Palaeolama'' species were relatives of modern lamines that lived in the New World from the Pleistocene around 1.9 million years ago to potentially the Holocene epoch around 3,353–4,231 years cal. Before Present (BP).<ref name="Ruez2005"/><ref>{{Cite journal |last1=Faria |first1=F. H. C. |last2=Carvalho |first2=I. S. |last3=Araújo-Júnior |first3=H. I. |last4=Ximenes |first4=C. L. |last5=Facincani |first5=E. M. |title=3,500 years BP: The last survival of the mammal megafauna in the Americas |year=2025 |journal=Journal of South American Earth Sciences |volume=153 |at=105367 |doi=10.1016/j.jsames.2025.105367 }}</ref> Fossil evidence suggests that it had a slender head, elongate snout, and stocky legs.<ref name="Dompierre2006">{{Cite book|last=Dompierre|first= H.|title= Observations on the diets of six late Cenozoic North American camelids: ''Camelops'', ''Hemiauchenia'', ''Palaeolama'', ''Procamelus'', ''Alforjas'', and ''Megatylopus''|year= 1995|publisher= National Library of Canada|isbn= 0-612-02748-1|oclc=46500746}}</ref><ref name="Fariña2013">{{Cite book|last1= Fariña|first1= R.A.|url= https://books.google.com/books?id=kUAKgNfiAvoC|jstor= j.ctt16gzd2q|title= Megafauna: Giant Beasts of Pleistocene South America|last2= Vizcaíno|first2= S.F.|last3=De Iuliis|first3= G.|date= 2013|publisher= Indiana University Press|isbn= 978-0-253-00230-3|oclc= 779244424}}</ref> They likely weighed around {{Convert|200|kg||abbr=}}<ref>Anthony J. Stuart, 2021, ''Vanished Giants: The Lost World of the Ice Age'', "7.19 Llamas: Palaeolama and Hemiauchenia", p.136, University of Chicago Press</ref> or up to {{Convert|300|kg||abbr=}}, surpassing the weight of modern llamas.<ref name="Fariña2013" /> They were specialized forest browsers and are often found in association with early equids, tapirs, deer, and mammoth.<ref name="Wheeler2006" /><ref name="Scherer2013" /><ref name="Guérin1999">{{cite journal|last1= Guérin|first1= C.|last2= Faure|first2= M.|title= ''Palaeolama'' (''Hemiauchenia'') ''niedae'' sp. nov. from Northeastern Brazil and its place among the South American Lamini|journal= Geobios|language= fr|volume= 32|issue= 4|year= 1999|pages= 629–659|doi= 10.1016/S0016-6995(99)80012-6}}</ref><ref name="Salas2003">{{Cite journal|last1= Salas|first1=R.|last2= Stucchi|first2=M.|last3= Devries|first3=T.J.|year= 2003|title=The presence of Plio-Pleistocene ''Palaeolama'' sp. (Artiodactyla: Camelidae) on the southern coast of Peru|journal= Bulletin de l'Institut français d'études andines|volume= 32|issue= 2|pages=347–359|doi=10.4000/bifea.6414|doi-access= free}}</ref>
=== Skull === ''Palaeolama'' had a long, slender skull with an elongated rostrum and robust jaw. This morphology more closely resembles the cranial morphology of ''Hemiauchenia'' than that of modern llamas.<ref name="Dompierre2006" />
=== Dental === The jaw and dental morphology of ''Palaeolama'' species distinguish them from other laminae. They tend to have a comparatively more dorsoventrally gracile mandible.<ref name="Dompierre2006" /><ref name="Ruez2005">{{Cite journal|last= Ruez|first= D.R.|date= 2005-09-30|title= Earliest record of ''Palaeolama'' (Mammalia, Camelidae) with comments on "''Palaeolama''" ''guanajuatensis''|journal= Journal of Vertebrate Paleontology|volume= 25|issue= 3|pages= 741–744|doi= 10.1671/0272-4634(2005)025[0741:eropmc]2.0.co;2|s2cid= 86522528}}</ref> Like ''Hemiauchenia'', ''Palaeolama'' species lack second deciduous premolars and can further be differentiated by the distinct size and shape of their third deciduous premolars. Their dentition has also been described as more brachyodont-like (short crowns, well-developed roots).<ref name="Ruez2005" />
=== Postcranial === Analyses of their limb elements reveal that they had shorter, stockier metapodials, and longer epipodials, giving them a short, stocky appearance.<ref name="Dompierre2006" /> Limbs such as these are typically associated with organisms adapted to walking on uneven and rugged terrains. This is also suggestive of being well-adapted to avoiding predators in forested areas.<ref name="Scherer2013" /><ref name="Dompierre2006" />
==Paleobiology== === Diet === Various dietary analyses have concluded that ''Palaeolama'' was a specialized forest browser that relied almost exclusively on plants high in C<sub>3</sub> for subsistence.<ref name="Guérin1999" /><ref name="Dompierre2006" /><ref name="EltinkEtAl2020">{{Cite journal |last1=Eltink |first1=Estevan |last2=Castro |first2=Mariela |last3=Montefeltro |first3=Felipe Chinaglia |last4=Dantas |first4=Mario André Trindade |last5=Scherer |first5=Carolina Saldanha |last6=de Oliveira |first6=Paulo Victor |last7=Langer |first7=Max Cardoso |date=March 2020 |title=Mammalian fossils from Gruta do Ioiô cave and past of the Chapada Diamantina, northeastern Brazil, using taphonomy, radiocarbon dating and paleoecology |url=https://linkinghub.elsevier.com/retrieve/pii/S0895981119302354 |journal=Journal of South American Earth Sciences |language=en |volume=98 |article-number=102379 |doi=10.1016/j.jsames.2019.102379 |bibcode=2020JSAES..9802379E |access-date=19 April 2024 |via=Elsevier Science Direct|url-access=subscription |hdl=11449/196635 |hdl-access=free }}</ref> Additionally, its shallow jaw and brachydont "cheek teeth" are highly suggestive of a mixed or intermediate seasonal diet consisting of primarily leaves and fruits, with some grass.<ref name="Marcolino2012">{{Cite journal|last1= Marcolino|first1= C.P.|last2= Isaias|first2= R.M. dos S.|last3= Cozzuol|first3= M.A.|last4= Cartelle|first4= C.|last5= Dantas|first5= M.A.T.|date= 2012|title=Diet of ''Palaeolama major'' (Camelidae) of Bahia, Brazil, inferred from coprolites |journal= Quaternary International |volume= 278|pages= 81–86|doi= 10.1016/j.quaint.2012.04.002|bibcode= 2012QuInt.278...81M}}</ref><ref name="MacFadden2000">{{Cite journal|last= MacFadden|first= B.J.|date= 2000|title= Cenozoic Mammalian Herbivores From the Americas: Reconstructing Ancient Diets and Terrestrial Communities|journal= Annual Review of Ecology and Systematics |volume= 31|issue= 1|pages= 33–59|doi= 10.1146/annurev.ecolsys.31.1.33|bibcode= 2000AnRES..31...33M}}</ref> Microwear analyses further validate this dietary interpretation.<ref name="Dompierre2006" /> Analysis of δ<sup>13</sup>C values from ''P. major'' remains in northeastern Brazil confirm it was primarily a consumer of C<sub>3</sub> plant matter.<ref name="EltinkEtAl2020" /> In the Melbourne Beds of Florida, ''P. mirifica'' shows evidence of being a leaf browser, while fossils of the same species from Ingleside, Texas are indicative of frugivory alongside folivory.<ref>{{Cite journal |last1=Semprebon |first1=Gina M. |last2=Rivals |first2=Florent |date=1 September 2010 |title=Trends in the paleodietary habits of fossil camels from the Tertiary and Quaternary of North America |url=https://www.sciencedirect.com/science/article/pii/S0031018210003202?casa_token=Kew0XUjDZqwAAAAA:OzpWNCQcR9BPBpO53SpyIWAXmwq3bHUMSRaJgP8fO_g09kU5s_BvjXbnaZ4QjzJPRmGhoyV6UUQ |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=295 |issue=1-2 |pages=131–145 |doi=10.1016/j.palaeo.2010.05.033 |access-date=10 July 2025 |via=Elsevier Science Direct|url-access=subscription }}</ref>
=== Group composition === [[File:Smilodon stalking Palaeolama.jpg|thumb|left|Two ''Smilodon populator'' stalking a ''Palaeolama major'' group in Brazil]] As inferred from observations of modern llama, ''Palaeolama'' probably organized into bands (consisting of a single male and multiple females) and troops (consisting exclusively of young males sometimes described as "bachelors"). Typically, band territories are defended by resident males, while troops remain more or less free-roaming until they form bands of their own.<ref name="Dompierre2006" />
== Habitat == thumb|right| Restoration of ''Palaeolama'' (below the gomphothere's tusks) and other mammals of Late Pleistocene Chile Fossil evidence suggests ''Palaeolama'' was primarily adapted to low-temperate, arid climates and preferred open, forested, and high-altitude mountainous regions.<ref name="Scherer2013">{{Cite journal|last= Scherer|first= C.S.|s2cid= 207195863|year= 2013|title=The Camelidae (Mammalia, Artiodactyla) from the Quaternary of South America: Cladistic and Biogeographic Hypotheses |journal= Journal of Mammalian Evolution |volume= 20|issue= 1|pages= 45–56|doi= 10.1007/s10914-012-9203-4}}</ref><ref name="Salas2003" /><ref name="Dompierre2006" /> The distribution of fossil evidence suggests that they had an altitudinal range limited exclusively by their dietary (vegetation) requirements.<ref name="Dompierre2006" /> Population density is shown to be highly dependent upon access and availability of subsistence resources.<ref name="Dompierre2006" /><ref>{{cite journal |last1=Ruez Jr. |first1=Dennis |title=Earliest Record of Palaeolama (Mammalia, Camelidae) with Comments on "Palaeolama" guanajuatensis |journal=Journal of Vertebrate Paleontology |date=September 30, 2005 |volume=25 |issue=3 |pages=741–744 |doi=10.1671/0272-4634(2005)025[0741:EROPMC]2.0.CO;2 |jstor=4524496 |s2cid=86522528 }}</ref>
== Range == The origins of this genus are a topic of much debate, as some of the earliest fossils occur during both the Irvingtonian in Florida and the Ensenadan in Uruguay.<ref name="Ruez2005" /> Despite this, agreement exists amongst paleobiologists on the dispersal of ''Palaeolama'' during the Great American Biotic Interchange.<ref name="Wheeler2006">{{cite book|chapter= Genetic Analysis of the Origins of Domestic South American Camelids|title= Documenting Domestication: New Genetic and Archaeological Paradigms |jstor= 10.1525/j.ctt1pnvs1.28 |pages= 329–341|author1= Wheeler, J.C.|author2= Chikhi, L.|author3= Bruford, M.W.|year= 2006|publisher= University of California Press|isbn= 978-0-520-24638-6|citeseerx= 10.1.1.603.88|editor1= Zeder, M.A.|editor2= Bradley, D.G.|editor3= Emshwiller, E.|editor4= Smith, B.D.}}</ref><ref name="Scherer2013" /><ref name="Ruez2005" /> Also, some evidence suggests a move to northern South America during the second of two Pleistocene Camilidae migration events.<ref name="Scherer2013" /> Fossil evidence ranges from the southern extent of North America (including California, Florida, and Mexico) south through Central America, and terminates in South America (Argentina and Uruguay).<ref name="Wheeler2006" /><ref name="Scherer2013" /><ref name="Salas2003" />
'''''Palaeolama mirifica''''', the "stout-legged llama", is known from southern California and the Southeastern U.S., with the highest concentration of fossil specimens found in Florida (specifically the counties of Alachua, Citrus, Hillsborough, Manatee, Polk, Brevard, Orange, Sumter, and Levy). Other fossil occurrences have been discovered in Mexico, Central America (El Salvador) and South America (Argentina, Bolivia, Brazil, Chile, Colombia, Ecuador, Paraguay, Peru, Venezuela and Uruguay).<ref>{{Cite web|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=52043|title=Fossilworks: Palaeolama mirifica|website=fossilworks.org}}</ref><ref>{{Cite web|url=https://www.theguardian.com/science/2020/nov/29/sistine-chapel-of-the-ancients-rock-art-discovered-in-remote-amazon-forest|title = 'Sistine Chapel of the ancients' rock art discovered in remote Amazon forest| website=TheGuardian.com |date = 29 November 2020}}</ref>
'''''Palaeolama major''','' identified by Liais in 1872, lived during the Late Pleistocene and was identified in fossil assemblages from northeastern and northern Brazil, the Pampean region of Argentina and Uruguay, northern Venezuela, and the coastal regions of Ecuador and northern Peru.<ref name="Scherer2013" />
'''''Palaeolama wedelli''''', identified by Gervais in 1855, lived during the Mid- to Late Pleistocene, with fossil specimens found in southern Bolivia and the Andean region of Ecuador.<ref name="Scherer2013" />
== Extinction == Climate change, changes and reductions in the types of vegetation on which they relied, and human predation are all hypothesized to have contributed to the extinction of ''Palaeolama'' during the Late Pleistocene or Early Holocene.<ref name="Dompierre2006" /><ref name="Ruez2005" /> Evidence from both the paleoecological and fossil records suggest that ''Palaeolama'', among other extinct camelids, weathered a number of glacial and interglacial episodes throughout their existence in North and South America. Their disappearance in some regions has been shown to coincide with a change in climate (to warmer, humid conditions) occurring at the end of the Pleistocene (also known as the Late Quaternary warming) suggesting an inability to persevere.<ref name="Dompierre2006" /><ref name="Barnosky2015">{{Cite journal|last1= Barnosky|first1= .D.|last2= Lindsey|first2= E.L.|last3= Villavicencio|first3= N.A.|last4= Bostelmann|first4= E.|last5= Hadly|first5= E.A.|last6= Wanket|first6= J.|last7 =Marshall|first7= C.R.|date=26 October 2015 |title= Variable impact of late-Quaternary megafaunal extinction in causing ecological state shifts in North and South America|journal= Proceedings of the National Academy of Sciences of the United States of America |volume= 113|issue= 4|pages= 856–861|doi= 10.1073/pnas.1505295112|pmid= 26504219|pmc= 4739530|doi-access= free}}</ref> This hypothesis is further supported by paleoecological evidence suggesting post-megafaunal extinction shifts in vegetation and whole ecosystems.<ref name="Barnosky2015" />
==See also== {{portal|Paleontology}} *List of North American animals extinct in the Holocene *List of South American animals extinct in the Holocene
==References== {{Reflist}}
{{Taxonbar|from=Q864122}}
Category:Prehistoric camelids Category:Pleistocene mammals of South America Category:Prehistoric mammals of North America Category:Pleistocene Artiodactyla Category:Prehistoric Artiodactyla genera Category:Fossil taxa described in 1869 Category:Pliocene Argentina