GEOLOGY OF THE INTERMOUNTAIN WEST an open-access journal of the Utah Geological Association ISSN 2380-7601 Volume 11 2024 This is an open-access article in which the Utah Geological Association permits unrestricted use, distribution, and reproduction of text and figures that are not noted as copyrighted, provided the original author and source are credited. Email inquiries to GIW@utahgeology.org. PIECING TOGETHER A PREHISTORIC PUZZLE: REGIONAL INFERENCES OF MICRO- AND MACROSCOPIC ANALYSES OF POSSIBLY ONE OF THE LAST HYBRID MAMMOTHS IN MAINLAND WESTERN NORTH AMERICA Kate Morrison, Natalya Usachenko, Jonathan Erdman, Shilah Waters, and Renee L. Love GEOLOGY OF THE INTERMOUNTAIN WEST an open-access journal of the Utah Geological Association ISSN 2380-7601 Production Cover Design and Desktop Publishing Douglas A. Sprinkel Cover Adolescent to young adult Jeffersonian Mammoth in Soda Springs, Idaho. 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Lund Utah Geological Survey, Emeritus 435.590.1338 williamlundugs@gmail.com Editors GEOLOGY OF THE INTERMOUNTAIN WEST an open-access journal of the Utah Geological Association Volume 11 2024 21 ABSTRACT We evaluated the depositional age, taxonomy, diagenetic alteration, and osteology of mammoth skele- tal remains from southeastern Idaho. In this study, we identified the first record of M. jeffersonii present in Idaho and only the second record of the species in Western North America that lived 13,586 to 13,444 cal BP. The mammoth remains were preserved in an ancient hot spring deposit and have indicators of possible pre-mortem injuries. The diagenetic processes post-mortem suggest that it was not immediately buried and was gnawed on by small and large carnivores. Our evaluation of the mammoth’s tusks, molars, and limb bones suggest that these remains belonged to a young adult male that had been around 29 years old at its time of death. This specimen lived at a time when mammoths were becoming endangered in western North America before their ultimate disappearance from the fossil record. Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Kate Morrison1, Natalya Usachenko2, Jonathan Erdman3, Shilah Waters4, and Renee L. Love5 1Department of Earth and Spatial Sciences, University of Idaho, Moscow, ID 83844-3022 USA; broo4823@vandals.uidaho.edu 2 Department of Geosciences, Mississippi State University, Mississippi State, MS, 39762 USA; nu52@msstate.edu 3Department of Earth and Spatial Sciences, University of Idaho, Moscow, ID 83844-3022 USA; erdm4467@vandals.uidaho.edu 4Johns Hopkins Bloomberg School of Public Health, Baltimore, MD 21205, USA; sloosle1@jh.edu 5Department of Earth and Spatial Sciences, University of Idaho, Moscow, ID 83844-3022, USA; rlove@uidaho.edu Citation for this article. Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L., 2024, Piecing together a prehistoric puzzle—regional inferences of micro- and mac- roscopic analyses of possibly one of the last hybrid mammoths in mainland Western North America: Geology of the Intermountain West, v. 11, p. 21–44, Supplemental Material, https://doi.org/10.31711/giw.v11.pp21-44. INTRODUCTION In the Late Pleistocene, 126 to 11.7 ka, Mammuthus primigenius (Woolly Mammoths) and Mammuthus colum- bi (Columbian Mammoths) had established populations in North America. Near the end of that epoch, between 14,690 to 12,890 Ka, Earth was experiencing the Bølling-Allerød Interstadial period, an abnormally warm period during the Ice Age that preceded the Younger Dryas (Broecker et al., 1988). It marks the retreat of the Laurentide and Cordille- ran Ice Sheets and is a period where climate change played a major role in altering the North American biome. As the ice sheets melted, cold, dry conditions transitioned into a wetter, more temperate environment typical of modern conditions (Doerner and Carrara, 2001). A north-south division of biomes allowed for multi- ple species of Mammuthus to cohabitate through niche partitioning (Yansa and Adams, 2012; Lister, 2017). For nearly 80 years, researchers (Osborn, 1942; Agu- ire, 1969; Saunders et al., 2010; Yansa and Adams, 2012; Enk et al., 2016; Widga et al., 2017) debated the validity of a third species: Mammuthus jeffersonii (Jeffersonian Mammoth). Recent genetic studies have suggested that the M. jeffersonii could either be a M. primigenius, M. columbi hybrid (Fisher, 2009; Enk et al., 2011), a sub- species of M. columbi (Aguire, 1969; Maglio, 1973; Gra- ham, 1986; Lister, 2017), or the product of introgression, which includes the hybridization and repeated back- crossing of genes between species through time (Enk et 22 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 al., 2016). Genes of the extant African elephant (Loxo- donta africana), close relative to both Mammuthus and Elephas, show introgressive behavior with recurrent backcrossing; this implies a genomic record response to ancient habitat changes (Roca et al., 2005). Palynological records (Doernner and Carrara, 2001) indicate a transition from a colder, sedge brush-domi- nated landscape to a higher concentration of spruce and pine, and more temperate conditions similar to those of the modern day during the Bølling-Allerød period. This is coincident with sedge-grasslands disappearing into more closed forests in the Great Lakes region and M. jeffersonii competing with mastodon food sources (Yan- sa and Adams, 2012). The long history of introgressive radiation within mammoth populations through major fluctuations of interglacial and glacial periods, shows that they were not only interbreeding but also compet- ing for food resources to survive (Yansa and Adams, 2012; Enk et al., 2016). Mammoth populations were de- clining, eventually leading to the megafaunal extinction event on the North American continental mainland (Bjorck et al., 1988; Agenbroad, 2005; Barnosky et al., 2015). Megafauna, including mammoths, disappeared from the rock record, and a total of 35 mammalian gen- era went extinct or became extremely endangered, with 90% of all mammals over 45 kg (99 lbs) disappearing from the continent (Gilmour et al., 2015). The primary driver of this extinction is still debated, although a com- bination of climate change and anthropogenic influenc- es appears to be the most likely cause (Fisher, 2018). A mammoth skeleton (specimen UISSM-001-CO- LA) was excavated from a hydrothermal spring depos- it 11 km (7 mi) north of Soda Springs, Idaho, in 1966 and is now located at the University of Idaho (Jones and Bowers, 1968; Figure 1). This deposit was postulat- ed to be Upper Pleistocene in age (Malde and Powers, 1962; Jones and Bowers, 1968) based on nearby geolog- ic mapping and seven species of freshwater gastropods that were discovered between 61 to 91 cm (2–3 ft) above the mammoth remains. Original reconnaissance by R. Jones in 1968 mapped a tufa rim of an ancient discontin- uous ‘Pleistocene Spring’ that was about 1.6 km (1 mi) in length. Nearly 100 springs have been mapped in the vicinity of Soda Springs and the Aspen Range to the east (Lewicki et al., 2013) and many have been associated with active and non-active accumulations of travertine deposits (Lewicki et al., 2013). The Pleistocene Spring is oriented against the east slope of a north-south-orient- ed fault block, along the trend with China Hat (Welhan et al., 2014; McCurry et al., 2015; Welhan and Breed- lovestrout, 2016). The north-south-oriented fault block is part of the Paris thrust fault system within the Sevier fold and thrust belt in southeastern Idaho (Lewicki et al., 2013) and hydrothermal activity is associated with the Quaternary Blackfoot Volcanic Field (Welhan et al., 2014). Although more bones were searched for in the vicinity, the Pleistocene Spring was the only deposit to contain mammoth bones, which were discovered 2 m (6 ft) beneath the modern ground surface. Here, we use several analytical techniques to eval- uate taxonomy and paleontological history of UISSM- 001-COLA. Specifically, the focus of this study was 5-fold: (1) precise radiometric dating of the molar, (2) pre- and post-mortem analysis of the health of the mammoth, (3) size, age, and gender determination, (4) diagenetic analyses, including recrystallization of bones and analysis of original material, and (5) taxonomic identification using the dimensions and morphology of the proboscidean’s molars and mitochondrial DNA (mtDNA) analysis. This study provides a snapshot into the life of one of the last mammoths recorded in the fos- sil record on mainland Western North America during a time of great stress for Ice Age megabeasts, and pro- vides unique insight into the impact of extinction driv- ers on Pleistocene megafaunal specimen. This megafau- na specimen also records the presence of Mammuthus jeffersonii in the western United States, an area largely devoid of confirmed fossil evidence of the species. MATERIALS AND METHODS Radiometric Dating Using a sterilized drill, material from the center of the molar was extracted and placed in a sterile plas- tic container. Samples were analyzed by Beta Analytic Testing Laboratory to obtain an Accelerator Mass Spec- trometry (AMS) radiocarbon date. Pretreatment meth- ods included washing the sample in hot hydrochloric 23 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 acid (HCl) and alkali (NaOH) to remove carbonates and organic acids. Next the sample was bathed in sodi- um chlorite (NaClO2) to eliminate wood cellulose that might be present. The techniques used to measure car- bon-14 in the tooth utilized accelerator mass spectrom- etry. In calibrating the conventional radiocarbon date to calendar year equivalents, the High-Probability Density Range Method, INTCAL13, was used. The standard de- ? Snake River Plain 5100 - 4300 ? Legend Occurrence Mammoth Species Dated Age 1 ( ) Colpen Spring, WA M. jeffersonii 12,000 2 ( ) Port of Clarkston, WA M. columbi 12,000 3 ( ) Tolo Lake, ID M. columbi (possible) M. primigenius 5100 - 4300? 4 ( ) Grove Mammoth Site, ID M. columbi 14,700 5 ( ) Ra�lesnake Cave, ID Mammuthus sp. 10,450 +/- 120 6 ( ) Owl Cave, ID Mammuthus sp. 12,850 +/- 150 12,250 +/- 200 10,920 +/- 150 7 ( ) Soda Springs, ID M. jeffersonii ~13,500 8 ( ) Preston, ID M. columbi 12,150 Figure 1. Mammoth distribution in North America. Gray shaded area represents the inferred range of mammoths during ter- minal Pleistocene (15 to 10 Ka) based on Agenbroad (2005) and Mammuthus sp. in Faunmap (Graham and Lundelius, 2010). White shaded areas show extent of glaciation at this time. Mammoth discoveries attributed to M. jeffersonii are represented with a black square. Other mammoth species discovery sites are indicated with a black circle. Inset: Mammoth discoveries in Washington and Idaho that have been radiocarbon dated within 15 to 10 Ka. Descriptions of localities are included. Sites 1 through 3 sourced from Sappington (2019) and sites 4 through 8 are sourced from Agenbroad (2005). 24 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 viations are not reported less than ±30 BP to prevent from misrepresenting the accuracy of the results. Pre- and Post-Mortem Analysis External and internal features such as calluses, fractures, enlarged pores, distinct porosity, and ulcers identified on the mammoth skeleton can indicate a pre-mortem event or skeletal-related disease (Krze- minska and Wedzicha, 2015; Krzeminska et al., 2015; Leshchinskiy, 2017). A total of 271 bones and bone frag- ments were imaged at Gritman Medical Center in Mos- cow, Idaho, using computed tomography (CT) technol- ogy (see Supplemental Material for inventory of bones). The scans provided 18 DICOM files and full images of the limbs, axial skeleton, and manus bones of the mam- moth. The images were analyzed using  the process- ing  software:  Analyze 14.0 (AnalyzeDirect, Overland Park, KS), MicroDicom Viewer (MicroDicom Ltd, So- fia, Bulgaria), and ITK-SNAP (Yushkevich et al., 2006), and viewed in 3D Microsoft Paint to measure any ab- normalities found internally such as enlarged pores and fractures and externally such as calluses and ulcers. The scans provided the ability to create 3D modeling of each bone for further assessment of surface features. The bones of the mammoth were then organized into a categorized inventory and evaluated based on ex- ternal features. To determine how much of the skeleton had been recovered, the specimen was divided into the following regions: limbs, vertebral column and ribcage, autopodial bones, crania, tusk, and pelvis (Supplemen- tal Material). These features were recorded, along with a detailed description of preservation quality. Destructive features were documented and measured, and preserva- tion quality was assessed using this scale: (1) very poor- ly preserved or severely fragmented, (2) poorly pre- served or fragmented, (3) mostly preserved or slightly fragmented, and (4) well preserved or not fragmented (Table 1). The regions were then given a total percent- age present value based on observational analysis and the preservation quality given. The total percentage of skeleton present was achieved from the average value of each region’s total percentage intact combined. Size, Age, and Gender Estimates The ratios of metacarpal (MTC) III length vs. radius length and manus height vs. radius length of the skele- ton were determined based on bone measurements and compared to those of different proboscidean species from Larramendi (2015) to show a comparison in body proportions. Manus height was calculated from Larra- mendi (2015) in which the length of the third metacar- pal is multiplied by two. This is under the assumption that the length of the third metacarpal represents ap- proximately 25% of the radius length in most probosci- deans (Table 2). Using the following equation: A = B x C where: A = length of the specimen’s frontal limb bone. B = established percentage that the length of the respective limb bone is of the animal’s total height. C = total height from manus to shoulder, the height was estimated. This height estimation was then compared to skeletal percentages for both M. primigenius and M. columbi (Larramendi, 2015; Table 3). To remain consistent with the calculations made for manus height in this study, the length of the radius was used to calculate the shoulder height of UISSM-001-COLA. To determine the molar set that the mammoth was on at the time of death, we assessed the quantities of plates that were fully erupted and in wear, plates that were actively erupting and in wear, and plates that were in the process of erupting. These were compared to the data sets in Roth and Shoshani (1988) and Lee and oth- ers (2012), which estimated molar set number accord- ing to these parameters. Some authors (Lee et al., 2012; Lister, 2017; Widga et al., 2017) use the designation of m1-m3 for the premolars and M1-M3 for the adult mo- lar sets. Here, we use the designation of M1-M6, which does not provide a distinction between the pre-mo- lars and molars but lists all molars in order of eruption (Stansfield, 2015). 25 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 Three methods were used to estimate UISSM-001- COLA’s age at its time of death: (1) Dentition – The approximate age of the mam- moth was first estimated using dental analysis as outlined by Roth and Shoshani (1988). The length, width, height, and plate number of the specimen’s right maxillary molar were measured to infer the re- spective molar set value and associated age range. In addition, states of eruption and wear were record- ed on the four exposed mandibular molars. This method utilized the comparison of these measure- ments to fossil and extant Elephas maximus (Asian elephant) molars of known age, which are the sister taxon to Mammuthus primigenius (Roca et al., 2015; Enk et al., 2016), to provide an accurate analogy of elephantid maturation. These results were addi- tionally compared to standards established in Laws (1966), as well as the records presented in Lee et al. (2012) and Haynes (2017), which assessed the mo- lar characteristics of Loxodonta africana (African elephant). (2) Post-Cranial Skeletal Assessment – Age param- eters were further determined through the exam- ination of the degree of epiphyseal fusion in the intact proximal and distal regions of UISSM-001- COLA’s humerus, ulna, radius, tibia, and femur, which were categorized as fused, partially fused, or unfused (Lister, 1999; Haynes, 2017; Table 4). Post- cranial bone growth increases in mammals until their epiphyses are fused to their diaphysis. African Elephant Equivalent Years (AEY; Lister,1999) are used to report relative mammoth ages. These were cross-referenced with the skeletal growth rates of modern proboscideans (Haynes, 2017), molar set number, and gender determination. (3) Age Reference Line – The final method utilized reference points along the lower jaw to determine the Age Reference Point (ARP) and Age Reference Line (ARL) following the methods of Stansfield (2015; Figure 2). Using observed tooth lamellae along the endpoint of the ARL, an additional age estimation was made. A hypothesis of the mammoth’s gender was made predominantly using its tusk characteristics, which in- cluded girth and curvature (Averianov, 1996; El Adli et Region Total % Present Preservation Notes Limbs 75 3 Left femur and ulna fragmented. Right femur not recovered. Vertebral Column and Rib Cage 25 2 Many ribs and vertebrate with identified gnaw marks. Several missing components on the spinous process and rib cage. Autopodial 95 4 Some phalanges missing. Skull 20 2 Primarily fragments except for three, fully intact molars. Skull currently being reconstructed. Tusks 90 1 Highly fragmented and fragile. Supported by plaster jacket. Pelvis 5 1 Highly fragmented. Currently being reconstructed. Entire Mammoth 51.70 2 Calculations based on the average percentage of each region. Table 1. Total percentage of specimen UISSM-001-COLA intact based on preservation quality. 26 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 al., 2015), and the degree of epiphyseal fusion observed in the radius, ulna, humerus, tibia, and femur (Lister, 1999; Stanfield, 2015; Haynes, 2017; Figure 3 and Table 4). Although an assessment of the pelvis would be ideal for more accurate gender determination, UISSM-001- COLA's pelvis was severely fragmented upon excavation and could not be analyzed. Instead, since mammoths were sexually dimorphic in body size and differed in their rates of skeletal maturation, we were able to use the size, shape, and length of their tusks to determine gender (Laws, 1966; Averionov, 1996; Haynes, 2017). The appearance of the mandibular condyles was also considered; in males, these structures are more circu- lar, whereas in females they are more ovular (Yacobi et Table 2. Ratios between metacarpal (MTC) III length vs. radius length and manus height vs. radius length of different pro- boscidean species found in Larraramendi (2015). Radius length from the proximal to distal end. Species Individual MTC III length (mm) Radius length (mm) Calculated manus height (mm) MTC III length vs. radius length (%) Manus height vs. radius length (%) Determined in this study UISSM-001-COLA 210 889 420 23.6 47.2 Mammuthus primigenius Pfannerhall 208 825 416 25.2 50.4 Mammuthus primigenius fraasi Steinheim 245 955 490 25.7 51.3 Mammuthus trogontherii Zhalainuoer III 255 985 500 25.9 51.3 Mammuthus meridionalis Scoppito 266 950 525 28 51.8 Mammuthus meridionalis Nogaisk 265 1040 530 25.5 55.3 Mammuthus columbi MSL-140 237 948 474 25 51 Mammuthus columbi NSM1597-62-2 194 823 388 23.6 50 Mammuthus columbi SDSM 124688 244 928 488 26.3 47.2 Table 3. Height estimation of specimen UISSM-001-COLA based on limb bone measurements compared to specimens of Larramendi (2015). Mammoth Species Larramendi (2015) Percentages UI-SSM-001COLA Radius Length (cm) Total Height of UISSM-001 COLA (cm) M. primigenius 26.67 88.9 333.33 M. columbi 27.10 88.9 328.04 27 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 al., 2004). The last indirect method used to determine gender was to assess the epiphyseal regions in the limb bones; in females, the epiphyses would typically finish fusing between ages 30 and 35, and in males, between 45 and 50 (Lister, 1999; Haynes, 2017). Diagenetic Analysis The fossilization of bone is characterized by the re- placement of protein and other organic biomolecules with inorganic material and the recrystallization of in- organic remains, such as hydroxyapatite in bone, into Table 4. Specimen UISSM-001-COLA's limb bones with degree of epiphyseal fusion. Findings are compared to Lister (1999) and suggest an age range of around 29 AEY. Bone Epiphyseal fusion Dental age (AEY) from Lister (1999) Haynes (2017) corrected age using Stansfield (2015) Distal humerus Fused ≥6 ≤24 Proximal tibia Fused ≥26 27-30 Distal tibia Fused ≥26 ≤24 Proximal ulna Fused ≤34 ≤24 Distal femur Fused ≤34 29-33 Proximal humerus Unfused ≤41 40-47 Proximal femur Unfused ≤43 37-42 Distal radius Unfused ≤43 Distal ulna Unfused ≤43 ≥52 Figure 2. Age estimation using the ARL (Stansfield, 2015). (A) A straight line was drawn from point A (the base of the man- dibular foramen) to point B (the most distal section of molar occlusal wear). A second line, C, was visualized along the ridge of the medial mandible, and intersects the first line at point D, which is known as the ARP. From this point, a distance of 10 cm (3.94 in) was measured, passing centrally, through the distal molar. This is the ARL. The yellow line E marks the endpoint of the 10-cm (3.94 in) measurement in this example from Stansfield (2015). (B) Age Reference Line calculation on specimen UISSM-001-COLA. 28 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 a more stable form (Pfretzschner, 2004; Buckley and Wadsworth, 2014; Keenan et al., 2015; Kendall et al., 2018). Environments most suitable to preservation are those that are unfavorable to microbial activity; remains most likely to avoid diagenesis are those consisting of or encased by hydrophobic, inorganic lattices that are impermeable to water and are, therefore, protectant against biotic and abiotic decay (Hedges et al., 1995; Trueman and Martill, 2002; Jans, 2008; Keenan, 2016). Therefore, some hard surface remains, such as bone and teeth that have inorganic casings, can protect biomole- cules including nucleic acids, proteins, lipids, and car- bohydrates from degradation. This can, in turn, enable certain chemical and molecular analyses, which could provide insights into genetic information and environ- mental conditions. To determine the diagenesis of the bones, and to assess whether original material was remaining in the bones for genetic analysis, the external surface structure of a tusk and tooth were imaged using scanning electron microscopy (SEM). Superficial samples were collected from the tusk and tooth for analysis. The samples were prepared with a carbon coating to gain high resolution and imaging under the SEM. Elemental composition of each specimen was determined utilizing SEM with en- ergy dispersive spectroscopy (SEM/EDS). This analysis was performed on both external surface samples and in- ternal samples. Internal samples were obtained by drill- ing into the tusk, tooth, and femur. Two samples were collected from the interior tusk, three from the tooth (outer, middle, inside), and two from the femur (epiph- ysis and diaphysis). Samples were then imaged without a carbon coat to obtain accurate elemental composition profiles from EDS. The molecular composition of the tusk was acquired using x-ray diffraction (XRD) per- formed at 40 kV. A total of 1561 data points were gener- ated over a spectrum of 0 to 80°. Steps equaled 0.05° in measurements of two-theta. Figure 3. Curvature of tusks, length, and girth. Green arrow showing curved length of 266 cm (105 in). Yellow arrow showing a non-curved length of 115 cm (45 in). Red arrow showing width of distal end at 18 cm (7.1 in). Inset image showing the circularity of mandibular condyle. 29 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 Taxonomic Identification Proteinaceous composition of UISSM-001-CO- LA was determined using methods of Cappellini et al. (2018) to determine if original material was remaining in the specimen for radiocarbon dating and taxonomic analysis. A femoral sample was collected using a steril- ized drill at the junction between the articular surface of the knee and the shaft, just proximal to the epicondyle, at a depth of approximately 1 cm (0.4 in). When drilling into the bone, the periosteum was discarded to mini- mize contamination. Treatment buffer was prepared using 0.125 M Tris-Cl, 4% SDS, 20% v/v glycerol, 0.2 M DTT, and 0.02% bromophenol blue with a pH of 6.8. A sample of 15 mg was boiled for 15 minutes and then the sample was loaded into an SDS gel composed of a 10% resolving gel and a stacking gel. Polyacrylamide gel electrophoresis (PAGE) was run for approximately 30 minutes to allow separation of the proteins and oth- er components of the femur. The gel was stained with Coomassie brilliant blue R-250 solution. Taxonomic identification of UISSM-001-COLA was determined in two ways: (1) by using measure- ments of molar characteristics adapted from Maglio (1973) and Lister (2017), and (2) by extracting mtDNA from a tooth sample. Measurements of UISSM-001- COLA’s molar included length (L), width (W), height of crown (H), enamel plate count (P), lamellar frequency (LF), and hypsodonty index (HI). The LF was calculat- ed on an average of six measurements taken from the upper, middle, and lower portion of the crown on both the lingual and buccal side. The HI was calculated as HI = H/W x 100, or height of the crown as an expres- sion of width standardized to a length of 100 mm (3.9 in). These measurements were obtained using a digital caliper on the maxillary right molar (Figure 4). No esti- mation of missing plates was required due to the pres- ence of the anterior-most root. The molar parameters of the specimen were then compared between holotypes or neotypes of M. primigenius, M. columbi, and M. jef- fersoni (Roth and Shoshani, 1988; Lee et al., 2012; Lister and Sher, 2015; Haynes, 2017; Lister 2017; Widga et al., 2017; Table 5). Elemental composition analysis provided guidance on where to sample for mtDNA. The anterior talon of the upper molar was sent to the University of California, Santa Cruz Paleogenomics Lab, for analysis. Extraction and processing were performed according to Dabney et al. (2013). Initial sample preservation and assessment was performed using a single-stranded library prepara- tion and sequencing on an Illumina NextSeq 2x150 run targeting 1 million raw reads. The reads were trimmed and aligned to Loxodonta africana nuclear genome and Mammuthus primigenius mitochondrial genome. Sub- sequently, the sample underwent mtDNA preparation and analysis using methods according to Kirillova et al. (2017), myBaits v4.01 protocol (Biodiscovery, LLC dba Arbor Biosciences, 2018), and Vershinina et al. (2020). Sequencing occurred on an Illumina NextSeq 2 x 150 run targeting 0.5 million raw reads. Afterwards, the reads were trimmed, merged, and filtered using Se- qprep2 (https://github.com/jstjohn/SeqPrep), FASTX_ ToolKit (http://hannonlab.cshl.edu/fastx_toolkit/), and PRINSEQ lite (Schmieder and Edwards, 2011). The final consensus sequence was generated using MIA (https:// github.com/mpieva/mapping-iterative-assembler) and compared to clades described in Enk et al. (2016) to as- sign UISSM-001-COLA to a haplogroup. RESULTS Radiometric Dating Radiocarbon dating was consistent with the esti- mated upper Pleistocene age derived from the gastro- pod identification and yielded a age of 13,586 to 13,444 cal BP with 95.4% probability (Figure 5; Beta Analytic Testing Laboratory specimen number: 524280). A C:N ratio was also obtained to test potential diagenetic alter- ation or contamination at 3.2, with an isotope ratio mass spectrometer reading of δ13C at –18.7 ‰ and δ15N at +11.32 ‰. Modern bone is in the range of 2.9 to 3.5, suggesting that little to no diagenetic alteration or con- tamination occurred (DeNiro, 1985; Enk et al., 2016). Pre- and Post-Mortem Analysis A total of 271 bones and bone fragments were item- ized (Supplemental Material). The pelvis and crania have significant damage and are currently in fragments. 30 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 Ta xo no m ic ID M ol ar Po si tio n Pl at e N um be r L en gt h H ei gh t W id th E na m el T hi ck ne ss L am el la r Fr eq ue nc y H yp so do nt y In de x Av er ag e R an ge Av er ag e R an ge Av er ag e R an ge Av er ag e R an ge Av er ag e R an ge Av er ag e R an ge Av er ag e R an ge M . c ol um bi M 3( M 6) R ig ht Si de 18 .9 18 -2 1 26 8. 6 18 0- 35 9 18 1. 1 16 1- 21 6 10 1. 3 69 -1 06 3. 1 2. 0- 3. 0 5. 6 5. 29 -7 .5 3 20 9. 1 17 6. 7- 25 3. 25 M 3( M 6) L ef t Si de 18 15 -1 9 26 8. 2 16 6- 31 5 17 9. 2 11 2- 22 0 91 .6 69 -1 08 2. 9 2. 0- 3. 0 6 5. 71 -7 .1 2 20 7 16 2. 32 - 24 3. 68 M 3( M 6) S id e U ns pe ci fie d 19 .1 18 -2 4 27 6. 3 26 0- 34 0 19 4. 5 15 2- 29 0 10 3. 8 74 -1 28 2. 4 1. 6- 3. 9 6. 1 5. 88 -7 .6 9 19 1. 4 15 8. 44 - 24 6 To ta l 18 .9 15 -2 4 27 2. 6 16 6- 35 9 19 0. 1 11 2- 29 0 10 2. 2 69 -1 28 2. 6 1. 6- 3. 9 6 5. 29 -7 .6 9 19 6. 1 15 8. 44 - 25 3. 25 M 2( M 5) S id e U ns pe ci fie d 10 .2 4. 0- 16 .0 11 9. 2 13 8- 19 9 14 7 13 1- 16 2 88 .1 68 -1 11 .4 2. 2 1. 5- 2. 5 6. 4 4- 8. 71 11 5 0- 21 7. 65 M . p ri m ig en iu s M 3( M 6) R ig ht Si de 24 .6 20 -2 8 27 2. 6 20 5- 32 8 17 3. 4 10 2. 1- 21 9 93 .5 72 -1 22 1. 8 1. 03 -2 .8 3 8. 5 6. 91 - 10 .9 4 18 9. 4 12 8. 27 - 23 0. 77 M 3( M 6) L ef t Si de 24 .7 20 -2 8 26 7. 4 19 5- 32 5 14 8. 8 10 2. 2- 17 9 96 .5 80 .9 -1 16 1. 8 1. 3- 2. 37 8. 3 7. 23 - 10 .1 6 15 8. 8 10 3. 44 - 18 7. 64 M 3( M 6) S id e U ns pe ci fie d 23 .8 21 -3 0 27 1. 8 26 0- 35 5 17 2. 6 15 5- 21 3 94 .9 75 -1 25 1. 5 1- 2. 1 8. 3 6. 15 -1 0 15 7. 6 16 8- 22 7. 6 To ta l 24 .2 20 -3 0 27 1 19 5- 35 5 16 7. 4 10 2. 1- 21 9 94 .9 72 -1 25 1. 6 1- 2. 83 8. 4 6. 15 - 10 .9 4 16 7. 4 10 3. 44 - 23 0. 77 M 2( M 5) S id e U ns pe ci fie d 13 .4 9. 0- 16 .0 16 6. 6 77 -2 14 13 1. 6 92 -1 62 .9 71 .2 50 -8 9 1. 6 1. 5- 1. 82 5 8. 8 6. 54 - 11 .7 6 11 3. 8 0- 20 1. 73 M . j eff er so ni i M 3( M 6) R ig ht Si de 22 .4 19 -3 1 28 7. 4 21 4- 38 0 17 8 13 4. 4- 24 9 10 1. 9 88 -1 20 2. 2 1. 5- 2. 9 7. 5 5- 10 .1 9 17 3. 5 74 .2 6- 23 5. 96 M 3( M 6) L ef t Si de 23 .4 22 -2 7 28 2 25 0- 30 5 15 3. 4 75 -2 10 94 .9 80 .7 - 10 8. 5 2. 5 1. 5- 2. 1 6. 6 7. 2- 8. 07 18 4. 7 13 3. 98 - 21 8. 56 M 3( M 6) S id e U ns pe ci fie d 27 27 29 5 29 5 20 6 20 2- 21 0 11 0. 6 10 3- 11 7 2 1. 3- 2. 7 8. 8 8. 8 18 8. 3 17 2. 65 - 20 3. 88 To ta l 23 19 -3 1 28 5. 8 21 4- 38 0 17 3. 4 75 -2 10 10 1. 6 80 .7 -1 20 2. 3 1. 3- 2. 9 7. 3 5- 10 .1 9 17 8 74 .2 6- 23 5. 96 M 2( M 5) S id e U ns pe ci fie d 15 24 1 83 1. 8 6. 2 0 U IS SM -0 01 - C O LA M 2( M 5) U pp er R ig ht 1 9 2 78 1 54 8 2. 8 2 .4 8 1 86 Ta bl e 5. T ot al av er ag es a nd ra ng es o f d en ta l p ar am et er s f ro m W id ga e t a l. (2 01 7) a nd L ist er ( 20 17 ) w ith co m pa ris on to th os e of sp ec im en U IS SM - 00 1- C O LA . N ot e: sp ec im en U IS SM -0 01 -C O LA 's m ea su re m en ts o f t he u pp er ri gh t M 5 m ol ar b et te r c oi nc id es w ith M . c ol um bi fo r t he p la te n um be r an d en am el th ic kn es s, an d M . p rim ig en iu s f or th e l am el la r f re qu en cy an d w id th . Th e l en gt h, h ei gh t, an d hy so do nt y in de x ar e s im ila r f or ea ch o f t he sp ec ie s. A ll m ea su re m en ts in m m . 31 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 The tusks have also experienced a large amount of frac- turing. The limb bones have significant breaks that sep- arate them into two to three fragments. The right hu- merus is missing except for the proximal epiphysis. The vertebral column only shows small amounts of fractur- ing, but the ribs have significant damage, and most are missing. An estimated 10% of the mammoth’s ribcage is present. Due to the large amount of fragmentation of the ribs, a full reconstruction was not possible. The ma- nus and pes bones are well-preserved and exhibit only small amounts of fragmentation. Observational analy- sis of the preservation quality showed that UISSM-001- COLA has an overall average quality number of 2 out of a scale of 4. The total percentage of UISSM-001-CO- LA’s intact skeleton was estimated to be 51.7% (Table 1). Whereas over half of the remains were present for anal- ysis, larger portions of the vertebral column, ribcage, crania, and pelvis were unrecovered or in fragments. External and internal observations revealed un- usual markings and features on the joints, limbs, and autopodial bones, and can be indicators of premortem health conditions and injuries. The ribs of the mam- moth contain gnaw markings with sizes ranging from presumed small mammalians to large carnivores. The marks on the ribs are currently under more investi- gation and will not be discussed further in this paper. An internal fracture, approximately 5.05 cm (2 in) in length, occurs inside the right calcaneum bone and is identified by a thickened callus around the fracture. The left calcaneum bone of the mammoth showed enlarged pore sizes of up to 3 mm (0.12 in) in diameter; this is in contrast to normal ranges for pores that are typically less than 0.1 mm (Leshchinskiy, 2012). Premortem de- formations by ulcers were identified by an area up to 6 cm2 (2.4 in2) with a linear dimension typically no larg- er than 7 cm (2.8 in). These features are common on Figure 4. Analysis of molars. (A through D) Assessment of molar parameters. (B through D) Characteristics of mammoth molars showing plate, lamellar frequency, and enamel thickness. Modified from Roth and Shoshani (1988). (E through F) Upper teeth. 32 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 the articular surface. This mammoth presents a feature that matches this description on the distal end of its hu- merus measuring 4 x 4.5 cm (1.6 x 1.8 in). These ulcers were compared to Leshchinskiy (2017) to confirm the identification. The seventh thoracic vertebra presents an oval-shaped deformation measuring 2 x 1.5 cm (0.8 x 0.6 in) which may be a premortem ulcer or a result of peripheral erosion on the articular surface. Specimen UISSM-001-COLA shows significant di- lation in the Haversian cavities of the axis. Studies have found that non-osteoporotic indications within Hav- ersian bones of these large animals would measure at typically less than 0.1 mm (Leshchinskiy, 2012). The vertebral column of mammoths can indicate external signs of osteological changes with distinct porosity rep- resented as numerous small, piercing holes (Krzemins- ka and Wojtal, 2015; Leshchinskiy, 2017). This feature is present in UISSM-001-COLA’s seventh thoracic verte- bra. These features are shown in Figures 6 and 7. Size, Age, and Gender Estimates Specimen UISSM-001-COLA’s jaw had a base alve- oli diameter of 18.9 cm (7.4 in) and was fairly circular, indicating the specimen may have been a male. Female alveolar diameters typically range between 8 and 10 cm (3 and 3.9 in) (Pilgrim and Western, 1983; Vereschcha- gin and Tikhonov, 1986; Averianov, 1996; Moss 1996). The length and curvature of the specimen’s tusk (Figure 3) suggest that UISSM-001-COLA was a sexually ma- ture male. Female tusks tend to be slenderer, straight- er, and more cylindrical in shape (Fisher, 2009), and whereas male tusks are generally more massive, more conical in shape, and turned medially at the adult stage (Lister and Bahn, 2007). Based on the specimen’s tusk characteristics, alveoli diameter, and the degrees of epiphyseal fusion in the limb bones (Figures 8 and 9; Table 4), it can be concluded that UISSM-001-COLA was a male mammoth. A morphological comparison of UISSM-001-CO- LA’s molars to those presented in Roth and Shoshani (1988) determined that UISSM-001-COLA would have already aged out of its first three sets of deciduous mo- lars. One of its adult molar sets was in wear at its time of death. Based off the dental aging standards of living elephants, mammoths would wear through and then lose each set of molars at approximately age 1, 5, 10, 22, Figure 5 . Conventional radiocarbon age for specimen UISSM-001-COLA is 13,586 to 13,444 cal BP. Date provided by Beta Analytics Laboratory. 33 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 35, and 65 (M1, M2, M3, M4, M5, and M6, respective- ly; Laws 1966; Lee et al., 2012). A comparison of mea- surements between UISSM-001-COLA’s molars and the values listed in Roth and Shoshani (1988) and Lister (2017) suggest that the specimen’s 4th set of molars (M4 molars) had fully erupted, were in wear, and were al- most completely gone, and its 5th molar set (M5) was al- most fully erupted and in wear, placing its approximate age between 22 and 35, respectively (Table 5; Figure 4). The Age Reference Line endpoint fell between the 10th and 11th lamella of the mammoth's left mandibular mo- Figure 6. (A) Premortem deformation by ulcer on the distal end of the humerus bone measuring 4 x 4.5 cm (1.6 x 1.8 in) outlined in white, dashed box. (A1) Premortem defor- mations by ulcers indicated by red arrows from Leshchinskiy (2012) for comparison. (B) Premortem ulcer or a result of peripheral erosion on the articular surface outlined in white, dashed box. (B1) Premortem deformations by ulcers indi- cated by white arrows from Leshchinskiy (2017) for com- parison. (C) Dilation of cavities opening into the vertebra foramen indicated by dashed, white box. (C1) Diliation of cavities opening into the vertebra foramen from Leshchins- kiy (2017) for comparison. (D) Distinct porosity in the sev- enth thoracic vertebra indicated by white arrows. (D1) Dis- tinct porosity shown with white arrows from Krzeminska et al. (2015) for comparison. Figure 7. CT rendering of the mammoth’s right (A) and left (B) calcaneum bones. Potential fracture indicated by red ar- row. Enlarged pores indicated by white circle. 34 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 lar (Figure 2). Since this molar is an M5, the estimat- ed age would be 28 to 29 AEY (using Lee et al., 2012, age class; Stansfield, 2015). Analysis of the mammoth’s front and rear limb bones revealed that none of the later epiphyseal plates had yet fully fused at its time of death, which places the estimated age range between 29 and 33 AEY (Figure 8) according to the classifications of Lister (1999) and Haynes (2017). Using the categories established by Lee et al. (2012), the specimen would lie somewhere between 27 and 31.5 years old. Stansfield’s (2015) age reference line corroborates these age ranges with an estimate of 28 to 29 years old (Figure 9). There- fore, the ontogenetic age of the specimen was postulat- ed to be 29 AEY, as this was the age in common between all these estimates. Diagenetic Analysis The SEM images revealed that the tusk exhibits high levels of porosity and permeability. The tooth, converse- ly, appears striated but generally remains predominantly non-porous (Figure 10). The EDS spectra demonstrated that most of the tusk, tooth, and femur are composed of calcium, phosphorus, and oxygen (Figure 11). Carbon is also present, but to a much smaller extent. Consid- ering the quality of the remains as well as the length of time since excavation, the major compound composing these samples is apatite – Ca10(PO4)6(OH)2. The EDS spectra further suggested some elemental replacement in the apatite, as illustrated by the presence of fluorine. These conjectures are confirmed according to the XRD performed on the tusk (Figure 12). Whereas the majority of the tusk, tooth, and femur samples contain only minimal amounts of carbon rel- ative to other elements, a high proportion of carbon occurs in some fragments of the interior tooth sample. However, the amount of carbon in the interior tooth sample is not homogenous; generally, the smaller frag- ments have higher fractions of carbon (Figure 11). The SEM revealed additional crystalline structures on the surface of the tusk and potential surface contaminants. EDS inspection of the crystalline structures demon- strates predominant components of calcium, phospho- rus, carbon, and oxygen. The main suspected surface contaminants appear fibrous and, according to EDS, have high levels of carbon. No protein occurred in the mammoth femur after processing and running PAGE. Taxonomic Identification The ratios found between the metacarpal (MTC) III length vs. radius length and manus height vs. radius length in UISSM-001-COLA were found to be small- Figure 8. Assessment of the epiphyseal fusion in limb bone fragments with scale. (A) Proximal tibia; fused. (B) Proxi- mal humerus; unfused. (C) Distal tibia; fused. (D) Proximal femur; unfused. (E) Proximal ulna; fused. (F) Distal radius; unfused. (G) Distal femur; fused. (H) Distal ulna; unfused. 35 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 Figure 9. Age estimation for specimen UISSM-001-COLA based on a correlation of the condition of its epiphyses (see Table 4) with prior correlations between molar wear and epiphyseal fusion in male and female specimens of Loxodonta africana. Modified from Haynes (2017). Overlap of the age estimate with the 5th set (M5) and degree of the epiphyseal fusion discounts that we could have only the 4th molar. 36 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 er than those in the comparison to other proboscide- an species, including Mammuthus primigenius, Mam- muthus primigenius fraasi, Mammuthus trogontherii, Mammuthus meridionalis, and Mammuthus columbi (Table 2; Larramendi, 2015). Individuals of M. colum- bi exhibited ratios nearest in value to those of UISSM- 001-COLA. The ratios found between the metacarpal (MTC) III length vs. radius length and manus height vs. radius length in UISSM-001-COLA were found to be smaller than those in the comparison to other pro- boscidean species, including Mammuthus primigenius, Mammuthus primigenius fraasi, Mammuthus trogonthe- rii, Mammuthus meridionalis, and Mammuthus columbi (Table 3; Larramendi, 2015). Individuals of M. columbi exhibited ratios nearest in value to those of UISSM-001- COLA. Specimen UISSM-001-COLA’s right maxillary had high lamellar frequency and thick enamel; The speci- men's tooth measurements match both those of M. jef- fersonii and M. columbi specimens (Table 5). Although different molars were measured (M5 in the case of this study compared to M6 in the holotypes), consistent similarities between these two molars have been con- sidered sufficient for taxonomic comparisons (Lister, 2017; Widga et al., 2017). After extracting and sequenc- ing mtDNA from the molar, the results confirmed that UISSM-001-COLA belonged to haplogroup F (Enk et al., 2016), the lineage of North American mammoths that includes M. columbi and M. jeffersonii (Figure 13). DISCUSSION Pre- and Post-Mortem Analysis Analysis of the percentage present and preservation quality (Table 1) of the bones indicate that some skeletal remains were fragmented and exposed to the surface. This is largely due to the absence of larger portions of the vertebral column and ribcage. There are rib bones missing and the ones that remain either have carnivo- rous gnaw marks or are in fragments. This leads us to conclude that the mammoth was exposed above the wa- ter before the remains were buried. Interpretation of the enlarged pores in the autopo- dium bones suggest that they are evidence of premor- tem lesions or deformation caused by an underlying bone disease. The pathologies, specifically osteopo- rosis, surrounding manus and pes bones are found to be commonly associated with ulcers and weakening of Figure 10. SEM images of specimen UISSM-001-COLA. (A through C) Tooth surface fragment. (D through F) Tuck surface fragment with scale. 37 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 the compact layer (Leshchinskiy, 2017). Pore widening within the vertebral column can be indicators of osteo- porosis or other skeletal-related diseases such as Kash- in-Beck disease (Krzeminska et al., 2015; Leshchinskiy, 2017). Defects within the spinous structure indicate the possibility of genetic defects or diseases (Krzeminska Figure 11. (A) SEM images of specimen UISSM-001-COLA tooth surface. (B through C) Interior tooth. (D) Tusk surface. (E through F) Interior bone with scale alongside respective energy dispersive X-ray spectroscopy (EDS) spectra. Red crosshairs represent the location of spot EDS analysis; images without crosshairs are alongside EDS spectra averaged across the full image. 38 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 et al., 2015). In addition, large marrow cavity openings into the vertebral foramen within the vertebrates of mammoths are also diagnostic of osteoporosis (Lesh- chinskiy, 2017). The internal fracture in the calcaneum could be an indicator of lameness (Leshchinskiy, 2015; Leshchins- kiy, 2017). There is also evidence that osteofibrosis-re- lated conditions can cause increased bone fragility that may have led to this fracture (Krezerminska et al., 2015). These observations provide insights into pos- sible bone diseases within the mammoth, whether it died because of bone disease or lameness, and pre- to postmortem states of the specimen. It is possible that the osteolytic conditions observed in UISSM-001-CO- LA could be representative of the broader implication that genomic defects may be correlated with the rise in osteolytic deformations within mammoths seen during the Late Pleistocene (Krezerminska et al., 2015; Lesh- chinskiy, 2015; Leshchinskiy, 2017; Rogers and Mont- gomery, 2017); however, further research on additional specimens would be required before this assertion can be made. The study also provides an insight into the life of the mammoth’s modern relative, the Asian elephant (Ele- phas maximus; Roca et al., 2015; Enk et al., 2016). Recent studies have shown a large trend in degenerative joint diseases within the Asian elephant populations (Luikart and Stover, 2005; Regnault et al., 2017). Studies into the endangered E. maximus also lead to the larger, ongoing question of a possible bottlenecking event among their ancient relatives. Studies suggest a low DNA diversity among both the modern Asian elephant and some spe- cies of mammoths including the M. primigenius (Vidya et al., 2005; Poinar et al., 2006; Vidya, 2016). Both E. maximus and mammoths have also faced similar popu- lation stressors such as habitat loss and over-predation (Saunders et al., 2010; Yansa and Adams, 2012; Gilmour et al., 2015; Vidya, 2016; Fisher, 2018). Size, Age, and Gender Estimates Podial to limb bone length ratios of UISSM-001-CO- LA (Table 2) suggest that the individual was smaller in body mass compared to the others within the Larrara- Figure 12. X-ray diffractogram of specimen UISSM-001-COLA tusk fragment. Analysis was performed at 40 kV. A total of 1561 data points were generated over a spectrum of 0 to 80°. Steps equaled 0.05° in measurements of two-theta. 39 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 mendi (2015) study. This may be due to the hypothesized age of the specimen at its time of death, corroborating that it was still a growing young adult. Specimen UISSM- 001-COLA's height estimate at approximately 3.28 m (10.76 ft) places it out of the range of adult M. primigenius heights, which did not typically exceed 3 m (9.8 ft) but lies within the height range of adult M. columbi, whose maximum heights reached about 4 m (11–13 ft) at full maturity (Lister and Bahn, 2007). M. jeffersonii is large- ly a morphological descriptor for a form that might be a hybrid from introgression or a subspecies of M. columbi. Gender differences exist in epiphyseal fusion rates, and it was pertinent to determine gender, even without a pelvis. If the mammoth were female, we would have observed many of its epiphysis fused to the diaphysis, or partially so, at the same time (Haynes, 2017). We observed many of the limb bones (proximal humerus, distal ulna and radius, and proximal femur) still un- fused, which aligns with the fusing sequence of male mammoths, not females (Haynes, 2017; Figures 8 and 9). This epiphyseal fusion is consistent with the M5 des- ignation (Figure 9). The proximal ulna and distal femur Figure 13. Maximum likelihood tree of speciemn UISSM-001-COLA’s (Erdmand_strict_Idaho_M.sp_HapF) mtDNA from 100 RAxML bootstrap replicates. Specimen UISSM-001-COLA’s mtDNA aligns strongly with Haplogroup F, which includes M. columbi, M. jeffersonii, and an unidentified mammoth. 40 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 were the key limb bones in intermediate stages of epiph- yseal fusion to examine. Specimen UISSM-001-COLA died when the proximal ulna was fully fused, and the distal femur was almost completely fused. Diagenetic Analysis SEM/EDS, XRD, and protein analysis suggest that most of the protein and other organic components have degraded in the superficial areas of the tooth and tusk. Because the tusk has a more homogenous anatomy than the tooth and given that the tusk is, currently, highly and easily fragmented, little to no recoverable organic materials remain in the tusk. The degradation of pro- teins, lipids, and genetic materials in the bones likely mirrored that of the tusk. This is evidenced by the lack of protein found by preliminary protein analysis. Fur- thermore, a femur fragment tested for isotope analysis indicated that little to no collagen existed in the sample. Due to their molecular structures, proteins and lipids can generally be analyzed on longer temporal scales than genetic material (Briggs and Summons, 2014; Buckley and Wadsworth, 2014; Cappellini et al., 2018). Thus, because proteins are not detectable in the femur at the time of this writing, most, if not all of, the ancient organic biomolecules likely vanished from the bone. The tooth, on the other hand, demonstrated more promising results of biomolecular preservation with the presence of high proportions of carbon in SEM/EDS analysis. The heterogeneity of elemental composition of the tooth may be explained by the protective barrier of enamel disrupting fossilization and degradation of the interior tooth (Trueman and Martill, 2002; Kend- all et al., 2018). During the SEM/EDS analysis, it ap- peared that the smaller particles from the tooth tended to contain more carbon, perhaps due to differences in fragmentation of the denser enamel and the more po- rous, interior dentin. Additionally, it should be noted that likely surface contaminants may occur on the exte- rior of the tooth, which contained high levels of carbon according to EDS analysis. Whereas some level of con- tamination is inevitable, and the presence of these sur- face contaminants should be considered, it is unlikely that this contamination is significantly contributing to the elemental and molecular composition results. The crystalline structures on the surface of the tusk may have arisen from one of two sources: first, these structures may be remnants of calcium carbonate from the depositional environment of UISSM-001-COLA or, second, they may represent a product of the process of efflorescence. Efflorescence occurs when carbon di- oxide reacts with water to form carbonic acid, which then combines with calcium carbonate to form calcium bicarbonate. The calcium bicarbonate, a more soluble compound than calcium carbonate, can distribute and permeate into many areas of tusk. If the water begins to evaporate, the calcium bicarbonate is deposited on the exterior surface of the tusk as calcium carbonate. The process of efflorescence is evidenced by the finding of the crystalline structures on the surface of the tusk, but not on the other remains. This likely occurred to a great- er extent in the tusk because of its porosity. The hard enamel on the surface of the tooth likely protected the teeth from calcite deposition. This finding is important to note, as the presence and absence of calcium carbon- ate crystals on the tusk and tooth, respectively, suggests that less water was able to seep into or evaporate off the tooth. Thus, the interior of the tooth is a good candidate for any future biomolecular analysis, as results further demonstrate that organic matter remains there. Taxonomic Identification Molars display different characteristics that are de- pendent on the tooth’s state of wear and the age of the mammoth, which can cause challenges in species iden- tification (Roth and Shoshani, 1988; Lister and Sher, 2015). With this said, the dimensions and characteristics of UISSM-001-COLA’s right maxillary were compared to type specimens of M. primigenius, M. columbi, and M. jeffersonii. Although UISSM-001-COLA displays a high lamellar frequency (Table 5) similar for those reported for the type specimen of M. jeffersonii and neotype of M. primigenius, the plate number is still characteristic of M. columbi. Whereas these characteristics were used to make the taxonomic assignment, the amount of overlap between the measurements on UISSM-001-COLA and the values presented for each taxon causes uncertainty (Lister, 2017; Widga et al., 2017). Haplogroup F is a clade of mammoths common to 41 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 the Great Plains and Great Basin, whereas haplogroup C is more common to the Yukon, Great Lakes, and East Coast. With that said, the differences between the mito- chondrial genomes in these regions are minor and sug- gest that they all share a common matriline (Enk et al., 2016; Widga et al., 2017). Widga et al. (2017) suggest morphological overlap between populations of Mam- muthus regionally and attribute this to varying environ- ments and population history. Specimen UISSM-001- COLA's taxonomic placement is most closely related to the Great Plains and Great Basin mammoth group (haplogroup F). It has also recently been proposed that mammoths occupying certain biogeographic ranges display vari- ability between members of the same species (Widga et al., 2017). While most mammoth occurrences in Ida- ho have not been taxonomically described according to current morphometrics, Widga et al. (2017) observed biogeographic variation between mammoth occurrenc- es in the western and eastern United States. Mammoths from the Great Plains, West Coast, and southwest have generally lower plate counts and thicker enamel than those from the Midwest and East Coast (Widga et al., 2017). A more detailed description of mammoth col- lections from Idaho, and by extension the northwestern U.S.A., is required to construct a more robust compari- son of this variation. CONCLUSION Here, we determined that UISSM-001-COLA spec- imen was most likely a male mammoth 3.28 m (10.76 ft) tall at shoulder height, young adult around 29 years old (AEY), living 13,586 to 13,444 cal BP. Specimen UISSM-001-COLA was among some of the last mam- moths to live on mainland of Western North America as mammoth populations became extremely endan- gered during the Bølling-Allerød climatic event. The mammoth died with potential lameness in one foot and potential bone disease and was preserved in an ancient terrestrial hot spring deposit. The cause of death is still undetermined. Over time, its remains underwent early stages of diagenetic alteration, and pores in the bones were infilled with recrystallized apatite and calcium carbonate. The incompleteness of the skeleton, along with the gnaw marks suggests partial disarticulation be- fore ultimate burial. Still, many of the large bones were preserved in a low-energy environment. While the driving forces of mammoth hybridization patterns are still not completely understood, they might represent interactions between distinct mammoth pop- ulations in response to the changing climate of the time. Whether this interbreeding was a direct result of cli- matic pressures remains unclear, but UISSM-001-CO- LA provides potential evidence of M. primigenius and M. columbi interaction. Extensive gene flow occurred between haplogroups C and F (Enk et al., 2016; Wid- ga et al., 2017), and UISSM-001-COLA is designated into haplogroup F, the M. jeffersonii line. Specimen UISSM-001-COLA’s death and subsequent preservation provides paleontologists with information about a key component to the ecosystem of Idaho. Further studies documenting other M. jeffersonii occurrences in West- ern North America will undoubtedly reveal more about the paleoecology and population dynamics of one of the Cenozoic’s most eponymous taxa, as well as insights into underlying causes of their eventual extinction. ACKNOWLEDGMENTS This project was funded by the Office of Under- graduate Research Grant and the Hill Undergraduate Research Fellowship from Dr. Brian and Gayle Hill at the University of Idaho. The authors would like to ac- knowledge the help of Dr. Tom Williams at the Univer- sity of Idaho for SEM/XRD expertise. Thank you to Dr. Lee Deobald at the University of Idaho Microchemistry Mass Spectrometry Laboratory and Gritman Medical Services in Moscow, Idaho, for donation of their CT scanner. Thank you to Shelby Dunn at UCSC Paleog- enomics Lab for performing the DNA analysis on the mammoth specimen. Thank you to C. Widga from East Tennessee State University for his thoughtful conversa- tions. Thank you to our reviewers for their insightful and helpful comments. REFERENCES Agenbroad, L.D., 2005, North American proboscideans—mammoths—the state of knowledge, 2003: Quaternary International, v. 126, p. 73–92.  42 Piecing Together a Prehistoric Puzzle—Regional Inferences of Micro- and Macroscopic Analyses of Possibly One of the Last Hybrid Mammoths in Mainland Western North America Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. Geology of the Intermountain West 2024 Volume 11 Aguire, E. E., 1969, Revision sistematica de los Elephantidae por su morfología y morfometría dentaria. 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S-1 Supplemental Material Inventory of the Bones Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-2 Spinose Process: Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length/width; cm) SMS001 1-4 Vertebrae (Portion) 20 SMS002 2-2 Lumbar Vertebrae 80 SMS003 2-13 Vertebrae (Portion) 20 SMS004 2-16 Sacrum 50 SMS005 1-5 Atlas 100 SMS006 2-12 Vertebrae (Portion) 10 SMS007 1-16 Vertebrae (Portion) 40 SMS008 2-17 Vertebrae (Portion) 50 SMS009 1-2 Lumbar Vertebrae 60 SMS010 1-8 Vertebrae (Portion) 20 SMS011 1-18 Vertebrae (Portion) 60 SMS012 1-11 Vertebrae (Portion) 40 SMS013 1-10 Vertebrae (Portion) 5 SMS014 2-1 Vertebrae (Portion) 15 SMS015 1-20 Vertebrae (Portion) 10 SMS016 1-19 Thoracic Vertebrae 70 SMS017 2-3 Vertebrae 55 SMS018 1-14 Thoracic Vertebrae 50 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-3 Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length/width; cm) SMS019 1-12 Vertebrae (Portion) 25 SMS020 1-15 Vertebrae (Portion) 10 SMS021 1-7 Vertebrae (Portion) 5 SMS022 2-6 Vertebrae 90 Ribs: Catalog # Scan ID (if Applicable) Bone ID % Intact Measurements (length/width; cm) R001 9-21 Rib N/A (so far) 26 / 7 R002 Rib N/A (so far) 27 / 6 R003 Rib N/A (so far) 10 / 4.5 R004 3-22 Rib N/A (so far) 16 / 7 R005 3-11 Rib N/A (so far) 14 / 9 R006 Rib N/A (so far) 13 / 8 R007 3-1 Rib N/A (so far) 16.5 / 7.5 R008 Rib N/A (so far) 11.5 / 7 R009 Rib N/A (so far) 21.5 / 5 R010 Rib N/A (so far) 12 / 5 R011 3-25 Rib N/A (so far) 15 / 6.5 R012 3-7 Rib N/A (so far) 14.5 / 7.5 R013 Rib N/A (so far) 10.5 / 4 R014 3-2 Rib N/A (so far) 25.5 / 7 R015 Rib N/A (so far) 22.5 / 6 R016 9-18 Rib N/A (so far) 11.5 / 9 R017 Rib N/A (so far) 10 / 5-7 R018 Rib N/A (so far) 7.5 / 7.5 R019 Rib N/A (so far) 12 / 6 R020 Rib N/A (so far) 9 / 3.5 R021 Rib N/A (so far) 7 / 4.5 R022 Rib N/A (so far) 17 / 4.5 R023 Rib N/A (so far) 13 / 5 R024 Rib N/A (so far) 17 / 6 R025 Rib N/A (so far) 11 / 6 R026 Rib N/A (so far) 33 / 4 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-4 Catalog # Scan ID (if Applicable) Bone ID % Intact Measurements (length/width; cm) R027 Rib N/A (so far) 17 / 3.5 R028 9-12 Rib N/A (so far) 27 / 3.5 R029 Rib N/A (so far) 14 / 3.5 R030 9-6 Rib N/A (so far) 33.5 / 4.5 R031 3-5 Rib N/A (so far) 33 / 5 R032 9-16 Rib N/A (so far) 31 / 5 R033 Rib N/A (so far) 14.5 / 3.5 R034 Rib N/A (so far) 11 / 3.5 R035 Rib N/A (so far) 26 / 4.5 R036 Rib N/A (so far) 36.5 / 5.5-4 R037 Rib N/A (so far) 29.5 / 5 R038 Rib N/A (so far) 27 / 5-4 R039 Rib N/A (so far) 17.5 / 4 R040 Rib N/A (so far) 9 / 4 R041 3-18 Rib N/A (so far) 26 / 5-4.5 R042 3-16 Rib N/A (so far) 28.5 / 5.5-4-3 R043 3-12 Rib N/A (so far) 34 / 6-5 R044 9-14 Rib N/A (so far) 13.5 / 5.5 R045 Rib N/A (so far) 19 / 4.5 R046 Rib N/A (so far) 9.5 / 5 R047 Rib N/A (so far) 19.5 / 5.5-5 R048 Rib N/A (so far) 10.5 / 4.5 R049 Rib N/A (so far) 24.5 / 4.5-4 R050 Rib N/A (so far) 14 / 5-4.5 R051 Rib N/A (so far) 11 / 4.5 R052 Rib N/A (so far) 11 / 5-4 R053 Rib N/A (so far) 9 / 4.5 R054 Rib N/A (so far) 46.5 / 3.5-3 R055 Rib N/A (so far) 55.5 / 4-3.5 R056 Rib N/A (so far) 16 / 7 R057 Rib N/A (so far) 21.5 / 8 R058 Rib N/A (so far) 23.5 / 4 R059 Rib N/A (so far) 10 /3.5 R060 Rib N/A (so far) 12 /3 R061 Rib N/A (so far) 5 / 4.5 R062 Rib N/A (so far) 9.5 / 3.5 R063 Rib N/A (so far) 9 / 2.5 R064 Rib N/A (so far) 6 / 2.5 R065 Rib N/A (so far) 7 / 3 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-5 Catalog # Scan ID (if Applicable) Bone ID % Intact Measurements (length/width; cm) R066 Rib N/A (so far) 6 / 5 R067 Rib N/A (so far) 9.5 / 4 R068 Rib N/A (so far) 7 / 5 R069 Rib N/A (so far) 9 / 3 R070 Rib N/A (so far) 5.5 / 3 R071 Rib N/A (so far) 16 / 3.5 R072 Rib N/A (so far) 13 / 5-4 R073 Rib N/A (so far) 13.5 / 3.5 R074 Rib N/A (so far) 8 / 4 R075 Rib N/A (so far) 6 / 3.5 R076 Rib N/A (so far) 6.5 / 3.5 R077 Rib N/A (so far) 11 / 3.5 R078 Rib N/A (so far) 8 / 3.5 R079 Rib N/A (so far) 11 / 3-2.5 R080 Rib N/A (so far) 13 / 3 R081 Rib N/A (so far) 8 / 2.5 R082 Rib N/A (so far) 8 / 3 R083 Rib N/A (so far) 4 / 3 R084 Rib N/A (so far) 10 / 3.5 R085 Rib N/A (so far) 7 / 2.5 R086 Rib N/A (so far) 6.5 / 2.5 R087 Rib N/A (so far) 7 / 4 R088 Rib N/A (so far) 9 / 3 R089 Rib N/A (so far) 11.5 / 3.5 R090 Rib N/A (so far) 12 / 2.5 R091 Rib N/A (so far) 8 / 4 R092 Rib N/A (so far) 6 / 2.5 R093 Rib N/A (so far) 6 / 3 R094 Rib N/A (so far) 7 / 3 R095 Rib N/A (so far) 9 / 3.5 R096 Rib N/A (so far) 6 / 1.5 R097 Rib N/A (so far) 4 / 3 R098 Rib N/A (so far) 6.5 / 4 R099 Rib N/A (so far) 15.5 / 4.5 R100 Rib N/A (so far) 14 / 3.5 R101 Rib N/A (so far) 12 / 3.5 R102 Rib N/A (so far) 22.5 / 4 R103 Rib N/A (so far) 8 / 5 R104 Rib N/A (so far) 12 / 4 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-6 Catalog # Scan ID (if Applicable) Bone ID % Intact Measurements (length/width; cm) R105 Rib N/A (so far) 24 / 3.5 R106 Rib N/A (so far) 14.5 / 3.5 R107 Rib N/A (so far) 11 / 2 R108 Rib N/A (so far) 9.5 / 3.5 R109 Rib N/A (so far) 9.5 / 3 R110 Rib N/A (so far) 8 / 2 R111 Rib N/A (so far) 5.5 / 3 R112 Rib N/A (so far) 6.5 / 4 R113 Rib N/A (so far) 15.5 / 4 R114 Rib N/A (so far) 7 / 4.5 R115 Rib N/A (so far) 16.5 / 4.5 R116 Rib N/A (so far) 10 / 4.5 R117 Rib N/A (so far) 17 / 3.5 R118 Rib N/A (so far) 9.5 / 4.5 R119 Rib N/A (so far) 11 / 4 R120 Rib N/A (so far) 10 / 3.5 R121 Rib N/A (so far) 8 / 3 R122 Rib N/A (so far) 9 / 4 R123 Rib N/A (so far) 16.5 / 5.5 R124 Rib N/A (so far) 6 / 3.5 R125 Rib N/A (so far) 11.5 / 5 R126 Rib N/A (so far) 6 / 3 R127 Rib N/A (so far) 15.5 / 4 R128 Rib N/A (so far) 7 / 2 R129 Rib N/A (so far) 10.5 / 4 R130 Rib N/A (so far) 5 / 3-2 R131 Rib N/A (so far) 11 / 4 R132 Rib N/A (so far) 6.5 / 2.5 R133 Rib N/A (so far) 9 / 4.5 R134 Rib N/A (so far) 4.5 / 3.5 R135 Rib N/A (so far) 8 / 3.5 R136 Rib N/A (so far) 6.5 / 3.5 R137 Rib N/A (so far) 10 / 4 R138 Rib N/A (so far) 10 / 3.5 R139 Rib N/A (so far) 8.5 / 2.5 R140 Rib N/A (so far) 12.5 / 3.5 R141 Rib N/A (so far) 11.5 / 4 R142 Rib N/A (so far) 13 / 3 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-7 Cranium: Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length/width; cm) SMC001 11-36 Cranium (Portion) 15 SMC002 11-45 Cranium (Portion) 15 SMC003 11-46 Cranium (Portion) 15 SMC004 11-4 Cranium (Portion) 15 SMC005 11-30 Cranium (Portion) 95 SMC006 11-24 Cranium (Portion) 95 SMC007 11-15 Cranium (Portion) 15 SMC008 11-16 Cranium (Portion) 5 SMC009 11-17 Cranium (Portion) 5 SMC010 11-18 Cranium (Portion) 2 SMC011 11-19 Cranium (Portion) 20 SMC012 11-21 Cranium (Portion) 10 SMC013 11-27 Cranium (Portion) 15 SMC014 11-41 Cranium (Portion) 5 SMC015 11-43 Cranium (Portion) 10 SMC016 11-44 Cranium (Portion) 10 SMC017 11-38 Cranium (Portion) 10 SMC018 11-40 Cranium (Portion) 10 SMC019 11-8 Cranium (Portion) 2 SMC020 11-6 Cranium (Portion) 10 SMC021 11-42 Cranium (Portion) 10 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-8 Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length/width; cm) SMC022 11-35 Cranium (Portion) 25 SMC023 11-12 Cranium (Portion) 10 SMC024 15-1 Lower Jaw/Mandible 80 76.2 / 50.8 SMC026 15-2 Right Tusk 90 228.6 SMC027 15-3 (possibly) Left Tusk 90 SMC028 15-1 Lower Left Molar (M1) 100 SMC029 15-1 Lower Right Molar (M1) 100 SMC030 Upper Left Molar 100 Front Legs: Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length; cm) SSMFL 1 7_1 Ulna 75 66.04 SSMFL 2 7_2 Humerus 90 38.10 SSMFL 3 7_3 Radius 85 63.50 SSMFL 4 7_4 Ulna 30 45.72 SSMFL 5 7_5 Ulna 30 59.63 SSMFL 6 7_6 Ulna 80 58.42 SSMFL 7 8_1 Humerus 90 90.17 SSMFL 8 8_2 Humerus 70 46.99 SSMFL 9 8_3 Radius 95 50.80 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-9 Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length; cm) SSMFL 10 8_4 Radius 45 19.05 SSMFL 11 8_5 Radius 50 22.86 SSMFL 12 8_6 Humerus 90 20.32 SSMFL 13 8_7 Radius 95 21.59 SSMFL 14 8_8 Radius 15 10.16 SSMFL 15 8_9 Radius 20 20.32 Rear Legs: Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length; cm) SMRL001 4-1 Femur(distal) 40 51 SMRL002 Femur(distal) 50 73 SMRL003 4-5 Femur 15 45 SMRL004 4-6 Femur 15 41 SMRL005 4-7 Femur 15 44.5 SMRL006 5-1 Tibia 40 30.5 SMRL007 Tibia 20 33.7 SMRL008 5-2 Tibia3 40 37 SMRL009 5-6 Tibia 4 30 34 SMRL010 4-8 Patella 1 100 15.4 SMRL011 5-4 Pelvis 1 10 35.6 SMRL012 5-3 Pelvis 2 15 33.8 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-10 Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length; cm) SMRL013 6-6 Pelvis 3 10 29.6 SMRL014 6-4 Pelvis 4 10 28.7 SMRL015 5-5 Pelvis5 10 26.8 SMRL016 6-11 Pelvis 6 10 20.3 SMRL017 6-12 Pelvis 7 5 17.7 SMRL018 6-5 Pelvis 8 5 18 SMRL019 4-2 Pelvis 9 5 20.4 SMRL020 6-9 Pelvis 10 5 14.6 SMRL021 5-7 Pelvis 11 10 28.3 SMRL022 6-13 Pelvis 12 5 23 SMRL023 6-1 Pelvis 13 5 19.5 SMRL024 6-10 Pelvis 14 5 21 SMRL025 6-7 Pelvis 15 5 20.5 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-11 Forefeet: Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements length and width; cm) SMF009 Cuneiform 80 10.5 / 9 SMF010 Magnum 50 8 / 10.5 SMF011 Unciform 96 15.5 / 10.5 SMF013 Trapezium 96 9 / 5 SMF014 Scaphoid 96 16 / 4 SMF016 14-8 Cuneiform 98 19 / 7 SMF017 14-7 Lunar 90 13.5 / 8 SMF018 14-11 Trapezium 98 9 / 4.5 SMF019 14-3 Pisiform 98 15 / 7 SMF020 14-4 Magnum 98 10.5 / 7.5 SMF021 14-6 Trapezoid 98 11 / 6 Hindfeet: Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length and width; cm) SMF001 Calcaneum 98 24 / 19 SMF002 Calcaneum 97 24 / 18 SMF003 Astragalus 65 9 / 7 SMF004 Navicular 75 13 / 4.5 SMF005 External Cuneiform 98 10.5 / 6.5 SMF006 External Cuneiform 96 11 / 7 SMF007 Internal Cuneiform 98 8 / 5 Supplemental Material: Inventory of the Bones Morrison, K., Usachenko N., Erdman, J., Waters, S., and Love, R.L. S-12 Catalog # Scan ID (If Applicable) Bone ID % Intact Measurements (length and width; cm) SMF008 Internal Cuneiform 96 7.5 / 5.5 SMF012 Astragalus 98 17.5 / 8 SMF015 Cuboid 98 14.5 / 5 SMF022 Navicular 50 11.5 / 5 Toes: Catalog # Scan ID Bone ID % Intact Measurements (length and width; cm) SMF023 14-9 Metacarpal 98 18.5 / 6.5 SMF024 14-1 Metacarpal 90 21 / 5.5 SMF025 14-2 Metacarpal 96 19 / 6 SMF026 10 -44 Metacarpal 90 20 / 6 SMF027 Metacarpal 96 19.5 / 7