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Taxonomy

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Originally, James W. Gidley named this animal Arctodus floridanus in 1928,[1] until affinities with the spectacled bear were realised by Chester Stock when specimens from San Josecito Cave were recovered in 1950. Described as Tremarctos mexicanus, it was recombined as T. floridanus by Kurten (1963), Lundelius (1972) and Kurten and Anderson (1980).[2][3] Despite at least 35 individuals being recovered by 1967, almost all specimens consist of isolated teeth & skull fragments,[4] save for the type specimen (from the Golf Course site of the Melbourne Bone Bed in Melbourne, Florida).[5]

Evolution

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Tremarctinae within Ursidae

†Hemicyoninae

†Ursavinae

Ailuropodinae

Ursinae

Tremarctinae (short-faced bears)

†Plionarctos

†Arctodus

Tremarctos

†Arctotherium

The closest living relative of the Florida cave bear is the spectacled bear (T. ornatus) of South America; they are classified together with the huge short-faced bears in the subfamily Tremarctinae.[6] Unlike its Neotropical sister species (T. ornatus), T. floridanus was a temperate species that has only been recovered from Nearctic sites.[7][8] The fossil record of T. ornatus is unknown, as T. ornatus remains do not appear in the South American record until the Holocene,[9] which may indicate that T. ornatus emerged from T. floridanus in the Holocene.[6] However, as the montane niche (1,800m or higher) was otherwise open, T. ornatus may have instead evolved in the Pleistocene.[10] Recent genetic research suggests that Arctotherium either emerged from the Tremarctos genus or was a sister lineage to Tremarctos.[11]

The Interior Plateaus preserve the oldest possible remains of T. floridanus from the Palm Spring Formation (Anza-Borrego, California, ~2.7Ma), Grand View fauna (Glenns Ferry Formation, Idaho, 2.3Ma),[12][13][14] and San Simon (Arizona ca. 2.2Ma),[15][16] although the Grand View specimen may instead represent Plionarctos.[13] Additionally, though originally described as Arctodus sp.,[17] researchers suggest that indeterminate ursid from the mid-Blancan Buckhorn fauna (New Mexico, 4Ma - 3Ma) may represent either Tremarctos sp. or Ursus abtrusus.[14][18] Plionarctos harroldum is the likely ancestor of Tremarctos; a Plionarctos harroldum specimen from Taunton (Washington, 2.9Ma)[19] appears evolutionarily intermediate between P. harroldum and T. floridanus.[20]

Diagnostics

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Kurtén described T. floridanus as being morphologically very similar to its sister species T. ornatus, with practically identical dentitions. However, T. floridanus was of somewhat larger size, with some slight differences in relative proportions, and much more robust limb bones. Kurtén compared the differences between the Tremarctos species as the differences between brown bears and Eurasian cave bears.[21]

Where T. floridanus and the American black bear are co-specific, the M2 molar is used to differentiate between the species. Despite their similar sizes, the M2 of T. floridanus has a subrectangular form with truncated ends, a flat crown profile, a straight line on the lingual side, a high anterior region, and lacks a cingulum ridge.[21][22] Meanwhile, the M2 of Arctodus is broader and more robust than Tremarctos.[23]

Description

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Skull

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As in other tremarctine bears, tooth size (particularly the P4) could be variable, as exemplified by an average sized female from Tennessee possessing the largest known teeth in T. floridanus, with some even comparable to Arctodus.[4] The canalis semicircularis lateral suggests that T. floridanus had a head posture of 38°, which is more oblique than its sister species T. ornatus (29°); as T. ornatus inhabits densely vegetated areas, the more oblique head posture in T. floridanus could infer a greater capacity for long distance vision.[24] Tremarctos floridanus has condyles raised well above the plane of the teeth,[23] while T. ornatus does not (and therefore potentially possessed a larger gape).[25]

Postcranial

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T. floridanus is thought to have been almost twice the size of T. ornatus, averaging around the size of a large black bear.[4] Like T. ornatus and A. simus, T. floridanus possessed a false thumb.[26]

Paleobiology

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Sexual dimorphism

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Like other tremarctine bears, T. floridanus was highly sexually dimorphic.[27][23] However, a likely female, the Grassy Cove Saltpeter Cave specimen, was the size of a large black bear[4]

Paleopathology

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A T. floridanus individual from Grassy Cove Saltpeter Cave, Tennessee possibly preserves a fractured arm, caused by the 9 m (30 ft) drop into the lower cave level it was recovered from.[4]

Range

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Blancan and Irvingtonian

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The distribution of Tremarctos floridanus changed across its natural history. In the Blancan (4.75Ma to 1.806Ma) and Irvingtonian (1.806Ma to 250,000 BP) faunal stages, T. floridanus remains are sparse, and solely recovered from western North America.[5][28] The Late Blancan records Tremarctos sp. from San Simon, Arizona,[29] and T. floridanus possibly the Grand View fauna of Idaho (though this may be Plionarctos),[30] while Irvingtonian remains have been recovered from Palm Spring Formation in California,[29] and El Golfo in Sonora.[29][31][32] T. floridanus was recorded from the Blancan Hagerman Fossil Beds,[33] but these have been reassigned to Protarctos abstrusus.[14] Irvingtonian remains were reported from Cumberland Bone Cave in Maryland,[34] but subsequent research establishes Arctodus pristinus was the only tremarctine bear present at the locality.[35]

Rancholabrean

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However, in the proceeding Rancholabrean epoch (250,000 to 11,000 BP), T. floridanus was widely distributed along the eastern Atlantic Plain and the broader Gulf Coast. Fossils have been recovered from the US states of Florida,[5][7][8][36][37][38][39][40][41][42] Georgia,[4] Mississippi,[43][44] South Carolina,[45][23][22][46] Tennessee,[4] and Texas,[21][47][48][49][50] and the Mexican states of Michoacán,[51] and Nuevo León.[52][53][54]

T. floridanus is suggested to have expanded its range into higher altitudes during warmer interstadials and interglacials, which would explain its presence in Inner Space Cavern on Edwards Plateau, Texas.[55] The Grassy Cove Saltpeter Cave specimen, from Cumberland Plateau, Tennessee, has been radiocarbon dated to within the climatically unstable MIS 3.[4][56]

T. floridanus has been described from Belize,[54][57] however the recent reassignment of a T. floridanus specimen to the morphologically similar Arctotherium wingei from the Hoyo Negro, and other positive identifications of A. wingei from Belize have put under question current T. floridanus identifications in Yucatán Peninsula. Current scholarly analysis asserts that A. wingei may have restricted the range of T. floridanus outside of Central & South America until the extinction of A. wingei, where subsequently Tremarctos begins to be found in the South America.[58][59]

Additionally, T. floridanus specimens assigned by Kurten in New Mexico (such as Isleta Caves and Conkling Cavern) have been reassigned to Arctodus simus - no western specimens of T. floridanus have been found in the Rancholabrean.[60]

Map of fossil localities

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SuperTah/sandbox is located in North America
San Simon Creek (Tremarctos sp.)
Grand View fauna (Tremarctos floridanus?)
Vallecito Creek? (Anza-Borrego)
El Golfo (Tremarctos cf. T. floridanus)
La Cinta Portalitos
San Josecito Cave (San Josito)
Ingleside Gravel Pit
Inner Space Cavern (Laubach No. 3)
Yazoo County, Golden Triangle Regional Airport (Town Creek?)
Myrtle Beach
Edisto Beach
Crowfield Local Fauna, Runnymede Plantation (Tremarctos sp.)
Ardis Local Fauna
Grassy Cove Saltpeter Cave (33,660 ± 3980 BP)
McFaddin Beach
Ladd's Quarry (Laddas)
Arredondo 1A; Arredondo 1G; Arredondo 2A
Ichetucknee River
Hornsby Sink / Spring
Merritt Island
Bone Cave
Lecanto 2A, Lecanto 3A
Sabertooth Cave
Cutler Hammock, Monkey Jungle Hammock 1
Peace River 3A, 8A
St. Johns River Dredge
Aucilla River 1A, Aucilla River 2C, Sloth Hole
Winter Beach / Luther Locality
Seminole Springs
Seminole Field
Millennium Park
Zeta Pond
Williston 3A
Manatee Springs (Manatee Sink)
Withlacoochee River 5A, Withlacoochee River 7A
Waccasassa River 9, Waccasassa River 10
Reddick 1, Reddick 1B, Reddick 1C, Reddick 1D
Eichelberger Cave
Ocklawaha River 2A
Orange Lake 2A
North Fernandina Beach
Ponte Vedra Beach
Rocky Creek
Daytona Beach Bone Beds
Blue Springs, St Petersberg?
Rock Springs
Haile 19C; Haile 7A; Haile 8A
Lake Rousseau
Santa Fe River 1B, Santa Fe River 2, Santa Fe River 6B
Melbourne
Vero Canal Site (Vero Beach)
Devil's Den, Octopus Cave
Interactive distribution map of Tremarctos floridanus (hover over dots for locality information)

Legend: Late Blancan (Pliocene / Early Pleistocene)

Irvingtonian (Early / Middle Pleistocene)

Rancholabrean (Late Pleistocene)

Paleoecology

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Atlantic Plain

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The Atlantic Plain (in particular Florida) hosts the greatest number of T. floridanus specimens on the continent, coming from well known sites such as Aucilla River, Cutler Fossil Site, Devil's Den Cave, Haile Quarry site, Melbourne Bone Bed and Rock Springs.[5] Despite this, Tremarctos floridanus is unknown before the Rancholabrean faunal stage, with Arctodus pristinus, a tremarctine bear inhabiting eastern North America during the Blancan and Irvingtonian, being very similar to T. floridanus in terms of size, skeletal anatomy, and dietary preferences.[61] Despite this, generally speaking large tremarctine fossils from the Early and Middle Pleistocene of Florida are considered to be A. pristinus, whereas those from the Late Pleistocene of Florida are considered to be T. floridanus. Indeed, black bears and Tremarctos floridanus are believed to have only colonized Florida with the extinction of A. pristinus (both of which only appear in Florida in the Late Pleistocene), however, T. floridanus could yet still be found from older sites in Florida.[61] T. floridanus was possibly an ecological replacement of A. pristinus, with T. floridanus finds becoming widespread in Rancholabrean Florida and the wider southeastern United States.[61][62][63]

At the Rainbow River and Lake Rousseau localities in Rancholabrean Florida, Tremarctos floridanus have been recovered alongside Smilodon, dire wolves, jaguars, ground sloths (Megalonyx, Paramylodon), llamas (Hemiauchenia, Palaeolama), Vero's tapir, giant beaver, capybara, Holmesina, horses, Bison antiquus, mastodon, Columbian mammoths and Arctodus, in a climate similar to today's. Furthermore, the abundance of black bears, and particularly Tremarctos floridanus in Florida, has led to a theorized niche partitioning of ursids in Florida, with Tremarctos floridanus being herbivorous, and black bears and Arctodus simus being omnivorous, with Arctodus being possibly more inclined towards carnivory.[64]

During the Rancholabrean, Florida and the Gulf Coast hosted a subtropical faunal region. Sometimes referred to as the Holmesina ("Chlamytherium")-glyptodont faunal province, T. floridanus was joined by other key indicator species such as Holmesina, Glyptotherium, capybaras, Eremotherium, Cuvieronius, Hesperotestudo, Mylohyus, Palaeolama, and Dasypus bellus.[65][66] The now submerged continental shelf in the Gulf of Mexico appears to have provided a dispersal corridor consisting of savanna and thorn scrub habitats between south Texas and Florida for these largely Neotropical fauna.[49][8] Generally in the southeast, interior regions were dominated by boreal forest, which transitioned into a unique spruce-pine-oak forest between the Last Glacial Maximum and Younger Dryas as the climate became warmer. However, in Florida this trend was reversed, with warm-wet adapted Pinus forest occupying central Florida during colder periods, and cooler & drier oak-scrub prairie becoming dominant during interstadials.[67]

Appalachian Highlands

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The Grassy Cove Saltpeter Cave specimen was radiocarbon dated to 33,660 ± 3980 radiocarbon BP.[56] Should T. floridanus have been present at the same locality after the Last Glacial Maximum, the Rancholabrean mid-Appalachians would have constituted open forest (dominated by jack pine), dense forest-edges and grasslands, with boreal, temperate and mid-western prairie elements. Other fauna would have included oxen (Ovibos, Bootherium), bison, horse, tapir, peccaries (Mylohyus, Platygonus), Cervalces, caribou, elk, white-tailed deer, ground sloths (Glossotherium, Megalonyx), mammoth, mastodon, giant beaver, brown bear, black bear, jaguar, the dire wolf, Smilodon, Acinonyx, and Arctodus.[68] An isotope study on a T. floridanus individual from Ladd's Quarry, Georgia broadly reconstructed the specimen as an omnivore.[69]

Interior Plains

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Laubach III in the Inner Space Cavern (dated to ca. 23,230 ± 490 radiocarbon BP) of the Interior Plains contains many animals that have modern relatives associated with subtropical climates, including Tremarctos floridanus, common opossum, Mexican free-tailed bat, jaguar, armadillo, and Glyptotherium. The fauna appears to have been deposited under a generally warm climate (inter-stadial).[55]

Intermontane Plateaus

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The Intermontane Plateaus is the only region to preserve T. floridanus remains from the Blancan and Irvingtonian faunal stages, in addition to the Rancholabrean.[5][28] T. floridanus may have evolved from Plionarctos harroldum, as a potential transitional fossil may have been recorded from Taunton (Washington, 2.9Ma).[19][20] Dubious remains have been recovered from New Mexico (Buckhorn Fauna, 4Ma - 3Ma)[14][17][18] and Idaho (Grand View fauna, 2.3Ma),[12][13][14] while validated remains are recorded from California (Palm Spring Formation, ~2.7Ma) and San Simon (Tremarctos sp., 2.2Ma).[15][16]

In the Anza-Borrego Desert, T. floridanus has been recovered from the Arroyo Seco local fauna (Late Pliocene, 3.041Ma - 2.581Ma), Vallecito Creek local fauna (Early Pleistocene, 1.95Ma - 1.77Ma) and the Borrego Badlands.[70][71][72] Appearing in the Late Blancan (~2.7Ma), the persistence of T. floridanus into the Irvingtonian age Vallecito Creek local fauna was part of a gradual faunal transition, which blended traditionally Blancan fauna (Stegomastodon, Titanotylopus, Bensonomys, Equus simplicidens) with typically Irvingtonian fauna (such as T. floridanus, Tetrametryx, other Equus sp., Lepus and Camelops) in the Late Blancan and Early Irvingtonian.[71]

In the Rancholabrean highlands of the Mexican Plateau, San Josecito Cave in Nuevo León hosted an extension of the Southern Plains, which in Nuevo León were flanked to the west by the Sierra Madre Oriental. T. floridanus would have inhabited a savanna, with trees primarily restricted to the grassland's waterways.[52] At La Cinta-Portalitos (Michoacán/Guanajuato) in the Trans-Mexican Volcanic Belt, prime habitat for Tremarctos floridanus were the gallery forests (Alnus, Fraxinus, Salix) and their wetlands (Phragmites, Typha), along with white-tailed deer, capybaras, Pampatherium, horses, and Cuvieronius, with its relative Arctodus simus preferring the closed temperate forests of the Madrean pine–oak woodlands.[73]

Extinction

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While once thought to have had a possible continuation into the Greenlandian stage of the Holocene from presumed 8,000 years old material from the Devil's Den Cave,[74] subsequent research indicates the fossils present were from the Rancholabrean epoch instead.[75] While T. floridanus may have dissapeared in North America, the species may have evolved into T. ornatus in South America, either in the Pleistocene or the Holocene.[6][10]

References

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  1. ↑ Gidley, James W. (1928). "A new species of bear from the Pleistocene of Florida". Journal of the Washington Academy of Sciences. 18 (15): 430–433. ISSN 0043-0439.
  2. ↑ E. L. Lundelius. 1972. Bureau of Economic Geology Report of Investigations 77.
  3. ↑ Kurtén and Anderson: 178-80
  4. 1 2 3 4 5 6 7 8 Guilday, John E.; Irving, David C. (October 1967). "Extinct Florida Spectacled Bear Tremarctos floridanus (Gidley) From Central Tennessee" (PDF). Bulletin of the National Speleological Society. 29 (4): 149–162.
  5. 1 2 3 4 5 Harrington, Arianna (April 10, 2015). "Tremarctos floridanus". Florida Museum (University of Florida). Archived from the original on October 30, 2021. Retrieved February 26, 2022.
  6. 1 2 3 Soibelzon, Leopoldo H.; Tonni, Eduardo P.; Bond, Mariano (2005-10-01). "The fossil record of South American short-faced bears (Ursidae, Tremarctinae)". Journal of South American Earth Sciences. Quaternary Paleontology and biostratigraphy of southern South Africa. 20 (1): 105–113. doi:10.1016/j.jsames.2005.07.005. ISSN 0895-9811.
  7. 1 2 Morgan, Gary S. (2002). "Late Rancholabrean Mammals from Southernmost Florida, and the Neotropical Influence in Florida Pleistocene Faunas". Biodiversity Heritage Library. Retrieved 2025-06-21.
  8. 1 2 3 Morgan, Gary S.; Emslie, Steven D. (2010-04-15). "Tropical and western influences in vertebrate faunas from the Pliocene and Pleistocene of Florida". Quaternary International. Faunal Dynamics and Extinction in the Quaternary: Studies in Honor of Ernest L. Lundelius, Jr. 217 (1): 143–158. doi:10.1016/j.quaint.2009.11.030. ISSN 1040-6182.
  9. ↑ Stucchi, Marcelo; Salas-Gismondi, Rodolfo; Baby, Patrice; Guyot, Jean-Loup; Shockey, Bruce J. (April 2004). "A 6,000+ year-old specimen of a spectacled bear from an Andean cave in Peru". Ursus. 20 (1): 63–68. doi:10.2192/08GR017R1.1. ISSN 1537-6176.
  10. 1 2 Soibelzon, Leopoldo H.; Rincón, Ascanio D. (2007-07-27). "The fossil record of the short-faced bears (Ursidae, Tremarctinae) from Venezuela. Systematic, biogeographic, and paleoecological implications". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen: 287–298. doi:10.1127/0077-7749/2007/0244-0287.
  11. ↑ Schubert, B. W.; Chatters, J. C.; Arroyo-Cabrales, J.; Samuels, J. X.; Soibelzon, L. H.; Prevosti, F. J.; Widga, C.; Nava, A.; Rissolo, D.; Erreguerena, P. L. (2019). "Yucatán carnivorans shed light on the Great American Biotic Interchange". Biology Letters. 15 (5): 20190148. doi:10.1098/rsbl.2019.0148. PMC 6548739. PMID 31039726. 20190148.{{cite journal}}: CS1 maint: article number as page number (link)
  12. 1 2 Janis, Christine M.; Scott, Kathleen M.; Jacobs, Louis L. (1998-05-28). Evolution of Tertiary Mammals of North America: Volume 1, Terrestrial Carnivores, Ungulates, and Ungulate Like Mammals. Cambridge University Press. ISBN 978-0-521-35519-3.
  13. 1 2 3 White Jr., Richard S.; Morgan, Gary S. (2005). "Arizonan Blancan Vertebrate Faunas in Regional Perspective". Vertebrate Paleontology of Arizona, Mesa Southwest Museum Bulletin. 11.
  14. 1 2 3 4 5 Ruez, Dennis R. (June 2009). "Revision of the Blancan (Pliocene) mammals from Hagerman Fossil Beds National Monument, Idaho". Journal of the Idaho Academy of Science. 45 (1) – via ResearchGate.
  15. 1 2 Thrasher, Larry (July 2022). "Fossils and Age Relationships of the Late Pliocene and Early Pleistocene (Blancan) 111 Ranch Beds and Bear Springs Wash Beds, Graham County, Arizona". Late Cenozoic Vertebrate Paleontology: Tribute to Arthur H. Harris. New Mexico Museum of Natural History and Science Bulletin. 88 – via ResearchGate.
  16. 1 2 Morgan, Gary S.; White Jr., Richard S. (2005). "Miocene and Pliocene vertebrates from Arizona". Vertebrate Paleontology in Arizona. New Mexico Museum of Natural History and Science Bulletin. 29 – via ResearchGate.
  17. 1 2 Morgan, G. S.; Seale, P. L. (May 1995). "New Mexico Geological Society Spring meeting abstracts-1 995" (PDF). New Mexico Geological Society. 17 (2).
  18. 1 2 Morgan, Gary S.; Sealey, Paul L.; Lucas, Spencer G.; Heckert, Andrew B. (1997). "Pliocene (Latest Hemphillian and Blancan) Vertebrate Fossils from the Mangas Basin, Southwestern New Mexico" (PDF). New Mexico Museum of Natural History and Science Bulletin No. 11 – via ResearchGate.
  19. 1 2 Morgan, James K.; Morgan, Neil H. (1995). "A New Species of Capromeryx (Mammalia: Artiodactyla) from the Taunton Local Fauna of Washington, and the Correlation with Other Blancan Faunas of Washington and Idaho". Journal of Vertebrate Paleontology. 15 (1): 160–170. ISSN 0272-4634.
  20. 1 2 Tedford, Richard H.; and Martin, James (2001-07-20). "Plionarctos, a tremarctine bear (Ursidae: Carnivora) from western North America". Journal of Vertebrate Paleontology. 21 (2): 311–321. doi:10.1671/0272-4634(2001)021[0311:PATBUC]2.0.CO;2. ISSN 0272-4634.
  21. 1 2 3 Kurtén, Björn (June 1, 1963). "Fossil Bears from Texas" (PDF). The Pearce-Sellard Series (1): 13 – via University of Texas Libraries.
  22. 1 2 Albright, L.; Sanders, Albert; Weems, Robert; Cicimurri, David; Knight, James (2019-10-31). "Cenozoic vertebrate biostratigraphy of South Carolina, U.S.A., and additions to the fauna". Bulletin of the Florida Museum of Natural History. 57 (2): 77–236. doi:10.58782/flmnh.qqgg4577. ISSN 2373-9991.
  23. 1 2 3 4 Sanders, Albert E. (2002). "Additions to the Pleistocene Mammal Faunas of South Carolina, North Carolina, and Georgia". Transactions of the American Philosophical Society. 92, Part 5: 33–40. ISBN 9780871699251 – via Google Books.
  24. ↑ Arnaudo, Maria Eugenia; Bona, Paula; Soibelzon, Leopoldo Hector; Schubert, Blaine W. (December 2016). "Anatomical study of the auditory region of Arctotherium tarijense (Ursidae, Tremarctinae), an extinct short-faced bear from the Pleistocene of South America". Journal of Anatomy. 229 (6): 825–837. doi:10.1111/joa.12525. ISSN 0021-8782. PMC 5108154. PMID 27460048.
  25. ↑ Figueirido; et al. (2010). "Demythologizing Arctodus simus, the 'short-faced' long-legged and predaceous bear that never was". Journal of Vertebrate Paleontology. 30 (1): 262–275. Bibcode:2010JVPal..30..262F. doi:10.1080/02724630903416027. hdl:10630/33066. S2CID 85649497.
  26. ↑ Salesa, M. J.; Siliceo, G.; Antón, M.; Abella, J.; Montoya, P.; Morales, J. (2006-12-30). "Anatomy of the "false thumb" of Tremarctos ornatus (Carnivora, Ursidae, Tremarctinae): phylogenetic and functional implications". Estudios Geológicos. 62 (1): 389–394. doi:10.3989/egeol.0662133. ISSN 1988-3250.
  27. ↑ Gazin, C. Lewis (15 December 1950). "Paleontology: Annotated list of fossil Mammalia associated with human remains at Melbourne, Fla" (PDF). Journal of the Washington Academy of Sciences – via U.S. National Museum.
  28. 1 2 Kurtén, B.; Anderson, E. Pleistocene Mammals of North America. Columbia University Press. ISBN 0230613993.
  29. 1 2 3 "Tremarctos floridanus-Florida spectacled bear". UTEP Biodiversity Collections. 3 April 2013. Retrieved February 27, 2022.
  30. ↑ White Jr., Richard S.; Morgan, Gary S. (2005). "Arizonan Blancan Vertebrate Faunas in Regional Perspective". Vertebrate Paleontology of Arizona, Mesa Southwest Museum Bulletin. 11.
  31. ↑ Croxen III, Fred W.; Shaw, Christopher A.; Sussman, David R. (March 2007). "Pleistocene Geology and Paleontology of the Colorado River Delta at Golfo de Santa Clara, Sonora, Mexico". Arizona Western College – via ResearchGate.
  32. ↑ Shaw†, Christopher A.; Croxen†, Fred W., III (2019-09-04), Pearthree, Philip A. (ed.), "Geology and paleontology of the early-middle Pleistocene El Golfo beds, Sonora, Mexico—A field guide", Geologic Excursions in Southwestern North America, vol. 55, Geological Society of America, p. 0, ISBN 978-0-8137-0055-7, retrieved 2025-06-03{{citation}}: CS1 maint: multiple names: authors list (link) CS1 maint: work parameter with ISBN (link)
  33. ↑ McDonald, Dr. Greg (January 1995). "The Fossil Record: Critter Corner" (PDF). Hagerman Fossil Beds National Monument. 4 (1).
  34. ↑ "PBDB Collection". paleobiodb.org. Retrieved 2025-06-05.
  35. ↑ Eshelman, Ralph E.; Bell, Christopher; Graham, Russel W.; Semken Jr., Holmes A.; Withnell, Charles B.; Scarpetta, Simon G.; James, Helen F.; Godfrey, Stephen J.; Mead, Jim I. (2025-04-18). Middle Pleistocene Cumberland Bone Cave Local Fauna, Allegany County, Maryland: A Systematic Revision and Paleoecological Interpretation of the Irvingtonian, Middle Appalachians, USA. Smithsonian Institution Scholarly Press.
  36. ↑ Hemmings, C. Andrew (2005). "An Update on Recent Work at Sloth Hole (8JE121), Aucilla River, Jefferson County, Florida" (PDF). Current Research In The Pleistocene. 22: 47–49.
  37. ↑ Schubert, Blaine W.; Hulbert, Richard C.; Macfadden, Bruce J.; Searle, Michael; Searle, Seina (2010-01). "Giant short-faced bears ( Arctodus simus ) in Pleistocene Florida USA, a substantial range extension". Journal of Paleontology. 84 (1): 79–87. doi:10.1666/09-113.1. ISSN 0022-3360. {{cite journal}}: Check date values in: |date= (help)
  38. ↑ "Peace River Paleo Project (PRiPP)". Vertebrate Paleontology Collection. Retrieved 2025-06-19.
  39. ↑ Carr, Robert S. (2012). Digging Miami. Gainesville, Florida: University Press of Florida. p. 32. ISBN 978-0-8130-4206-0.
  40. ↑ Purdy, Barbara A. (2008). Florida's People During the Last Ice Age. Gainesville, Florida: University Press of Florida. ISBN 978-0-8130-3204-7.
  41. ↑ "Seminole Field". Florida Vertebrate Fossils. Retrieved 2025-06-17.
  42. ↑ Gut, H. James; Ray, Clayton E. (1963). "The Pleistocene Vertebrate Fauna of Reddick, Florida". Quarterly Journal of the Florida Academy of Sciences. 26 (4): 315–328. ISSN 0015-3850.
  43. ↑ "Fossil Friday – MDEQ". www.mdeq.ms.gov. Retrieved 2025-06-03.
  44. ↑ "Bulletin 128: A List, Bibliography And Index Of The Fossil Vertebrates Of Mississippi – MDEQ". www.mdeq.ms.gov. Retrieved 2025-06-03.
  45. ↑ "Tremarctos sp., Bear tooth". The Charleston Museum. Retrieved February 27, 2022.
  46. ↑ Bentley, Curtis C.; Bentley, Curtis C.; Knight, James L.; Knoll, Martin A. (1994). "The mammals of the Ardis local fauna (late Pleistocene), Harleyville, South Carolina". Brimleyana. 21: 1––35.
  47. ↑ Smith, Felisa; Tomé, Catalina; Smith, Emma Elliott; Lyons, S.; Newsome, Seth; Stafford, Thomas (2016-01-01). "Unraveling the consequences of the terminal Pleistocene megafauna extinction on mammal community assembly". School of Biological Sciences: Faculty Publications.
  48. ↑ Austin~cjbell@jsg.utexas.edu, Christopher J. Bell~The University of Texas at; Wbg004@shsu.edu, William Godwin~Sam Houston State University~; University~kelsey.jenkins@yale.edu, Kelsey M. Jenkins~Yale; University~pjl001@shsu.edu, Patrick J. Lewis~Sam Houston State (2020-09-28). "First fossil manatees in Texas: Trichechus manatus bakerorum in the Pleistocene fauna from beach deposits along the Texas Coast of the Gulf of Mexico". Palaeontologia Electronica. Retrieved 2025-06-05.{{cite web}}: CS1 maint: numeric names: authors list (link)
  49. 1 2 Moretti, John A.; Flores, Deanna; Bell, Christopher J.; Godwin, Will; Hartstone-Rose, Adam; Lewis, Patrick J. (2024-04-23). "The scimitar-cat Homotherium from the submerged continental shelf of the Gulf Coast of Texas". Anatomical Record (Hoboken, N.J.: 2007). doi:10.1002/ar.25461. ISSN 1932-8494. PMID 38654480.
  50. ↑ "McFaddin Beach". www.texasbeyondhistory.net. Retrieved 2025-06-21.
  51. ↑ Eng-Ponce, Joaquin (August 2021). "Reconstruccion paeloambiental del yacimiento La Cinta-Portalitos, Michoacan-Guanajuato, Mexico (thesis)" (PDF). Faculty of Biology, Universidad Michoacana de San Nicolás de Hidalgo.
  52. 1 2 Arroyo-Cabrales, Joaquin; Johnson, Eileen; Cruz, J. Alberto (2021-10-26). "San Josecito Cave and Its Paleoecological Contributions for Quaternary Studies in Mexico". Quaternary. 4 (4): 34. doi:10.3390/quat4040034. ISSN 2571-550X. Archived from the original on 2023-01-12.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  53. ↑ Stock, Chester (1950). "Bears from the Pleistocene Cave of San Josecito, Nuevo León, Mexico". Journal of the Washington Academy of Sciences. 40 (10): 317–321. ISSN 0043-0439.
  54. 1 2 Churcher, C. S. (March 2020). "Pleistocene mammals from Extinction Cave, Belize". Canadian Journal of Earth Sciences. 57 (3): 366–376. Bibcode:2020CaJES..57..366C. doi:10.1139/cjes-2018-0178. S2CID 182629185.
  55. 1 2 Toomey, III, Rickard S. (1994). "Vertebrate Paleontology of Texas Caves". In Elliott, W. R.; Veni, G. (eds.). The Caves and Karst of Texas (PDF). National Speleological Society, Huntsville, Alabama. pp. 51–68.
  56. 1 2 Jr, Robert Stuckenrath; Mielke, James E. (January 1970). "Smithsonian Institution Radiocarbon Measurements VI". Radiocarbon. 12 (1): 193–204. doi:10.1017/S0033822200036286. ISSN 0033-8222.
  57. ↑ Czaplewski, Nicolas J.; Krejca, Jean; Miller, Thomas E. (2003). "Late Quaternary Bats from Cebada Cave, Chiquibul Cave System,Belize" (PDF). Caribbean Journal of Science. 39: 23 – via Belize Foundation for Research and Environmental Education.
  58. ↑ Schubert, B. W.; Chatters, J. C.; Arroyo-Cabrales, J.; Samuels, J. X.; Soibelzon, L. H.; Prevosti, F. J.; Widga, C.; Nava, A.; Rissolo, D.; Erreguerena, P. L. (2019). "Yucatán carnivorans shed light on the Great American Biotic Interchange". Biology Letters. 15 (5): 20190148. doi:10.1098/rsbl.2019.0148. PMC 6548739. PMID 31039726. 20190148.{{cite journal}}: CS1 maint: article number as page number (link)
  59. ↑ Schubert, Blaine W.; Chatters, James C.; Arroyo-Cabrales, Joaquin; Soibelzon, Leopoldo H.; Awe, Jaime J.; Griffith, Cameron S.; De Anda, Guillermo; Luna Erreguerena, Pilar (October 28, 2016). "The South American Short -Faced Bear Arctotherium from the Yucatán Peninsula of Belize and Mexico: Implications for their Biogeography, Paleobiology, Evolution and Extinction" (PDF). Society of Vertebrate Paleontology. SVP 76th Annual Meeting: 220.
  60. ↑ Lucas, Spencer G.; Sullivan, Robert M. Vertebrate Paleontology in New Mexico: Bulletin 68. New Mexico Museum of Natural History and Science.
  61. 1 2 3 Emslie, Steven D. (1995). "The fossil record of Arctodus pristinus (Ursidae: Tremarctinae) in Florida" (PDF). Bulletin of the Florida Museum of Natural History. 37 (15): 501–514. doi:10.58782/flmnh.hduf9651. S2CID 168164209.
  62. ↑ Russell, Dale A.; Rich, Fredrick J.; Schneider, Vincent; Lynch-Stieglitz, Jean (May 2009). "A warm thermal enclave in the Late Pleistocene of the South-eastern United States". Biological Reviews. 84 (2): 173–202. doi:10.1111/j.1469-185X.2008.00069.x. PMID 19391200. S2CID 9609391.
  63. ↑ Esker, Donald; Wilkins, William; Agenbroad, Larry (2010-08-13). "Esker, Wilkins, and Agenbroad—Multivariate Analysis Of Ursids: A multivariate analysis of the ecology of North American Pleistocene bears, with a focus on Arctodus simus". ResearchGate.
  64. ↑ Schubert, Blaine; Hulbert, Richard; MacFadden, Bruce; Searle, Michael; Searle, Seina (2010-01-01). "Giant Short-faced Bears (Arctodus simus) in Pleistocene Florida USA, a Substantial Range Extension". Journal of Paleontology. 84 (1): 79–87. Bibcode:2010JPal...84...79S. doi:10.1666/09-113.1. S2CID 131532424.
  65. ↑ Martin, Larry; Neuner, A. (1978-01-01). "The End of the Pleistocene in North America". Transactions of the Nebraska Academy of Sciences and Affiliated Societies.
  66. ↑ "A Cultural Resource Survey of the Continental Shelf from Cape Hatteras to Key West. Volume I, Introduction and Physical Environment" (PDF). Interior Department, Bureau of Land Management. 1. June 1981. {{cite journal}}: |first= missing |last= (help)CS1 maint: multiple names: authors list (link)
  67. ↑ Widga, Chris; Anderson, Derek T; Walker, Renee B (2021). "Plant and Animal Communities in the Southeastern US during the Late Pleistocene". doi:10.13140/RG.2.2.14378.13766. {{cite journal}}: Cite journal requires |journal= (help)
  68. ↑ Guilday, John E. (1982). "Appalachia 11,000-12,000 Years Ago: A Biological Review". Archaeology of Eastern North America. 10: 22–26. ISSN 0360-1021.
  69. ↑ Zenako, Paula. "What's For Dinner? An Isotopic Analysis of Pleistocene Mammals of the American Southeast" (PDF). University of Maryland (thesis).
  70. ↑ Murray, Lyndon Keith (December 2008). "Effects of taxonomic and locality inaccuracies on biostratigraphy and biochronology of the Hueso and Tapiado formations in the Vallecito Creek-Fish Creek section, Anza-Borrego Desert, California (Masters thesis)". University of Texas at Austin Libraries.
  71. 1 2 Cassiliano, Michael L. (1999). "Biostratigraphy of Blancan and Irvingtonian Mammals in the Fish Creek-Vallecito Creek Section, Southern California, and a Review of the Blancan-Irvingtonian Boundary". Journal of Vertebrate Paleontology. 19 (1): 169–186. ISSN 0272-4634.
  72. ↑ Cassiliano, Michael (1997-03-01). "Taphonomy of mammalian fossils across the Blancan-Irvingtonian boundary: Palm Spring Formation, Anza-Borrego desert of southern California". Palaeogeography, Palaeoclimatology, Palaeoecology. 129 (1): 81–108. doi:10.1016/S0031-0182(96)00123-X. ISSN 0031-0182.
  73. ↑ Eng-Ponce, Joaquin (August 2021). "Reconstruccion paeloambiental del yacimiento La Cinta-Portalitos, Michoacan-Guanajuato, Mexico (thesis)" (PDF). Faculty of Biology, Universidad Michoacana de San Nicolás de Hidalgo.
  74. ↑ Kurtén and Anderson: 56, 178-79
  75. ↑ Purdy, Barbara A.; Rohlwing, Kathryn M.; MacFadden, Bruce J. (2015-07-01). "Devil's Den, Florida: Rare Earth Element Analysis Indicates Contemporaneity of Humans and Latest Pleistocene Fauna". PaleoAmerica. 1 (3): 266–275. doi:10.1179/2055556315Z.00000000032. ISSN 2055-5563.
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