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Systematic Paleontology of a Late Pleistocene Avifauna from Saint Lucie County, Florida

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Title:
Systematic Paleontology of a Late Pleistocene Avifauna from Saint Lucie County, Florida
Creator:
Kilmer, John A
Place of Publication:
[Gainesville, Fla.]
Florida
Publisher:
University of Florida
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Language:
english
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1 online resource (74 p.)

Thesis/Dissertation Information

Degree:
Master's ( M.S.)
Degree Grantor:
University of Florida
Degree Disciplines:
Zoology
Biology
Committee Chair:
STEADMAN,DAVID W
Committee Co-Chair:
MACFADDEN,BRUCE J
Committee Members:
KIMBALL,REBECCA T
Graduation Date:
5/2/2015

Subjects

Subjects / Keywords:
Beaches ( jstor )
Birds ( jstor )
Eggshells ( jstor )
Fossils ( jstor )
Humerus ( jstor )
Paleontology ( jstor )
Parakeets ( jstor )
Species ( jstor )
Ulna ( jstor )
Vertebrates ( jstor )
Biology -- Dissertations, Academic -- UF
aves -- dickerson -- florida -- fossil -- pleistocene
St. Lucie County ( local )
Genre:
bibliography ( marcgt )
theses ( marcgt )
government publication (state, provincial, terriorial, dependent) ( marcgt )
born-digital ( sobekcm )
Electronic Thesis or Dissertation
Zoology thesis, M.S.

Notes

Abstract:
Dickerson Coquina Pit is a sand and coquina mine located in Saint Lucie County, Florida. Before open-pit operations ceased at the site, collections were made from a vertebrate fossil bed of Middle Pleistocene (Late Irvingtonian) age in the Okeechobee Formation. Specimens from the coquina pit were also discovered in fill material used in beach replenishment on Hutchinson Island, St. Lucie County. The avifauna of the two sites contains over 70 elements representing 12 orders, 16 families, and 26 species. A surprisingly wide variety of species with affinities ranging from woodlands to the open ocean are represented in this relatively small fossil assemblage. Extant and recently extinct species found include a transitional turkey Meleagris aff. gallopavo, an albatross Phoebastria aff. albatrus, Great Cormorant Phalacrocorax carbo, a large stork Ciconia maltha, a large crane Grus sp., Great Auk Pinguinus impennis, and Carolina Parakeet Conuropsis carolinensis. The Dickerson Coquina Pit taphonomy and paleoecology are most similar to those of other shell beds in Central and South Florida such as the Irvingtonian Leisey Shell Pit of Hillsborough County, Florida. ( en )
General Note:
In the series University of Florida Digital Collections.
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Includes vita.
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Includes bibliographical references.
Source of Description:
Description based on online resource; title from PDF title page.
Source of Description:
This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis:
Thesis (M.S.)--University of Florida, 2015.
Local:
Adviser: STEADMAN,DAVID W.
Local:
Co-adviser: MACFADDEN,BRUCE J.
Statement of Responsibility:
by John A Kilmer.

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UFRGP
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Copyright Kilmer, John A. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Classification:
LD1780 2015 ( lcc )

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1 SYSTEMATIC PALEONTOLOGY O F A MIDDLE PLEISTOCENE AVIFAUNA FROM SAINT LUCIE COUNTY, FLORIDA By JOHN ANDREW KILMER A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIR EMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 201 5

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2 © 201 5 John Andrew Kilmer

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3 This thesis is gratefully dedicated to my parents , Galen and Laura Kilmer .

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4 ACKNOWLEDGMENTS I thank m y a dvisor, David Steadman, for his supervision and patience during the course of this study. I am indebted to my committee members Rebecc a Kimball and Bruce MacFadden for their time and sound advice . I am also thankful for the help and advice of Roger Portel l and Richard Hulbert , whose conversations aided me early into this work. Paul Roth and the dozens of fossil collectors and preparators that discovered, cleaned, and catalogued the specimens cannot be forgotten . Most of all, I wish to thank my friends and family members who constantly buoyed my spirits with their support : Tina Man Chen , Rachael Kilmer , and, most importantly, my parents Galen and Laura Kilmer. I co without you!

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5 TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................ ................................ ................................ .. 4 LIST OF TABLES ................................ ................................ ................................ ............ 7 LIST OF FIGURES ................................ ................................ ................................ .......... 8 LIST OF ABBREVIATIONS ................................ ................................ ........................... 10 ABSTRACT ................................ ................................ ................................ ................... 11 CHAPTER 1 INTRODUCTION ................................ ................................ ................................ .... 12 Discovery of the Dickerson Coquina Pit Specimens ................................ ............... 12 Geology of the Dickerson Coquina Pit ................................ ................................ .... 13 Biochronology of the Dickerson Coquina Pit ................................ ........................... 16 North Hutchinso n Island Beach Fill Specimens ................................ ...................... 17 2 METHODS ................................ ................................ ................................ .............. 22 3 RESULTS ................................ ................................ ................................ ............... 23 Systemat ic Paleontology ................................ ................................ ......................... 23 Order ANSERIFORMES ................................ ................................ ......................... 23 Family ANATIDAE ................................ ................................ ............................ 23 Genus A ix Boie, 1828 ................................ ................................ ................ 23 Genus Anas Linnaeus, 1758 ................................ ................................ ...... 24 Genus Aythya Boie, 1822 ................................ ................................ .......... 25 Genus Bucephala Baird, 1858 ................................ ................................ ... 26 Order GALLIFORMES ................................ ................................ ............................ 26 Family PHASIANIDAE ................................ ................................ ...................... 26 Genus Meleagris Linnaeus, 1758 ................................ .............................. 26 Order PODICIPEDIFORMES ................................ ................................ .................. 28 Family PODICIPEDIDAE ................................ ................................ .................. 28 Genus Podilymbus Lesson, 1831 ................................ .............................. 28 Order PROCELLARIIFORMES ................................ ................................ ............... 29 Family DIOMEDEIDAE ................................ ................................ ..................... 29 Genus Phoebastria Reichenbach, 1853 ................................ .................... 29 Order CICONIIFORMES ................................ ................................ ......................... 30 Family CICONIIDAE ................................ ................................ ......................... 30 Genus Ciconia Brisson, 1760 ................................ ................................ ..... 30 Order SULIFORMES ................................ ................................ .............................. 31 Family SULIDAE ................................ ................................ .............................. 31

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6 Genus Morus Vieillot, 1816 ................................ ................................ ........ 31 Family PHALACROCORACIDAE ................................ ................................ ..... 32 Genus Pha lacrocorax Brisson, 1780 ................................ .......................... 32 Family ANHINGIDAE ................................ ................................ ....................... 33 Genus Anhinga Brisson 1760 ................................ ................................ .... 33 Order PELECANIFORMES ................................ ................................ ..................... 34 Family ARDEIDAE ................................ ................................ ........................... 34 Genus Ardea Linnaeus, 1758 ................................ ................................ .... 34 Genus Nyctanassa Stejneger, 1887 ................................ .......................... 34 Family THRESKIORNITHIDAE ................................ ................................ ........ 35 Genus Eudocimus Wagler, 1832 ................................ ............................... 35 Order ACCIPITRIFORMES ................................ ................................ ..................... 35 Family ACCIPITRIDAE ................................ ................................ ..................... 35 Genus Haliaeetus Savigny, 1809 ................................ ............................... 35 Genus Buteo Lacépède, 1799 ................................ ................................ ... 36 Order GRUIFORMES ................................ ................................ ............................. 36 Family RALLIDAE ................................ ................................ ............................ 36 Genus Rallus Linnaeus, 1758 ................................ ................................ .... 36 Genus Fulica Linnaeus, 1758 ................................ ................................ .... 37 Fami ly GRUIDAE ................................ ................................ ............................. 37 Genus Grus Brisson, 1760 ................................ ................................ ......... 37 Order CHARADRIIFORMES ................................ ................................ ................... 38 Family ALCIDAE ................................ ................................ .............................. 38 Genus Pinguinus Bonnaterre, 1791 ................................ ........................... 38 Order STRIGIFORMES ................................ ................................ .......................... 40 Family STRIGIDAE ................................ ................................ .......................... 40 Genus Bubo Duméril, 1805 ................................ ................................ ........ 40 Genus Strix Linnaeus, 1758 ................................ ................................ ....... 40 Order PSITTACIFORMES ................................ ................................ ...................... 41 Family PSITTACIDAE ................................ ................................ ...................... 41 Genus Conuropsis Salvadori, 1891 ................................ ........................... 41 4 DISCUSSION ................................ ................................ ................................ ......... 64 5 CONCLUSIONS ................................ ................................ ................................ ..... 67 REFERENCES ................................ ................................ ................................ .............. 68 BIOGRAPHICAL SKETCH ................................ ................................ ............................ 74

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7 LIST OF TABLES Table page 3 1 Birds identified from the Dickerson Coquina Pit (DCP) and from beach fill derive d from DCP on North Hutchinson Island (HI), St. Lucie County, Florida ... 43 3 2 Measurements (mm ± SD) of various skeletal elements in select Irvingtonian and modern Meleagris ................................ ................................ ........................ 45 3 3 Measurements (mm ± SD) of the carpometacarpus in Phoebastria ................... 46 3 4 Measurements (mm ± SD) of the distal tibiotarsus in Grus ................................ 46

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8 LIST OF FIGURES Figure page 1 1 Map of Florida and St. Lucie County (inset) showing the locations of Pleistocene sites discussed in the text ................................ ............................... 18 1 2 Dickerson Coquina Pit 2 (DCP) in 2006. Photo by Roger Portell ........................ 19 1 3 North Hutchinson Island (HI) between Jenson Beach and Walton Rocks Beach wh ere fill from the Dickerson Coquina Pit (DCP) was deposited ............. 19 1 4 Stratigraphic section of Dickerson Coquina Pit 2 (DCP), St. Lucie County, Florida (ad apted from Herrera et al. 2006) ................................ ......................... 20 1 5 A brief overview of previous stratigraphic nomenclature of the Pleistocene deposits of St. Lucie County ................................ ................................ ............... 21 3 1 Pectoral and hindlimb elements of dabbling ducks (Anatidae) from Dickerson Coquina Pit, Florida compared w ith modern specimens ................................ ..... 47 3 2 Pectoral and hindlimb elements of Mallards ( Anas platyrhynchos ) from Dickerson C oquina Pit, Florida compared with modern specimens .................... 48 3 3 Wing elements of diving ducks (Aythyini and Mergini) from Dickerson Coquina Pit, Florida compared with modern specimens ................................ ..... 49 3 4 Pectoral and hindlimb elements of turkeys ( Meleagris aff. gallopavo ) from Dickerson Coquina Pit, Florida compared with modern specimens .................... 50 3 5 Additional pectoral and hindlimb elements of turkeys ( Meleagris aff. gallopavo ) from Dickerson Coquina Pit, Florida compared with modern specimens ................................ ................................ ................................ .......... 51 3 6 The humeri in palmar A B) aspect a nd tarsometatarsi in anterior C E) aspect of Pied billed Grebes ( Podilymbus podiceps ) from Dickerson Coquina Pit, Florida compared with modern specimens ................................ ......................... 52 3 7 The coracoids and carpometacarpi of Phoebastria albatrosses ......................... 53 3 8 The proximal carpometacarpi and distal tarsometatarsi of fossil storks (Ciconiidae) ................................ ................................ ................................ ........ 54 3 9 Fossi l Northern Gannet ( Morus bassanus ) and Anhinga ( Anhinga anhinga ) from Dickerson Coquina Pit, Florida compared with modern specimens ............ 55 3 10 Wing and hindlimb elements of cormorants ( Phalacroco rax ) from Dickerson Coquina Pit, Florida compared with modern specimens ................................ ..... 56

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9 3 11 Fossils of herons (Ardeidae) and White I bises ( Eudocimus albus ) from Dickerson Coquina Pit, Florida compared wit h modern specimens .................... 57 3 12 Fossils of hawks and eagles (Accipitridae) from Dickerson Coquina Pit, Florida compared with modern specimens ................................ ......................... 58 3 13 The right humeri of rails ( Rallus elegans ) and the right distal ulna and right distal tibiotarsus of coots ( Fulica americana ) from Dickerson Coquina Pit, Florida compared with modern specimens ................................ ......................... 59 3 14 The phalanxes and distal tibiotarsi of cranes ( Grus ) ................................ ........... 60 3 15 Humeri, scapulae, and tibiotarsi of Great Auks ( Pinguinus impennis ) ................ 61 3 16 Ulnae of Great Auks ( Pinguinus impennis ) ................................ ......................... 62 3 17 The humeri, tarsometatarsi, and femora of owls (Strigidae) from Dickerson Coquina Pit, Florida compared with moder n specimens ................................ ..... 62 3 18 The humeri of parakeets ................................ ................................ ..................... 63

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10 LIST OF ABBREVIATIONS BP before present DCP Dickerson Coquina Pit locality HI Hutchinson Island beach fill ma terial NALMA North American Land Mammal Age MIS Marine Isotope Stage(s) UF Florida Museum of Natural History, University of Florida, Gainesville and the c ollections prefix for fossils housed in the Vertebrate Paleontology Collection and for modern speci mens in the Ornithology Collection at th is institution UF/PB The collections prefix for certain fossils housed in the Vertebrate Paleontology Collection that were once in the private collection of Pierce Brodkorb

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11 Abstract of Thesis Presented to the Grad uate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science SYSTEMATIC PALEONTOLOGY OF A MIDDLE PLEISTOCENE AVIFAUNA FROM SAINT LUCIE COUNTY, FLORIDA By John Andrew Kilmer May 20 1 5 Chair: David Steadman Major: Zoology Dickerson Coquina Pit is a sand and coquina mine located in Saint Lucie County, Florida . Before open pit oper ations ceased at the site , collections were made from a vertebrate fossil bed of Middle Pleistocene ( Late Irvingtonian ) age in the Okeechobee Formation . Specimens from the coquina pit were also d iscovered in fill material used in beach replenishment on Hutchinson Island , St. Lucie County. The avifauna of the two sites contain s over 70 elements representing 12 orders, 1 6 families, and 26 species. A surprisingly w ide variety of species with affinities ranging from woodlands to the open ocean are represented in this relatively small fossil assemblage. E xtant and rece ntly extinct species found include a transition al turkey Meleagris aff. gallopavo , an albatross Phoebastria aff . albatrus , Great Cormorant Phalacrocorax carbo , a large stork Ciconia maltha , a large crane Grus sp., Great Auk Pinguinus impennis , and Carolina Parakeet Conuropsis carolinensis . The Dickers on Coquina taphonomy and paleoecology are most similar to those of other shell beds in Central and South Florida such as th e Irvingtonian Leisey Shell Pit of Hillsborough County, Florida.

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12 CHAPTER 1 INTRODUCTION Peninsular Florida is well known for i ts wealth of fossil avifaunas. Important and well studied localities range in age from the Early Miocene Thomas Farm (Steadman 2008) to Late Pleistocene sites such as the Ichetucknee River (Campbell 1980) . The bones of birds are generally small, light, and hollow; a ny locality with an extensive avian fossil record represents an exceptional pr eservational environment. However, without screenwashing fossil matrix bird fossils usually go unnoticed. While the Irvingtonian is well studied in Florida, nearly all known localities of this age are from inland sinkhole or riverine deposits or from the Gulf Coast ( Emslie 1998; Hulbert 2001) . The first major vertebrate fossil locality in Saint Lucie County, the Dickerson Coquina Pit, is unique; it is o ne of the o nly Irv ingtonian localities from represent s a wide assortment of terrestrial, aquatic, and marine avifauna . While the number of bird s pecimens recovered was small and the sediments were never screenwashed, the diversity and importance of these specimens is surprising. I describe the first occurrence of Carolina Parakeet ( Conuropsis carolinensis ), the first occurrence of Great Cormorant ( Phalacrocorax carbo ) in North America, and an early record of Northern Gannet ( Morus bassanus ) in Fl orida. Here, I present the systematic paleontology of the Dickerson Coquina Pit avifauna, provide description of the geology and biochronology of the site, and interpret the paleoecology of the area at the time of deposition. Discovery of the Dickerson Co quina Pit Specimens The Florida Museum of Natural History , University of Florida (UF) was first alerted to the fossils being found in St. Lucie County by George H. Means of the Florida

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13 Geological Survey in 2002. Dickerson Aggregates operators were extracti ng sand and coquina from two large, contiguous borrow pits near Indrio, known as Dickerson Coquina Pits 1 and 2 ( Fig ure 1 1 , 1 2) . A vertebrate fossil layer was discovered in the bottommost reaches of the pits. In addition to the large mammals typically fo und at Floridian fossil localities, a number of avian specimens w ere also recovered . The pits were excavated in the Pleistocene Anastasia and Okeechobee Formation s C oast ( Scott et al. 2001 ) . The mamma l and bird fossils at the site appear to be most similar in age to Late Irvingtonian assemblages in Florida such as those at Coleman IIA and IIIC (Ritchie 1980; Steadman 1980) or possibly the Rancholabrean assemblages at the nearby Vero Beach and Melbourne localitie s and inland sites such as the Iche tucknee River (Webb 1974, Campbell 1980, Hulbert 2001) . The depositional environment is comparable to those at other Irvingtonian vertebrate localities from marine shell beds in South Florida, such as Leisey She ll Pit (Emslie 1995 a ) and Shell Materials Pit (Emslie 1998). Dickerson Coquina Pit 1 was flooded in 2004 , shortly after collections were made there . Dickerson Coquina Pit 2 was active between 2004 and 2008. The second pit was allowed to fill with groundwat er in 2008 , and was eventually joined with the first pit in 2009. Dickerson Coquina Pits 1 and 2 are hereafter referred to as DCP. Geology of the Dickerson Coquina Pit The geology of the DCP is complex and generally composed of interbedded sand and coquin a limestone (Fig ure 1 4 ). Located in the Florida P eninsula sedimentary province (Perkins 1977), the DCP is approximately 16 km inland in an area that was subject to marine transgressions and regressions in the Pleistocene. The modern elevation of the area surrounding the site is from 7.0 to 8.8 m above sea level. Most

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14 facies at the site were formed by accreted beach dune complexes in high energy environments (Per kins 1977; Herrera et al. 2006). Lateral correlation of beds at the site to those at other locat ions is difficult at best. Several authors have offered their own nomenclature for the strata in the area (Fig ure 1 5 ) based on lithology (Sellards 1912; Scott et al. 2001), invertebrate biostratigraphic units (Petuch and Roberts 2007 ) , or eustatic sea lev els (Perkins 1977) . The layer of undifferentiated, organic rich eolian quartz sand that makes up the first 5 m of sediment at the DCP may be Holocene in origin ( Herrera et al. 2006 ; Oleinik and Comas 2012 ) . The underlying Anastasia Formation was mapped at or near the surface at the DCP by Scott et al. (2001). The formation was first described by Sellards C oast of eastern Florida, but has come to refer to a layer of Late Pleistocene sand s and limestones underlying the Atlantic Coastal Ridge from St. Johns County south to Palm Beach County and extending as much as 30 km inland in St. Lucie and Martin Counties (Scott et al. 2001). Believed to be the youngest lithified marine deposit in Flor ida (Perkins 1977), an age range for this unit is difficult to determine (Portell et al. 2003). Radiometric (U234/Th230) and amino acid racemization dating of shells and limestones from upper portions of the Anastasia place the age of the formation betwee n 136 ,500 and 8 ,000 years before present (BP) (Osmond et al. 1970; Murphy 1973; Mitterer 1974), al though age estimates for many minor facies attributed to the Anastasia have never been obtained. Major facies of the Anastasia were likely deposited during th e l ast interglacial (Marine Isotope Stages (MIS) 5e) between 130,000 100,000 BP

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15 (Oleinik and Comas 2012 ; Osmond et al. 1970 ) when sea levels rose up to 5 m above current levels (Harmon et al. 1983). Most indurated sand and shell facies of the DCP are att ributable to the Anastasia Formation (Herrera et al. 2006) . The modern echinoids collected at depths of 7 10 m and modern mollusks at all depths bespeak the relative youthfulness of the facies (Fig ure 1 4 ), although Herrera et al. (2006) cautioned that the lower facies at DCP may be older than typical Anastasia sediments. Recent electron spin resonance optical dating studies by Burdette et al. (2013) confirm these suspicions. A sample taken just above the vertebrate fossil layer at a depth of 11.5m was date d between 730,000 430,000 BP. The boundary between the Anastasia and beds below is not well defined and the lithologies of these formations are similar. The majority of these beds were deposited during times of increased sea level and marine transgression in warm interglacial periods and eroded during periods of lower sea level in cool glacials. Scott (1992 ; 2001) grouped almost all shelly sediments of latest Pliocene and Pleistocene age into the informal Okeechobee formation. Previous attempts to codify th e stratigraphic nomenclature of these sediments were based on biostratigraphic criteria without lithologic descriptions. Faunal units including the Caloosahatchee, Bermont, Fort ch and Roberts (2007) were ormation. The vertebrate fossil layer was the lowest regularly exposed (above water) facies at the DCP (Fig ure 1 4 ). While its lower boundary was not measured, Herrera et al. (2006) reported that this layer was only a few meters thick. Fossilized wood and several

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16 invertebrates were recovered from the layer. Most of the gastropods and bivalves were of freshwater origin, although terrestrial gastropods were also collected. Aquatic and marine vertebra te taxa recovered include sharks, stingrays, various freshwater and marine bony fish, freshwater turtles, and alligator. Terrestrial tortoises ( Hesperotestudo crassicutata ), turtles ( Terrapene carolina ) , and snakes also were collected. Mammalian taxa colle cted at the site are discussed below. Beds below the vertebrate fossil layer were occasionally exposed at the DCP. Petuch (2004) measured some sections at the site and believed that the lowest layers of the pit, below the vertebrate fossil bed, corresponde d to the Late Pliocene to Early and Burdette et al. (2013) dated a sample from a depth of 14m to be from between 1,370,000 610,000 BP . If these assessment s are correct, then the fossil layer is certainly older than typical fa cies of the Anastasia Formation and . It is likely that the vertebrate fossil bed represents a drowned tidal creek or marsh covered in a marine transgression that followed a glacial. Biochronology of the D ickerson Coquina Pit Hundreds of mammalian fossils were collected from the DCP and HI localities. Taxa such as Holmesina septentrionalis, Paleolama mirifica, Procyon lotor, and Panthera onca place the maximum age of the vertebrate layer firmly in the Pleis tocene rather than the Pliocene (Hulbert 2001; Bell et al. 2004). First occurrence dates of Mammuthus columbi in Florida are in the Irvingtonian North American Land Mammal Age (NALMA); its presence in the DCP material further narrows the age of the site. S everal other specimens found at the DCP and HI localities place the vertebrate fossil layer either in the Late Irvingtonian or the Rancholabrean NALMA. Canis dirus and

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17 Tapirus veroenis made their first appearance in the Late Irvingtonian but become common in the Rancholabrean (Hulbert 1995; Bell et al. 2004). Bison sp., the defining taxon of Rancholabrean faunas (Bell et al. 2004), was not recovered at the site. It is unclear how to interpret this absence. The avifauna reported below , with the exception of Phoebastria aff. albatrus, Ciconia maltha, and Grus sp., contains none of the species with western North American or Neotropical affinities seen in Early Pleistocene avifaunas (Emslie 1998). Turkey ( Meleagris aff. gallopavo ) fossils recovered at the DCP a re similar to the Rancholabrean modern M. gallopavo , but are smaller , similar in size to the transitional forms from the Late Irvingtonian Coleman IIIC locality described by Steadman (1980). A ll mollusk specimens and two of the three echinoid specimens in layers above the vertebrate fossil layer represent living species (Herrera et al. 2006 ). Taken together, the fossils suggest a Late Irvingtonian NALMA for the vertebrate fossil layer at DCP or possibly a very Early Rancholabrean age . North Hutchinson Islan d Beach Fill Specimens After two direct hits by hurricanes Frances and Jeanne in September, 2004 (Clark 2004; Franklin et al. 2006), material dredged from the DCP was used in an emergency beach replenishment project on North Hutchinson Island, St. Lucie Co unty, Florida (Fig ure s 1 1 , 1 3 ). The sand from the pit was delivered unwashed and fossils arrived mostly intact. The dark, fossiliferous replenishment material is easily differentiated from the native beach sand and subsequent beach fill. H undreds of foss ils have been collected from the beach fill material and on the beach berm . The provenance of the North Hutchinson Island (HI) specimens is clear; preservation is identical to that at the DCP, the faunal makeup is nearly the same , and no vertebrate

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18 fossil material was found in the native beach sand on the island prior to beach replenishment. Figure 1 1. Map of Florida and St. Lucie County (inset) showing the locations of Pleistocene sites discussed in the text : 1) Dickerson Coquina Pit, Late Irvingtonian ; 2) North Hutchinson Island ; 3) Vero Beach , Late Rancholabrean ; 4) Melbourne , Late Rancholabrean; 5) I chetucknee , Late Rancholabrean; 6) Shell Material Pit, Irvingtonian; 7 ) Leisey Shell Pit , Irvingtonian ; 8) Coleman IIA and IIIC, Late Irvingtonian .

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19 F igure 1 2. Dickerson Coquina Pit 2 (DCP) in 200 6 . Photo by Roger Portell . Figure 1 3. North Hutchinson Island (HI) between Jenson Beach and Walton Rocks Beach where fill from the Dickerson Coquina Pit (DCP) was deposited . Photo by Paul Roth.

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20 Fig ure 1 4. Stratigraphic section of Dickerson Coquina Pit 2 (DCP) , St. Lucie County, Florida ( a dapted from Herrera et al. 2006). Scale is in meters.

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21 Figure 1 5. A brief overview of previous stratigraphic nomenclature of the Pleistocene deposits of St. Lucie County. Time is in millions of years (Ma).

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22 CHAPTER 2 METHODS Fossils at the DCP were collected by staff of UF or donated to UF by private collectors until 2008, when the pits were allowed to flood and dredge mining began. In the years following the replenishment project on HI, beachcombers and amateur paleontologists also donated specimens to UF . Screenwashing was not performed at either locality but were washed by the tides on the beach at HI . The fossils were cemented with a coquinoid matrix and so me cavities were filled with calcite crystals which often had to be removed for identification. Avian fossils were identified by direct comparison with modern specimens from the Ornithology Collection and the V ertebrat e Paleontology C ollection at UF . All m easurements were taken with Fowler & NSK Max Cal digital calipers, rounded to the nearest 0.1 mm. Specimens were photographed with a Canon EOS 5D Mark II camera and composed with Adobe Lightroom, Helicon Focus, and Adobe Photoshop CC . The anatomical termin ology used follows Baumel (1993). Taxonomy follows that of the fourth supplement (Chesser et al. 2013 ). The fossils are housed and catalogued in the Division of Vertebrate Paleontology at UF . Most f os sils housed in the Vertebrate Paleontology Collection and modern skeletons housed in the Ornithology Collection use the prefix UF but some specimens in the Vertebrate Paleontology Collection that were once in the private collection of Pierce Brodkorb use t he prefix UF/PB .

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23 CHAPTER 3 RESULTS The composite avifauna of the DCP and HI sites contains 65 elements representing 12 orders, 16 families, and 26 species (Table 3 1). Two genera, and five of the species represented are extinct. Two families and an additi onal genus no longer occur in the region today. Most taxa represented in the D CP fauna are wetland species that are extant or only very recently extinct. Systematic Paleontology Order ANSERIFORMES Family ANATIDAE Genus Aix Boie , 1 828 Aix sponsa Linnaeus, 1758 Figure 3 1 (B, D) Material. Right proximal humerus, UF 274109; left distal tibiotarsus, UF 274114. Remarks. The humerus shows the restricted circular opening of the pneumatic fossa common in perching ducks and also the curved, robust shaft and a lack of inflection of the humeral head which are diagnostic in Aix as described by Woolfenden (1961) . The condyles of the tibiotarsus are displaced m edially and the condylus medialis extends more anteriorly than the condylus lateralis (Woolfenden 1961) . The ex act proportions and size of this specimen are only represent ed in Aix sponsa . T his spec ies has been found at several Pleistocene fossil localities in Florida.

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24 Genus Anas Linnaeus, 1758 Anas cf. A. americana Gmelin, 1789 Figure 3 1 (H) Material. Partial l eft proximal coracoid, UF 274123. Remarks. This specimen compares well with A . americana but is too fragmentary for definite identification. The coracoids of dabbling ducks show few qualitative diagnostic characters (Woolfenden 1961). A. americana is rare ly preserved in the Pleistocene record of Florida but has been found in the Ichetucknee River , Leisey Shell Pit , and a handful of other localities. Anas cf. A. platyrhynchos or A. rubripes Linnaeus, 1758 Figure 3 2 Material. Left coracoid, UF 274124; left proximal scapula, UF 274127; left distal femur, UF 274148; right shaft of humerus, UF 274156; left distal ulna, UF 274159. Remarks . The angle of the crista deltopectoralis of the humerus is diagnostic of Anas species, but the other elements presented her e are not easily distinguishable within members of this genus (Woolfenden 1961). These specimens are comparable to A . platyrhynchos or perhaps A. rubripes in size but cannot be satisfactorily assigned a specific identification. The two species are so simil ar osteologically that it is extremely difficult, if not impossible to differentiate them ( Campbell 1980). Fossils of Mallards are common ly recovered in the Pleistocene of Florida; they are nearly as numerous in the Ichetucknee River fossil record as they are in the area today.

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25 Anas discors (Linnaeus, 1758) Figure 3 1 (F) Material. Left partial carpometacarpus, UF 274131. Remarks . The carpometacarpi of Anas can be distinguished by the external surface of the trochlea carpalis and the scars of the ligament attachments. The carpometacarpus of this species is differentiated from that of A. carolinensis by the deeply excavated anterior carpal fossa (Woolfenden 1961). A. discors fossil s are commonly found in the Pleistocene localities of Florida. Genus Aythya Boie, 1822 Aythya collaris (Donovan, 1809) Figure 3 3 ( B C, E ) Material. Left proximal humerus, UF 274105; right proximal humerus, UF 274110; left proximal coracoid, UF 274145. Remarks. Unlike those of the dabbling ducks, hu meri of Aythya have a closed pn eumatic fossa and a thinner shaft (Woolfenden 1961). Aythya can also be distinguished from Clangula and Lophodytes by the well formed impressio brachialis anticus (Woolfenden 1961). T he coracoids of Aythya and its closest relatives are di stinct. The ventra l portion of the head of the coracoid is reduced and the bicipital muscle attachment is a thin groove (Woolfenden 1961). The more gracile depth of the shaft distinguishes Aythya from othe r diving ducks. The size and shape of these specimens conform most cl osely to those of A. collaris . This species is a common fossil in Pleistocene localities of Florida.

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26 Genus Bucephala Baird, 1858 Bucephala clangula (Linnaeus, 1758) Figure 3 3 ( G, I ) Material. Left distal humerus, UF 274134; right proximal humerus, UF 274 143. Remarks. The humeri of Bucephala can be separated by those of the mergansers by their relatively small and rounded crista deltopectoralis and from the eiders by its shallow, closed fossa pneumotricipitalis and curved shaft (Woolfenden 1961). These hum eri conform most closely to those of B. clangula . The anatids found in the DCP material are generally common year round or winter residents in Florida today and inhabit a variety of brackish and freshwater wetlands. B . clangula , the Common Goldeneye, is th e only uncommon member of this group. A colder water species, B. clangula is an occasional winter resident in St. Lucie County today; it typically ranges no farther south than northern Florida . B. clangula was also rarely preserved in the Pleistocene recor d of Florida. Only three other specimens have been recovered , all from the Ichetucknee River locality. Order GALLIFORMES Family PHASIANIDAE Genus Meleagris Linnaeus, 1758 Meleagris aff . gallopavo Linnaeu s, 1758 Table 3 2, Figure 3 4 , Figure 3 5 Material. Two left proximal coracoids, UF 274125, UF 274126; right proximal humerus, UF 274122; right distal ulna, UF 274161; right proximal radius, UF 274166;

PAGE 27

27 left distal femur, UF 274106; two left distal tibiotarsi, UF 274116, UF 274136; left partial shaft of a ta rsome tatarsus, UF 274169; two phalang es, UF 274152, UF 274153. Remarks. The turkey specimens from DCP are similar to modern M. gallopavo , but are slightly smaller . They are also similar to Meleagris sp. specimens recovered from the Irvingtonian Coleman II IC locality which Steadman (1980) described as intergrade between M. gallopavo and M. anza or M. leopoldi . The characters and measurements of turkeys described below and in Table 3 2 were reported in monograph (1980). The characters of tarsometa tarsus were found to be the most diagnostic by Steadman but only one of these was measurable . UF 274116 lacks the bony spur of adult male turkeys so it probably represents that of a female. The least wi d th of the shaft of the tarsometatarsus was higher tha n average modern female turkeys and only slightly larger than those of Coleman females. The characters observed in the coracoids, the width of the head of the coracoid through the scapular facet and the depth of the head , were slightly smaller than those o f male modern and smaller than Coleman turkeys but larger than those of most females. The proximal width of the humerus is similar to that of male Coleman specimens, lower than average modern males and greater than modern females. The distal width, depth o f the internal condyle, external condyle, and fibular condyle observed in the femur are most similar to those of the male Coleman specimens and smaller than average modern males . In the tibiotarsus, the distal width and depths of the internal and external condyles were within the range of modern female turkeys and certainly smaller than those of males . The phalanges of the DCP turkey are shorter and thinner than modern specimens. The ulna

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28 and radius of the DCP turkey are either fragmentary or non diagnostic . I refer to the DCP as Meleagris aff. gallopavo . The large, thick bones of the DCP t urkey were likely washed into the area from nearby woodlands and were the most commonly recovered avian specimens. M. gallopavo ean but is thought to have evolved in the Late Irvingtonian from forms such as those found at Coleman IIIC (Steadman 1980) and the DCP . Order PODICIPEDIFORMES Family PODICIPEDIDAE Genus Podilymbus Lesson, 1831 Podilymbus podiceps (Linnaeus, 1758) Figure 3 6 Material. Left distal humerus, UF 274130; right proximal tarsometatarsus, UF 2741 6 5, left shaft tarsometatarsus, UF 274168 . Remarks. P. podiceps The size and shape of the fossa medialis brachialis, c ondyl i dorsalis and ventralis, tuberculum supracondylare ventral e , and corpus humeri are diagnostic of P. podiceps in UF 274130 . The position of the foramina vascularia proximalia , cotyla e medialis and lateralis , and the impressio ligamentum collateralis l ateralis are diagnostic in UF 2741 65. Diagnostic characters in UF 274168 include the foramen vasculare distale, facies subcutanea lateralis, and cristae plantares medial i s and lateralis .

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29 Order PROCELLARIIFORMES Family DIOMEDEIDAE Genus Phoeba stria Rei chenbach, 1853 Phoebastria aff. albatrus (Pallas, 1769) Table 3 2, Fig ure 3 7 Material. Right partial coracoid, UF 274144; left proximal carpometacarpus, UF 274140. Remarks. Albatrosses, rare vagrants in the North Atlantic today, were common off the easte rn coast of Nor th America in the Pliocene and have been recovered from sites in Florida, North Carolina, and England (Ols o n and Rasmussen 2001; Ols o n and Hearty 2003). T he Pliocene taxa were closely related to the extant North Pacific albatrosses of the ge nus Phoebastria . A species similar to P. albatrus , a nearshore species threatened with extinction in the North Pacific today, is known to have survived into the Pleistocene in the N orth Atlantic on Bermuda (Ols o n and Hearty 2003). A single carpometacarpus (UF 205745) from Cochran Shell Pit is the only previous Pleistocene albatross record from Florida. The two DCP specimens most closely resemble those of the genus Phoebastria . The great albatrosses ( Diomedea ) available for study were m uch larger than these specimens. Ols o n and Rasmussen (2001) and others (Harrison and Walker 1978; Dyke et al. 2007) have compared fossil albatrosses from across the North Atlantic and have generally based their diagnosis on the size of the specimens in question in addition to t he osteological details of the tarsometatarsus, humerus, and other elements not found in the DCP assemblage. Though some of the smaller subspecies of Diomedea may approach those of the largest fossil Phoebastria specimens, there is presently no

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30 evidence of naturally occurring Diomedea in the North Atlantic, past or present ( AOU 1998; Olson and Rasmussen 2001). UF 274144 and 274140 are slightly larger than average modern P. albatrus in several respects , al though the sample size for both carpometacarpi and coracoids is small (Table 3 2) . UF 274140 is generally intermediate in size between the modern specimens of P. albatrus and a Pliocene fossil species ( P. anglica ) from Florida (Brodkorb 1960 , Olson and Rasmussen 2001 ) . The length of the step in the sternal margin of extremitas sternalis, the curvature of the processus procoracoideus, position of the foramen nervi supracoracoidei, and impressio musculi sternocoracoi dei are most similar to those of P. albatrus . I refer to the DCP specimens as Phoebastria aff. albatrus . Order CICONIIFORMES Family CICONIIDAE Genus Ciconia Brisson, 1760 Ciconia maltha Miller, 1910 Figure 3 8 Material. Partial right distal tarsometatarsus, UF 274120; partial right distal tarsometatarsus, UF 274121; left prox imal carpometacarpus , UF 274138 . Remarks. These specimens are similar to modern C . ciconia specimens but are much larger, approaching Jabiru mycteria specimens in size. The specimens are also much larger than the fossil stork Mycteria wetmorei and the tarsometatarsus lacks t he distal portion of that is characteristic of members of that genus ( Suárez and Olson 2003). They compare well with C. maltha specimens in the V ertebrate

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31 P aleontology C ollection (UF /PB 001234 , UF 8 0 433 , and UF 15708 ) from the Ichetucknee River and Leisey Shell Pit , Florida . While only the distal ends of t he tarsometatarsi were preserved, UF 274121 possess es an anteroposteriorly shaft and UF 274120 possesses a less posteriorly situated outer trochlea as Suárez and Olson (2003) described . The foramen vascular e distale is rounded posteriorly and the m uscle scars on the dorsal face are extremely similar to those in other C. matha specimens. The sulcus tendineus and facies articular is digitalis major of the carpometacarpus are also characteristic. C. maltha is common in the Ichetucknee River, Leisey Shell Pit, and other Pleistocene localities , likely due to its large size and aquatic affinities. Order SULIFORMES Family SULIDAE Genus Morus Vieillot, 1816 Morus bassanus ( Linnaeus , 17 58 ) Figure 3 9 (B) Material. Right distal partial dentary, UF 273107. Remarks. The fenestrae , muscle scars , and crista tomialis on this specimen correlate with those of modern M. bassanus. T his is the first recorded fossil specimen of this species in Flo rida. Specimens of a similar age are known from Virginia (Spencer and Campbell 1987) and possibly England (Seward et al. 2006). A Holocene specimen was found in the Summer Haven midden in Florida (Brodkorb 1960).

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32 Family PHALACROCORACIDAE Genus Phalacroc orax Brisson, 1780 Phalacrocorax cf. P. auritus (Lesson, 1831) Figure 3 10 (B, F) Material. Left proximal femur, UF 274117; left proximal tarsometatarsus, UF 274149. Remarks. The se specimens resemble P . auritus in size but are too fragmentary for definite identification. P. auritus Phalacrocorax carbo ( Linnaeus , 17 58 ) Figure 3 10 (C) Material. Left femur, UF 274118. Remarks. The North American subspecies of Great Cormorant, P . carbo carbo , is mostly mar ine . It is common in and around its breeding colonies in the Canadian Maritime Provi nces and northern New England. Non breeding individuals may range far to the south, even to Florida (Stevenson and Anderson, 1994). colonial distribution is not well known , although specimens have been recovered from the Holocene in Florida at Lewis Camp Mound (Kratt 2005) 2005) ; many populations were locally endangered or extirpated by hunters near the turn of the 20 th century , an d P. carbo is easily confused in the field with P. auritus (Erskine 1972; Stevenson and Anderson 1994) but is larger in comparison . Records of P. carbo in Florida increased in the 1970s coinciding with the gradual recovery of the species at th is time . The presence of P. carbo in the DCP material is suggestive of a more abundant population of these birds in the Pleistocene. This specimen is the first

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33 described fossil record of P. carbo in North America and is diagnosable by its large size and the shape and p roportions of the trochlea fibularis, linea intermuscularis caudalis and cranialis, and the facies articularis acetabularis . Phalacrocorax cf. P. carbo ( Linnaeus , 17 58 ) Figure 3 10 (H, J) Material. Left distal ulna, UF 274162; left distal tibiotarsus, UF 274115. Remarks. These specimens are slightly smaller than modern P. carbo specimens but much larger than in P. auritus . Family ANHI N GIDAE Genus Anhinga Brisson 1760 Anhinga anhinga ( Linnaeus , 17 66 ) Figure 3 9 (D) Material. Partial synsacrum, UF 274146. Remarks. This specimen exhibits a sulcus ventralis synsacri and recessus caudalis fossae identical to that of A. anhinga . The size and shape of the fused synsacro lumbar vertebrae and the recesses between the synsacro caudal vertebrae are also characteris tic. The only other fossil Anhinga described from the Pleistocene of Florida, A. beckeri , was much larger than A. anhinga (Emslie 1998). Miocene species A. subvolans and A. grandis have also been found in Florida. Anhinga fossils are somewhat common in Fl orida but have not been found at in any other area in North America.

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34 Order PELECANIFORMES Family ARDEIDAE Genus Ardea Linnaeus, 1758 Ardea herodias Linnaeus, 1758 Figure 3 11 (B) Material. Left distal tibiotarsus, UF 274108. Remarks. This specimen is mos t similar to modern A. herodias in size and proportion. It possesses a well defined fibular ridge and a narrow pons supratendineus as compared with the similar sized A. alba . The depth of the canalis extensorius is also most similar to A. herodias. This sp ecies is common in the Pleistocene record of Florida at riverine sites such as the Ichetucknee River. It is found in a wide variety of aquatic habitats ranging from beaches to small freshwater streams today. Genus Nyctanassa Stejneger, 18 8 7 Nyctanassa vio lacea (Linnaeus, 1758) Figure 3 11 (D) Material. Right partial quadrate, UF 274113. Remarks. Th e cranial osteology of birds is diagnostic. The length and shape of the processus orbitalis and the size and shape of the capitulum squamosum and oticum of the p rocessus oticum, the shape of the condyli caudalis, pterygoideus, and lateralis on the processus mandibularis, and cotyla quadratojugalis are identical to modern N. violacea . This species is rarely recovered in the Pleistocene of Florida.

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35 Family THRESKIOR NITHIDAE Genus Eudocimus Wagler, 1 8 32 Eudocimus albus (Linnaeus, 1758) Figure 3 11 (F) Material. Left proximal coracoid, UF 274133. Remarks. The shape of the cotyla scapularis and facies articularis humeralis, and curvature of the head of the coracoid ide ntify this specimen as a member of the genus Eudocimus . Another Pleistocene species, E. leiseyi was much smaller and gracile than the DCP and modern specimens (Emslie 1995 a ) . Eudocimus fossils are rarely found in the Pleistocene of Florida. Order ACCIPIT RIFORMES Family ACCIPITRIDAE Genus Haliaeetus Savigny, 1809 Haliaeetus leucocephalus (Linnaeus, 1766) Figure 3 12 (B, D, F) Material. Right proximal scapula, UF 274164; phalanx ungualis , UF 2741 37 ; phalanx, UF 294468 . Remarks. T he positions and proportio ns of the acromion and tuberculum coracoideum confirm the identity of the scapula as that of a large eagle ( Haliaeetus or Aquila ). T he size of the scapula and ungal phalanx is closest to that of a male H. leucocephalus while UF 294468 is likely a female. B ald Eagles are somewhat common in the Rancholabrean of Florida but ha ve not been recorded earlier until now .

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36 Genus Buteo Lacépède, 1799 Buteo lineatus ( Gmelin , 17 88 ) Figure 3 12 (H, J) Material. Left distal humerus , UF 274167; right distal tarsometatarsus , UF 274119. Remarks. The condyli ventralis and dorsalis, fossa muscluli brachialis, and the processus supracondylaris dorsalis of the humerus are most similar to those of Buteo and the size of these elements is most similar to those of B. lineatus. B. li neatus is commonly recorded from the Pleistocene of Florida. Order GRUIFORMES Family RALLIDAE Genus Rallus Linnaeus, 1758 Rallus elegan s Audubon , 1 834 Figure 3 13 (B) Material. Right humerus, UF 274142. Remarks. The shape and proportions of the corpus h umerus, crista deltopectoralis, processus flexorius, and margo caudalis on this humerus are most similar to those of R. elegans , though it is slightly larger than modern specimens. R. elegans is common in the Pleistocene record of Florida.

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37 Genus Fulica Linnaeus, 1758 Fulica americana Gmelin, 1789 Figure 3 13 ( D, F ) Material. Right distal ulna, UF 274163; right distal tibiotarsus, UF 274128. Remarks. The tibiotarsus exhibits a large condylus lateralis and pons supratendineus similar to that of F. american a . The condyli dorsalis and ventralis and tuberculum carpale of the ulna are nearly identical to this species. F. americana is a very commonly preserved fossil at Pleistocene sites in Florida and is quite common today . Family GRUIDAE Genus Grus Brisson , 1 760 Grus sp. Table 3 3, Figure 3 14 Material. Right distal tibiotarsus, UF 274132; phalanx, UF 274157. Remarks. The right distal tibiotarsus (UF 274132) was compared to modern and select fossil Grus species from North America (Table 3 3 ). The specimen is larger than in either extant North American species ( G. ameri c ana, G. canadensis ) , but compares well to an Early Pleistocene specimen ( a distal tibiotarsus , UF 91223 from the Irvingtonian Shell Material s Pit. Emslie (1995) described this large crane of th e Shell Materials and Leisey Shell Pits of Hillsborough County as Grus sp. and believed this species was closely related to or possibly even conspecific to Grus cubanensis . Thought to be flightless by its discoverers, G. cubanensis lived in the La te Pleist ocene of Cuba (Fischer 1968; Fischer and Stephan 1971). If G. cubanensis was flightless , the

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38 Floridian specimens would likely be distinct enough to warrant identification as a separate species. Because G. cubanensis material was not available for study and the Floridian record is so poor , I refrain from assigning the DCP specimen a specific epithet. UF 274157 was compared with specimens of extant Ardea , Ciconia , Jabiru , and Grus. Th is phalanx was most similar to those of Grus americana , but was much larger . The width of the shaft, the angular nature of the tuberculum, and the shape of the trochlea was closest to G. americana . The specimen was not similar to Ciconia maltha specimens in the V ertebrate P aleontology C ollection from Leisey Shell Pit, Florida (UF 80435, 85009) in shape or size. I refer to both UF 274157 and UF 274132 as Grus sp. Order CHARADRIIFORMES Family ALCIDAE Genus Pinguinus Bonnaterre, 1791 Pinguinus impennis ( Linnaeus, 1758) Fig ure s 3 15, Figure 3 16 Material. Right proximal scapula, UF 274102; right proximal humerus , UF 274101; left proximal ulna, UF 274104; left ulna, UF 274103; right ulna, UF 266756; right pro ximal tibiotarsus, UF 274155; right proximal radius, UF 274139. Remarks. These specimens clearly correspond to modern specimens of P. impennis such as UF 33837 (composite). Two of the fossils (UF 274103, 274104) exhibit a porous bone surface and represent juvenile individuals. No modern radii of P . impennis were available for study; nevertheless , UF 274139 is more massive and disti nctive than the radii of other modern alcids such as Alca torda and Uria lomvia .

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39 The number of Great Auk elements found at the DCP and HI localities is surprising ; it was the second most common avian tax on identified. Several specimens of P. impennis have been recovered from archeological sites in Florida: Cotton midden , Volusia County (Hay, 1902); the Castle Windy site , Volusia County (Weigel 1958) ; Green Mound mid den, Volusia County (Hamon 1959); Summer Haven midd en, St. Johns County (Bro dkorb 1960); and Boca Weir, Palm Beach County (Fradk in 1980). A fossil specimen has also been reported from the Middle Pleistocene of Bermuda (Olson 2003). Great Auks were driven to extinction at their last breeding colonies on Funk Island , Canada and Eldey Island , Icelan d by the mid nineteenth century ( Meldgaard 1988 ) . The presence of both adult and juvenile Great Auk fossils at the DCP may indicate the presence of more southerly breeding colonies in the Middle Pleistocene. Y oung P. impennis were at least moderately prec ocial and may have left their nesting areas shortly after hatching to join their parents at sea (Gaskell 2004; Houston et al . 2010). It may have taken juvenile birds a number of months to complete skeletal development ( Meldgaard 1988 ) . T he presence immatur e P. impennis specimens in the Holocene middens of Herriko Barra, Basque country, Spain suggest that the young birds may have traveled hundreds of kilometers from their breeding areas before their bon es were fully ossified (Stewart 2002). Another explanati on is that these specimens represent a die off event caused by a large storm or hurricane that forced these individuals southward to Florida. In 2012, the unusual weather patterns brought about by Hurricane Sandy caused a similar event in another alcid, th e Razorbill ( Alca torda ) . Hundreds of individuals were blown out of

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40 their northern range and many washed up, emaciated, on the shores of Florida that year . Order STRIGIFORMES Family STRIGIDAE Genus Bubo Duméril, 1805 Bubo virginianus (Gmelin, 178 8) Figu re 3 17 (B, D) Material. Left distal humerus, UF 274111; right distal tarsometatarsus, UF 2741 4 7. Remarks. The condyles of the tarsometatarsus, UF 2741 4 7, are somewhat eroded but are proportional to B. virginianus as is the shaft and the foramen vasculare distale. The position and proportions of the crista supracondylus medialis, condylus lateralis, and tuberculum m usculi gastrocnemialis medialis in UF 274111 approach those of this species, also. The size of UF 274111 is most similar to that of a male B. vi rginianus while UF 2741 4 7 is closer to that of a female. These owls are somewhat common in Floridian cave deposits but rare in other Pleistocene localities. Genus Strix Linnaeus, 1758 Strix varia Barton, 1799 Figure 3 17 (F) Material. Right proximal femur , UF 274135. Remarks. The proportions and shape of the crista tr o chanteris, fovea ligamentosus capitis, facies articularis acetabularis, and linea intermuscularis caudalis

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41 are identical to those of S. varia . This owl is common in the riverine and cave depo sits of Florida . Order PSITTACIFORMES Family PSITTACIDAE Genus Conuropsis Salvadori, 1891 Conuropsis carolinensis (Linnaeus, 1758) Fig ure 3 18 Material. Right proximal humerus, UF 274112. Remarks. The sulcus transversus, impressio coracobr achialis, and the di stinctive shape and size of this humerus confirm its identity as a parakeet. While no osteological material of the extinct C . carolinensis was available, the specimen is identical in shape and proportion to drawings by Shufeldt (1886) a s shown in Fig ure 3 18 . F ew prehistoric ly parakeet exist . Parmalee (1958, 1967) identified the remains of several C. carolinensis elements from archeological sites in central Illinois . T hree elements were also found at a burial site in southwestern On tario (Prevec 1984). A fossil parrot humerus from the Miocene of Nebraska was described as a member of the genus Conuropsis by Wetmore (1926), al though the validity of this generic placement is questionable (Ols o n 198 5 ). The HI specimen of C. carolinensis is the earliest record of the species and the first pre Holocene fossil ever described. Historically , C . carolinensis lived mostly in the southern United States along the Gulf of Mexico from Texas to Florida to the Carolinas, as well as and north and west along major waterways. L ast seen in St. Lucie County

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42 sometime in the 1880s (McKinley 1 985), t he species became extinct in the 1920s or 1930s ( Snyder and Russe l l 2002 ).

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43 Table 3 1 . Birds identified from the Dickerson Coquina Pit (DCP) and from beach fill derived from DCP on North Hutchinson Island (HI), St. Lucie County, Florida. Current status in Florida categories: E, extinct/extirpated; V, vagrant; M I , migrant; R, resident; W, widespread. Generalized habitat categories: A, aquatic (f resh or brackish water) ; MA, marine; T, terrestrial (non aquatic). FAMILY Species DCP HI Current status Habitat ANATIDAE Aix sponsa (Wood Duck) 2 R, W A cf. Anas americana (probable American Wigeon) 1 M I , W A cf. Anas platyrhynchos or A. rubripes ( probable Mallard or Black Duck ) 4 1 R, W A Anas discors (Blue winged Teal) 1 M I , W A Aythya collaris (Ring necked Duck) 1 2 M I , W A Bucephala clangula (Common Goldeneye) 2 M I A , MA PHASIANIDAE Meleagris aff. gallopavo ( t ur key) 3 8 R, W T PODICIPEDIDAE Podilymbus podiceps (Pied billed Grebe) 3 R, W A DIOMEDEIDAE Phoebastria aff. albatrus (albatross) 1 1 E MA CICONIIDAE Ciconia maltha (large stork) 1 3 E T, A SULIDAE Morus bassanus (North ern Gannet) 1 M I , W MA PHALACROCORACIDAE cf. Phalacrocorax auritus (probable Double crested Cormorant) 1 1 R, W A Phalacrocorax carbo (Great Cormorant) 1 V A, M A cf. Phalacrocorax carbo (probable Great Cormorant) 2 V A, M A ANHINGI DAE Anhinga anhinga (Anhinga) 1 R, W A ARDEIDAE Ardea herodias (Great Blue Heron) 1 R, W A Nyctanassa violacea (Yellow crowned Night Heron) 1 R, W A

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44 Table 3 1. Continued. FAMILY Species DCP HI Current status Habitat THRESKIORNIT HIDAE Eudocimus albus (White Ibis) 1 R, W A ACCIPITRIDAE Haliaeetus leucocephalus (Bald Eagle) 3 R, W A Buteo lineatus (Red shouldered Hawk) 2 R, W T RALLIDAE Rallus elegans (King Rail) 1 R, W A Fulica americana (American Coot) 1 1 R, W A GRUIDAE Grus sp. (large crane) 1 1 E A ALCIDAE Pinguinus impennis (Great Auk) 5 2 E MA STRIGIDAE Bubo virginianus (Great Horned Owl) 1 1 R, W T Strix varia (Barred Owl) 1 R, W T PSITTACIDAE Conurop sis carolinensis (Carolina Parakeet) 1 E T

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45 Table 3 2 . Measurements (mm ± SD) of various skeletal elements in select Irvingtonian and modern Meleagris . The number of specimens examined is in parentheses. Asterisks denote slightly da maged specimens. Characters are from Steadman (1980). Measurements of turkeys from Coleman IIIC and modern turkeys were reported by Steadman. Coleman male Meleagris sp. fossils modern male M . gallopavo Dickerson Coquina Pit M . aff. gallopavo fossils C oleman female Meleagris sp. fossils modern female M . gallopavo coracoid head through scapular facet 37.80 37.0 38.7 (7) 40.00 ± 1.74 35.3 43.1 (39) 34.46* 33.8* 35.12 (2) 31.40 31.0 31.8 (2) 31.08 ± 1.52 28.0 34.2 (32) depth of head 14.84 ± 0.49 14.2 15.8 (8) 15.60 ± 0.89 13.3 17.2 (40) 12.94 12.23 13.65 (2) 11.85 11.8 11.9 (2) 11.64 ± 0.84 10.0 13.8 (32) humerus proximal width 36.83 33.6 37.8 (7) 40.19±1.52 36.0 43.2 (39) 37.2 (1) 31.33* 30.1* 32.0 (3) 31.23 ± 1.36 28.2 34.1 (31) tarsometa tarsus least width of shaft 9.60 ± 0.48 9.1 10.5 (11) 9.26 ± 0.61 8.0 10.8 (45) 7.55 (1) 8.0 (1) 7.21 ± 0.44 6.3 8.2 (34) tibiotarsus distal width 21.09 ± 0.47 20.3 21.6 (10) 21.66 ± 1.03 19.0 21.5 (39) 18.86 (1) 17.88 17.0 18.8 (5) 17.12 ± 0.86 15.7 19.3 (28) depth of internal condyle 21.08 20.6 21.8 (6) 21.38 ± 1.10 18.1 23.1 (38) 15.65 (1) 17.8* 17.2 18.7* (4) 17.00 ± 0.86 15.4 19.6 (28) depth of external condyle 18.95 ± 0.25 18.6 19.5 (12) 19.29 ± 0.74 17.0 20.4 (38) 17.24 (1) 16.05 15.6 16.5 ( 2) 15.43 ± 0.66 14.2 17.3 (28) femur distal width 28.70 28.2 29.4 (5) 29.38 ± 1.29 26.0 32.0 (40) 28.63 (1) 23.7 (1) 22.36 ± 1.23 20.0 25.0 (30) depth of internal condyle 23.56* 22.3* 24.7* (5) 24.45 ± 1.00 21.6 26.0 (38) 23.62 (1) 21.2* (1) 18. 92 ± 0.80 17.8 21.4 (30) depth of external condyle 23.50 22.4 24.8 (7) 23.93 ± 0.98 21.4 25.5 (38) 23.42 (1) 20.10 19.7 20.5 (2) 18.26 ± 0.83 16.9 20.6 (30) depth of fibular condyle 20.47 19.6 21.7 (6) 20.87 ± 0.90 19.0 22.5 (39) 19.55 (1) 16.9* (1) 15.90 ± 0.86 14.8 18.6 (30)

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46 Table 3 3 . M easurements (mm ± SD) of the carpometacarpus in Phoebastria . Species Proximal breadth Proximal depth Distance between bridge of metacarpal process and pollical facet Minimum shaft depth P. aff . albatrus, Di ckerson Coquina Pit (UF 274140 ) 22.8 9.6 17.5 6.2 P. anglica, Bone Valley (UF 65765) 24.0 10.3 18.0 7.0 P. albatrus (n = 4) 20.7 ± 0.5 9.1 ± 0.3 15.2 ± 0. 2 6.0 ± 0. 5 Table 3 4 . Measurements (mm ± SD) of the distal tibiotarsus in Grus . Measure ments of G. cubanensis were reported by Fischer and Stephan (1971). Species Sex Distal width Medial condyle depth Dickerson Coquina Pit fossil (UF 274132) 25.6 26.1 Shell Materials fossil (UF 91223) 26.5 27.2 G. cub an ensis fossils 27.0 32.0 25.6 31.0 G. americana (n = 8) M 22.0 ± 0.5 20.7 ± 0.3 G. americana (n = 8) F 21.1 ± 0.7 19.7 ± 0.8 G. canadensis (n = 8) M 19. 2 ± 0.8 18.3 ± 0.9 G. canadensis (n = 8) F 18.6 ± 1.0 17.9 ± 0.8

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47 Figure 3 1. Pectoral and hindlimb elements of dab bling ducks (Anatidae) from Dickerson Coquina Pit, Florida compared with modern specimens. A D: The right humeri in palmar aspect A B) and left distal tibiotarsi in anterior aspect C D) of Aix sponsa . A, C: modern A. sponsa UF 47798. B: fossil A. sponsa , U F 274109. D: fossil A. sponsa , UF 274114. E F: The left carpometacarpi of Anas discors in internal aspect. E : modern A. discors , UF 47143. F : fossil A. discors , UF 274131 . G H: The left coracoids of Anas americana in dorsal aspect . G : modern A. americana , UF 47091. H : fossil A. cf. americana , UF 274123. Scale bar = 20 mm.

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48 Figure 3 2. Pectoral and hindlimb elements of Mallards ( Anas platyrhynchos ) from Dickerson Coquina Pit, Florida compared with modern specimens. Right humeri in palmar aspect A B), le ft coracoids in dorsal aspect C D), left scapulae in dorsal aspect E F), left ulnae in dorsal aspect G H), and left femurs in anterior aspect I J). A, C, E, G, I: modern A. platyrhynchos , UF 47807. B: fossil A. cf . platyrhynchos , UF 274124 . D: fossil A. cf . platyrhynchos , UF 274156. F: fossil A. cf . platyrhynchos , UF 274127. H: fossil A. cf . platyrhynchos , UF274154. J: fossil A. cf . platyrhynchos , UF274148. Scale bar = 20 mm.

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49 Figure 3 3. Wing elements of diving ducks (Aythyini and Mergini) from Dickerso n Coquina Pit, Florida compared with modern specimens. A C: The humeri of Aythya collaris in aconal aspect. A: modern A. collaris , UF 43819. B: fossil A. collaris , UF 274105. C: fossil A. collaris , UF 274110. D E: The left coracoids of A. collaris in dorsa l aspect. D: modern A. collaris , UF 43819. E: fossil A. collaris , UF 274145. F I : The humeri of Bucephala clangula in aconal F G) and palmar H I ) aspects. F , H : modern B. clangula , UF 46119. G: fossil B. clangula , UF 274134. I : fossil B. clangula , UF 27414 3. Scale bar = 20 mm.

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50 Figure 3 4. Pectoral and hindlimb elements of turkeys ( Meleagris ) from Dickerson Coquina Pit, Florida compared with modern specimens. Proximal coracoids in dorsal aspect A C), proximal humeri in aconal aspect D E), distal femora in posterior aspect F G), and distal ulnae in internal aspect H I). A, D, F, H: modern M. gallopavo , UF 42955. B: M. aff. gallopavo fossil, UF 274125. C: M. aff. gallopavo fossil, UF 274126. E: M. aff. gallopavo fossil, UF 274122. G: M. aff. gallopavo fossil, UF274106. I: M. aff. gallopavo fossil, UF274161. Scale bar = 20 mm.

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51 Figure 3 5. Additional p ectoral and hindlimb elements of turkeys ( Meleagris ) from Dickerson Coquina Pit, Florida compared with modern specimens. Proximal radii in palmar aspect A B), distal tibiotarsi in anterior aspect C E), tarsometatarsi in internal a spect F G) , and phalanges in dorsal aspect H K). A, C, F , H , J : modern M. gallopavo , UF 42955. B: M. aff. gallopavo fossil, UF 274166. D: M. aff. gallopavo fossil, UF 274116. E: M. aff. gallopavo fossil UF 274136. G: M. aff. gallopavo fossil, UF 274169. I: M. aff. gallopavo fossil, UF 274152. K: M. aff. gallopavo fossil UF 274153. Scale bar = 20 mm.

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52 Figure 3 6. The humeri in palmar A B) aspect and tarsometatarsi in anterior C E) aspect of Pied billed Grebes ( Podilymbus podiceps ) from Dickerson Coquina Pit, Florida compared with modern specimens. A: modern P. podiceps , U F 22422. B: fossil P. podiceps , UF 274130. C: modern P. podiceps , UF 22422. D: fossil P. podiceps , UF 274165. E: fossil P. podiceps , UF 274168. Scale bar = 20 mm.

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53 Figure 3 7. The right coracoids in dorsal A B) and ventral C D) aspects and the carpom etacarpi in internal E G) and external H J) aspects of Phoebastria albatrosses. A, C, E, H: modern P. albatrus UF 39050 (composite). B, D: P. aff . albatrus , fossil from Dickerson Coquina Pit, Florida, UF 274144. F, I: P. aff . albatrus , fossil from Dickerso n Coquina Pit, Florida, UF 274140. G, J: P. anglica, fossil from Bone Valley, Florida, UF 65765. Scale bar = 20 mm.

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54 Figure 3 8. The proximal carpometacarpi in dorsal A B) aspect and distal tarsometatarsi in anterior C F) and posterior G J) aspects of fossil storks (Ciconiidae). A: Ciconia maltha fossil from Ichetucknee River, Florida, UF / PB 001234. B: C. maltha fossil from Dickerson Coquina Pit, Florida, UF 274138 . C, G: C. maltha fossil from Leisey Shell Pit, Florida, UF 80433 . D, H: C. maltha foss il from Dickerson Coquina Pit, Florida, UF 274121. E, I: C. maltha fossil from Dickerson Coquina Pit, Florida, UF 274120. F, J: Mycteria wetmorei from Ichetucknee River, Florida, UF 15594. Scale bar = 20 mm.

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55 Figure 3 9. Fossil Northern Gannet ( Morus ba ssanus ) and Anhinga ( Anhinga anhinga ) from Dickerson Coquina Pit, Florida compared with modern specimens. A B: The right distal dentaries of M. bassanus in dorsal aspect. A: modern M. bassanus , UF 42081. B: fossil M. bassanus , UF 274107. C D: The synsacra of A. anhinga in dorsal aspect . C: modern A. anhinga , UF 40342. D: fossil A. anhinga , UF 274146. Scale bar = 20 mm.

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56 Figure 3 10. Wing and hindlimb elements of cormorants ( Phalacrocorax ) from Dickerson Coquina Pit, Florida compared with modern specime ns. Femora in anterior aspect A D), proximal tarsometatarsi in anterior aspect E F), distal ulnae in dorsal aspect G H), and proximal tibiotarsi in anterior aspect I J). A, E : modern P. auritus , UF 22806. D, G, I: modern P. carbo , UF 22784. B: fossil P. cf P. auritus , UF 274117. C: fossil P. carbo , UF 274118. F: fossil P. cf P. auritus , UF 274149. H: fossil P. cf P. carbo , UF 274162. J: fossil P. cf P. carbo , UF 274115. Scale bar = 20 mm.

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57 Figure 3 11. Fos sils of herons (Ardeidae) and White I bises ( Eud ocimus albus ) from Dickerson Coquina Pit, Florida compared with modern specimens . A B: The left distal tibiotarsi of Ardea herodias in anterior aspect. A: modern A. herodias , UF 23074. B: fossil A. herodias , UF 274108. C D: The right quadrates of Nycta nassa violacea in posterior aspect. C: modern N . violacea , UF 22983. D : fossil N. violacea , UF 274113. E F: The left proximal coracoids of Eudocimus in dorsal aspect. E: modern E. albus , UF 23123. F: fossil E. albus , UF 274133. Scale bar = 20 mm.

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58 Figur e 3 12. Fossils of hawks and eagles (Accipitridae) from Dickerson Coquina Pit, Florida compared with modern specimens. A E: The right proximal scapulae in ventral A B) aspect, phalanges in dorsal C D) aspect, and phalanxes in lateral E F) aspect of Haliae etus leucocephalus . A, E: modern H. leucocephalus , UF 23837. B: fossil H. leucocephalus , UF 274164. C: modern H. leucocephalus , UF 23842. D: fossil H. leucocephalus , UF 274137. F: fossil H. leucocephalus, UF 274468. G J: The left distal humeri in palmar aspect G H) and right distal ta rsometatarsus in anterior aspect I J) of Buteo lineatus . G, I: modern B. lineatus , UF 23893. H: fossil B. lineatus , UF 274167. J: fossil B. lineatus , UF 274119. Scale bar = 20 mm.

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59 Figure 3 13. The right humeri of rails ( Rallus elegans ) in aconal A B ) aspect and right distal ulna in ventral C D) aspect and right distal tibiotarsus in anterior E F) aspect of coots ( Fulica americana ) from Dickerson Coquina Pit, Florida compared with modern specimens. A: modern R. elegans , UF 24313. B: fossil R. elegans , UF 274164. C, E: modern F. americana , UF 46733. D: fossil F. americana , UF 274163. F: fossil F. americana , UF 274128. Scale bar = 20 mm.

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60 Figure 3 14. The phalanxes in dorsal A B) aspect and distal tibiotarsi in anterior C E) and distal F H) aspects o f cranes ( Grus ). A, C, F: modern G. americana , UF 40254. B: fossil Grus sp. from Dickerson Coquina Pit, Florida, UF 274157. D, G: Grus sp. fossil from Dickerson Coquina Pit, Florida, UF 274132. E, H: Grus sp. fossil from Shell Materials Pit, Florida, UF 91 223. Scale bar = 20 mm.

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61 Figure 3 15. Humeri, scapulae, and tibiotarsi of Great Auks ( Pinguinus impennis ) . Proximal humeri in aconal A B) and palmar C D) aspects, scapulae in dorsal E F) and ventral G H) aspects, proximal tibiotarsi in anterior I J) a nd posterior K L) aspects. A, C, E, G, I, K: modern P. impennis UF 33837. B, D: P. impennis , fossil from Dickerson Coquina Pit, Florida, UF 274101. F, H: P. impennis , fossil from Dickerson Coquina Pit, Florida, UF 274102. J, L: P. impennis , fossil from Dic kerson Coquina Pit, Florida, UF 274155. Scale bar = 20 mm.

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62 Figure 3 16. Ulnae of Great Auks ( Pinguinus impennis ) in dorsal A C) and ventral D F) aspects . A, D: modern P. impennis UF 33837. B, E: P. impennis, fossil from Dickerson Coquina Pit, Florida, U F 274103. C, F: P. impennis , fossil from Dickerson Coquina Pit, Florida, UF 274156. Scale bar = 20 mm. Figure 3 17. The humeri, tarsometatarsi, and femora of owls (Strigidae) from Dickerson Coquina Pit, Florida compared with modern specimens. Left dis tal humeri in aconal A B) aspect and right distal tarsometatarsi in anterior C D) aspect of Bubo virginianus . A: modern B. virginianus , UF 40631. B: fossil B. virginianus , UF 274111. C: modern B. virginianus , 40642 D: fossil B. virginianus , UF 274147. E F: Right proximal femora of Strix varia in posterior aspect. E: modern S. varia , UF 40369. F: fossil S. varia , UF 274135. Scale bar = 20 mm.

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63 Figure 3 18. The humeri of parakeets in aconal A D) and palmar E H) aspects. A, E: modern Nandayus nenday , U F 26047. B, F: modern Aratinga solstitialis , UF 25934. C, G: Conuropsis carolinensis, fossil from Dickerson Coquina Pit, Florida, UF 274112. D, H: modern C. carolinensis, adapted from R. W. Shufelt (1886). Scale bar = 20 mm.

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64 CHAPTER 4 DISCUSSION The DCP l ocality was likely located in an open embayment with several streams nearby during fossil deposition , a situation akin to that of other marine shell beds in South Florida ( Pratt and Hulbert 1995; Emslie 1995 a ; Emslie 1998). Most avian fossils recovered fro m DCP and HI represent typical lagoonal, estuarine, and freshwater wetland birds found in the area today. Wading birds ( e.g., Ardea herodias, Nyctanassa violacea, Eudocimus albus ) that fed on a variety of invertebrates and small vertebrates are common in shallow water and muddy areas. N. violacea forage s in areas with high concentrations of crustaceans like tida l marshes, beaches, lagoons, creeks , and rivers; it relies more on coastal areas in the winter (Watts 2011). E. albus also moves to coastal areas in the non breeding season and frequents f reshwater wetlands, salt marshes, tidal creeks, and lagoons. This species typically feeds in shallow pools with depths less than 20 cm (Heath et al. 2009). Rallus elegans typically inhabit shallow freshwater and br ackish marshes with abundant plant cover (Poole et al. 2005). While modern North American storks ( Mycteria americana ) feed almost exclusively in aquatic habitats, C. maltha may have foraged in open uplands in addition to aquatic habitats (Suárez and Olson 2003). Double crested Cormorants ( Phalacrocorax auritus ) forage in shallow waters (<8 m deep) within sight of the shoreline (Hatch and Weseloh 1999 ), while Great Cormorants ( P . carbo ) forage deeper (<20 m deep) , sometimes up to 30 km from shore (Hatch et a l. 2000) . P. auritus frequent coastal areas in the non breeding season. A . anhinga are found in shallow, sheltered waters with nearby drying and sunning perches

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65 such as bays, lagoons, and tidal and freshwater streams (Frederick and Siegel Caussey 2000). D abbling ducks ( Anas spp.) and coots ( Fulica americana ) prefer shallow bodies of water with emergent vegetation during most of the year. Coots utilize coastal bays, lagoons, and other brackish habitats during winter (Brisbin et al. 2002). T he diving ducks represented at the DCP ( Aythya collaris, Bucephala clangula ) are typically found in deeper waters (usually <5 m deep) where they feed on underwater vegetation or small mollusks and crustaceans (Eadie et al. 1995; Roy et al. 2012) . In its breeding range B. clangula resides in freshwater wetlands, lakes and rivers but is concentrated in more saline habitats in the non breeding season such as shallow bays and estuaries (Eadie et al. 1995). M arine taxa , such as Phoebastria aff . albatrus , Morus bassanus, and Pin guinus impennis, could have been pushed into the DCP locality in a large storm or may represent vagrant individuals which became sick and/or malnourished . These taxa along with several species of marine fish found in the vertebrate fossil layer were likely deposited in higher sections o f the vertebrate fossil layer which represent the start of a marine transgression. The remains of species such as Aix sponsa, M eleagris aff. gallopavo , Buteo lineatus , Bubo virginianus, Strix varia, and Conuropsis carolinensi s may have been transported to the area of deposition from n earby woodlands and riparian areas . Some mammalian taxa at the site , such as Paleolama mirifica and Equus sp . , indicate that open habitats such as prairies or grasslands were nearby as does the pr esence of Grus sp .

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66 The presence of several specimens of northern, nearshore and marine t axa such as Bucephala clangula, Phalacrocorax carbo, and Pinguinus impennis at the DCP may indicate that sea surface temperatures were cooler than those at present . Cyc lical cold and warm cli matic variations are well known from the last glacial cycle (Heinrich, 1988; Bond et al. 1992 , Dansgaard et al. 1993, NGRIP members 2004 ). That being said, it must be made clear that the avian species present at a fossil site such a s the DCP may not be indicative of the specie s regularly present in the area and that i ndividuals that die in an area did not necessarily live there for long. Fossil assemblages are death assemblage s , n ot life assemblage s . It could be that a n event such as a large storm or hurricane forced these animals to the area where they later died. A Late Pliocene assemblage from western Florida contained hundreds of individuals of a single species of cormorant that likely died in a single event, likely a result of to xic poisoning in a red tide (Emslie 1995 b ). It should also be noted that Phalacrocorax carbo and Pinguinus impennis specimens have been recovered in Holocene shell mounds when conditions could have only been slightly colder than at present (see Remarks abo ve) .

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67 CHAPTER 5 CONCLUSIONS Only a handful of terrestrial vertebrate localities have been identified from the Okeechobee Formation or from the Late Irvingtonian of Eastern Florida . While the DCP avifaunal assemblage is small and sediments were not screenw ashed , a surprising number of species from disparate habitats is represented. A number of Great Auks ( Pinguinus impennis ) , a Northern Gannet ( Morus bassanus ) , and an extinct albatross ( Phoebastria aff. albatrus ) likely represent diseased or malnourished in dividuals. More terrestrial species such as transitional turkeys ( Meleagris aff. gallopavo ) , large cranes ( Grus sp.) , and Carolina Parakeets ( Conuropsis carolinensis ) were likely washed into the area by a freshwater stream. Other species described inhabit the tidal streams, marshes, and shallow embayments that typify the depositional environment of the site. Most of the avian taxa represented at the DCP are extant or only recently extinct. Alt hough some marine species have north ern affinities, none of the b irds represent s the western or tropical influences that characterize the avi faunas of several other , earlier Pleistocene localities. The fossils identified and described above provide a new insight into the riverine and co a stal Late Irvingtonian avifauna o Atlantic Coast , a tidal wetland community remarkably similar to those observed in the area today with a few major exceptions .

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68 REFERENCES Check list of North American Birds, 7th ed. American Ornithologist , Washington, D.C . Baumel, J. J. 1993 . Handbook of avian anatomy: N omina anatomica avium , 2 nd ed. Nutta ll Ornithological Club, Cambridge. Bell, C. J., E. L. Lundelius , A. D. Barnosky, G. W. Graham , E. H. Lindsay, D. R. Ruez , H. A. Semken, S. D. Web b, and R. J. Zakrzewski . 2004 . The Blancan, Irvingtonian, and Rancholabrean mammal ages. Pages 232 314 in Late Cretaceous and Ce nozoic Mammals of North America ( M. O. Woodburne , Ed .). Columbia University Press, New York . Bond , G. et al. 1992. Evidence for massive discharges of icebergs into the North Atlantic Ocean during the last glacial period. Nature 360:245 249. Brisbin, Jr., I. Lehr, and T. B. Mowbray. 2002. American Coot ( Fulica americana ) . In The Birds of North America Online (A. Poole, Ed.) , Ithaca. Brodkorb, P. 1960. Great Auk and Common Murre from a Florida Midden. Auk 77:342 343. Burdette, K. E, W . J. Rink, D . J. Mallinson, G. H. Means, and P. R. Parham. 2013 . Electron spin resonance optical dating of marine, estuarine, and ae olian sediments in F lorida, USA. Quaternary Research 7 9 : 66 74. Campbell, K. E. Jr. 1980. A review of the Rancholabrean avifauna of the Ichetucknee River, Florida. N atural History Museum of Los Angeles County Contributions in Science 330:119 129. Chesser, R. T., R. C. Banks, F . K. Barker, C. Cicero, J. L. Dunn, A. W. Kratter, I. J. Lovette, P. C. Rasmussen, J. V. Remsen, J. D. Rising, D. F. Stotz, and K. Winker. 2013. Fifty Fourth Supplement to the American Ornithologists' Union Check List of North American Birds . Auk 130:558 5 72. Clark, R. R. 2004. Hurricane Frances & Hurricane Jeanne: Post Storm Beach Conditions and Coastal Impact Report with Recommendations for Recovery and Modifications of Beach Management Strategies. Florida Department of Environmental Protection Division o f Water Resource Management Bureau of Beaches and Coastal Systems, Tallahassee, Florida. Dansgaard, W. et al. 1993. Evidence for general instability of past climate from a 250 kyr ice core record. Nature 364:218 220. Dyke, G. J., R. L. Nudds, and C. A. Wa lker. 2007. The Pliocene Phoebastria Diomedea anglica 631.

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69 Eadie, J. M., M. L. Mallory , and H. G. Lumsden. 1995. Common Goldeneye ( Bucephala clangula ). In The Birds of North America Online (A. Poole, E d.). Ithaca . Emslie, S. D. 1995 a . An early Irvingtonian avifauna from Leisey Shell Pit, Florida. Bulletin of the Florida Museum of Natural History 37:299 344. Emslie, S. D. 1995 b. A catastrophic death assemblage of a new species of cormorant and other seab irds from the Late Pliocene of Florida. Journal of Vertebrate Paleontology 15:313 330. Emslie, S. D. 1998. Avian community, climate, and sea level changes in the Plio Pleistocene of the Florida Peninsula. Ornithological Monographs 50: 1 113. Erskine, A. J. 1972. The Great Cormorants of eastern Canada. Canadian Wildlife Service Occasional Papers 14. Fischer, K. 1971. Ein flugunfähiger Kranich aus dem Pleistozän von Kuba. Monatsschrift für Ornithologie und Vivarienkunde 15:270 271. Fischer, K., and B. Stephan. 1971. Ein flugunfähiger Kranich ( Grus cubensis n. sp.) aus dem Pleistozän von Kuba: Eine o steologie der Familie der Kraniche (Gruidae). Wissenschaftliche Zeitschrift der Humboldt Universität zu Berlin, Mathematisch N aturwissenschaftliche Reihe 20: 541 592. Fradkin, A. 1980. Bird remains from two south Florida p rehistoric sites. Florida Scientist 43: 111 115. Franklin, J. L., R. J. Pasch, L. A. Avila, J. L. Beven, M. B. Lawrence, S. R. Stewart, and E. S. Blake. 2006. Atlantic hurricane season of 20 04. Monthly Weather Review 134: 981 1025. Frederick, P. C. and D . Siegel Causey. 2000. Anhinga ( Anhinga anhinga ). In The Birds of North America Online (A. Poole, Ed.). Ithaca. Gaskell, J. 2004. Remarks on the terminology used to describe developmental behaviour among the auks (Alcidae), with particular reference to that of the Great Auk Pinguinus impennis . Ibis 146: 231 240. Hay, O. P. 1902. On the finding of the bones of the Great Auk ( Plautus impennis ) in Florida. Auk 19:255 258. Hamon, J. H. 1959. Northern birds from a Florida Indian midd en. Auk 76: 533 534. Harmon, R.S., R.M. Mitterer, N. Kriausakul, L.S. Land, H.P. Schwarcz, P. Garrett, G.J. Larson, H.L. Vacher, and M. Rowe. 1983. U Series and a mino a cid r acemization g eochronology of Bermuda: Implications for e ustati c s ea level fluctuation over the past 250,000 y ears. Palaeogeography, Palaeoclimalolo gy, Palaeoecology 44:41 70.

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72 Ritchie, T. L. 1980. Two mid Pleistocene avifaunas from Coleman, Florida. Bulletin of the Florida Museum of Natural History 26:1 36. Roy, C. L., C. M. Herwig, W. L. Hohman, and R. T. Eberhardt. 2012. Ring necked Duck (Aythya collaris). In The Birds of North America Online (A. Poole, Ed.). Ithaca. Schubert, B. W., R. C. Hulbert Jr. , B. J. MacFadden, M. Searle, and S. Searle. 2010. Giant short faced bears ( Arctodus simus ) in Pleistocene Florida USA, a substantial range extension. Journal of Paleontology 84:79 87 . Scott, T. M. 1992. Coastal plains stratigraphy: The dichotomy of biostratigrap hy and lithostratigraphy a philosophical approach to an old problem. Pages 21 26 in the Plio Pleistocene Stratigraphy and Paleontology of Southern Florida (Scott, T. M. and W. D. Allmon Eds.) . Florida Geological Survey Special Publication 36. Florida Geolo gical Survey , Tallahassee, Florida. Scott, T. M. 2001. Text to Accompany the Geologic Map of Florida. Tallahassee: Florida Geological Survey Open File Report 80. Tallahassee, Florida. Scott, T. M., K. M. Campbell, F. R. Rupert, J. D. Arthur, T. M. Missimer , J. M. Lloyd, J. W. Yon, and J. G. Duncan. 2001. Geologic Map of the State of Florida . Florida Geological Survey, Florida Department of Environmental Protection, Florida Map Series 146. Florida Geological Survey . Tallahassee, Florida. Sellards, E. H. 1912 . The soils and other surface residual materials of Florida. Pages 7 79 in Florida Geological Survey Fourth Annual Report. Florida Geological Survey . Tallahassee, Florida. Seward, L., S. D. Chapman, and A. P. Currant. 2006. A catalogue of British Pleistoce ne birds identified by Colin JO Harrison and stored in the Natural History Museum, London, (Department of Paleontology ). Historical Biology 18 :235 254. Shufeldt, R. W. 1886. Osteology of Conurus (sic) carolinensis . Journ al of Anatomy and Physiology 20 : 405 425 . Snyder, N. F. R., and K. Russell. 2002. Carolina Parakeet ( Conuropsis carolinensis ). In The Birds of North America, no. 667 (A. Poole and F. Gill, Eds.). Birds of North America, Philadelphia. Spencer, R. S. and L. D. Campbell. 1987. The fauna and pale oecology of the Late Pleistocene marine sediments of southeastern Virginia. Bulletins of American Paleontology 92 :1 124. Steadman, D. W. 1980. A review of the osteology and paleontology of turkeys (Aves: Meleagridinae). N atural History Museum of Los Angeles County Contributions in Science 330:131 207.

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73 Steadman, D. W. 2008. Doves (Columbidae) and cuckoos (Cuculidae) from the early Miocene of Florida. Bulletin of the Florida Museum of Natural History 47:49 72. Stevenson, H. M. , and B. H. Anderson. 1994. The bird life of Florida. University Press of Florida, Gainesville. Stewart, J. R. 2002 . Sea birds from coastal and non coastal, archaeological and or igi n. Acta Zoologica Cracoviensia 45 :167 178 . Su á rez, W. and S. L. Olson. 2003. New records of storks (Ciconiidae) from Quaternary asphalt deposits in Cuba. The Condor 105:150 154. Watts, B. D. 2011. Yellow crowned Night Heron (Nyctanassa violacea). In The Birds of North America Online (A. Poole, Ed.). Ithaca. Webb, S. D. 1974. Pleistocene mammals of Florida. The University Press of Florida, Gainesville. Weigel , P. H. 1958. Great Auk remains f rom a Florida shell midden. Auk 75:215 216.

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74 BIOGRAPHICAL SKETCH John Andrew Kilmer was born in 1987, in DuBois, Pennsylvania. He graduated from DuBois Area High School in June, 2006. He attended Susquehanna University and received a Bachelor of Science degree in e cology in May, 2010. He worked and volunteered i n the Florida Museum of Natural History and as a graduate teaching assistant from 2010 to 2013 and now works as a field technician with the National Ecological Observation Network (NEON) . Andy enjoys backpacking , canoeing, caving, and birding.