Monograph |
Corresponding author: Jean-Jacques Jaeger ( jean-jaques.jaeger@univ-poitiers.fr ) Corresponding author: Somsak Panha ( somsak.pan@chula.ac.th ) Academic editor: Raquel López-Antoñanzas
© 2016 Kantapon Suraprasit, Jean-Jacques Jaeger, Yaowalak Chaimanee, Olivier Chavasseau, Chotima Yamee, Pannipa Tian, Somsak Panha.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Suraprasit K, Jaeger J-J, Chaimanee Y, Chavasseau O, Yamee C, Tian P, Panha S (2016) The Middle Pleistocene vertebrate fauna from Khok Sung (Nakhon Ratchasima, Thailand): biochronological and paleobiogeographical implications. ZooKeys 613: 1-157. https://doi.org/10.3897/zookeys.613.8309
|
The fluviatile terrace deposits of Khok Sung, Nakhon Ratchasima province, have yielded more than one thousand fossils, making this the richest Pleistocene vertebrate fauna of Thailand. The excellent preservation of the specimens allows precise characterization of the faunal composition. The mammalian fauna consists of fifteen species in thirteen genera, including a primate, a canid, a hyaenid, proboscideans, rhinoceroses, a suid, cervids, and bovids. Most species correspond to living taxa but globally (Stegodon cf. orientalis) and locally (Crocuta crocuta ultima, Rhinoceros unicornis, Sus barbatus, and Axis axis) extinct taxa were also present. The identification of Axis axis in Khok Sung, a chital currently restricted to the Indian Subcontinent, represents the first record of the species in Southeast Asia. Three reptilian taxa: Crocodylus cf. siamensis, Python sp., and Varanus sp., are also identified. Faunal correlations with other Southeast Asian sites suggest a late Middle to early Late Pleistocene age for the Khok Sung assemblage. However, the Khok Sung mammalian fauna is most similar to that of Thum Wiman Nakin, dated to older than 169 ka. The Khok Sung large mammal assemblage mostly comprises mainland Southeast Asian taxa that migrated to Java during the latest Middle Pleistocene, supporting the hypothesis that Thailand was a biogeographic pathway for the Sino-Malayan migration event from South China to Java.
Large mammals, taxonomy, Ailuropoda–Stegodon assemblage, paleobiogeography, late Middle Pleistocene, Quaternary, northeastern Thailand, mainland Southeast Asia
In the Pleistocene, mammalian faunas in mainland Southeast Asia as well as in South China are characterized by the occurrence of Ailuropoda (giant panda) and/or Stegodon (extinct proboscidean), also called “Ailuropoda–Stegodon faunal complex”. This faunal association is a characteristic of the long period ranging from the Early to Late Pleistocene (
Map of Southeast Asia showing A the Sundaland boundaries and the migration route hypothesis: Siva-Malayan route (black), Sino-Malayan route (red), and Taiwan-Philippine Archipelago route (blue) and B the location of the Khok Sung sand pit (star) and other Middle (red circle) and Late (yellow circle) Pleistocene sites. The Sunda shelf boundaries at the sea level about 120 m lower than the present day are compiled from
Dating from the Middle to Late Pleistocene, there are numerous paleontological and archaeological sites with mammalian fossil faunas discovered in Southeast Asian mainland (Indochinese) and islands (Sundaic) (Fig.
An ongoing survey of Pleistocene deposits in Thailand has led to the discovery of numerous mammalian fossils by the Thai-French paleontological team in limestone caves from the northern to southern part of the country. Several fissure-filling and cave deposits: Thum Wiman Nakin (
In 2005, the Khok Sung sand pit (Nakhon Ratchasima province, northeastern Thailand) was excavated (Fig.
The sand pit was open for the pond construction (approximately 50 m long × 50 m wide × 8 m deep) (Fig.
All fossil specimens are housed at the Khok Sung local museum (Nakhon Ratchasima) and at the
iPHEP
Institut International de Paléoprimatologie et de Paléontologie Humaine: Evolution et Paléoenvironnements,
NHMP
THNHM-M
Mammal collection, Thailand
The dental nomenclature follows
All specimens were measured using digital callipers to the nearest 0.01 mm. The tooth dimensions for all mammals were measured at the base of the crown along the anterior-posterior margins for the maximum length (L) and from the labial (incisors and canines)/buccal (premolars/molars) to lingual margins for the maximum width (W). In the case of measurements of stegodontid cheek teeth, the methods and parameters used for molar and ridge dimensions were given in Fig.
Parameters and measurement methods used for the lower third molar of Stegodon. Lengths and widths of the molar ridges are abbreviated as “LR” and “WR”, respectively. An illustration of two right m3 (lateral and occlusal views) of Stegodon orientalis is duplicated from the specimen
Scapula | |
HS | Height along the spine |
DHA | Diagonal height from the most distal point of the scapula to the thoracic angle |
Ld | Greatest dorsal length |
SLC | Smallest length of the Collum scapulae (neck of the scapula) |
GLP | Greatest length of the Processus articularis (glenoid process) |
LG | Length of the glenoid cavity |
BG | Breadth of the glenoid cavity |
Long bones | |
GL | Greatest length |
GLl | Greatest length of the lateral part |
GLC | Greatest length from the caput (head) |
PL | Physiological length (for radius only) |
Ll | Length of the lateral part |
Bp | Greatest breadth of the proximal end |
BFp | Greatest breadth of the Facies articularis proximalis (for radius only) |
BPC | Greatest breadth across the coronoid process (=greatest breadth of the proximal articular surface) (for ulna only) |
SD | Smallest breadth of diaphysis |
Dp | Depth of the proximal end |
Bd | Greatest breadth of the distal end |
BFd | Greatest breadth of the Facies articularis distalis (for radius only) |
Dd | Greatest breadth of the distal end |
DC | Greatest depth of the Caput femoris |
DD | Smallest depth of the diaphysis (for metapodials only) |
BT | Greatest breadth of the trochlea (for humerus only) |
LO | Length of the olecranon (for ulna only) |
DPA | Depth across the Processus anconaeus (for ulna only) |
SDO | Smallest depth of the olecranon (for ulna only) |
Pelvis | |
GL | Greatest length of one half |
LA | Length of the acetabulum including the lip |
LS | Length of the symphysis |
SH | Smallest height of the shaft of ilium |
SB | Smallest breadth of the shaft of ilium |
SC | Smallest circumference of the shaft of ilium |
LFo | Inner length of the foramen obturatum |
GBTc | Greatest breadth across the Tubera coxarum–greatest breadth across the lateral angle |
GBA | Greatest breadth across the acetabula |
GBTi | Greatest breadth across the Tubera ischiadica |
SBI | Smallest breadth across the bodies of the Ischia |
The body mass of ruminants was estimated using the equations of
We compared differences in species compostion of Southeast Asian large mammal fauna during the Middle Pleistocene, using an analysis of the faunal similarity. According to unequal sampling conditions for our data, we applied two criteria for undertaking this analysis: localities are disqualified when they have fewer than 10 taxa identified at the species level and taxa are excluded when their appearances are still doubtful (i.e. poor taxonomic description or identification). We therefore selected Simpson’s Faunal Resemblance Index (FRI) because it has the smallest influence of sample sizes and emphasizes faunal resemblances (
A right tibia, DMR-KS-05-04-04-1.
The right tibia is complete (Fig.
Postcranial remains of Macaca sp. A–D and Cuon sp. E–H from Khok Sung: A–D DMR-KS-05-04-04-1, a right tibia in anterior (A), medial (B), proximal (C), and distal (D) views E–F DMR-KS-05-04-11-34, a right ulna in medial (E) and anterior (F) views G–H DMR-KS-05-04-28-13, a right femur in proximal (G) and posterior (H) views.
Tibial morphology is relatively conservative within and among primates. Particularly, the morphological differences of tibiae among cercopithecoids are minimal (
Ratios of the greatest lengths of tibiae (GL) to the lengths and widths of proximal and distal tibiae (Bp, Dp, Bd, and Dd) of Khok Sung macaques compared to recent Southeast Asian primates.
DMR-KS-05-04-04-1 | Presbytis (N = 30) | Hylobates (N = 24) | Macaca (N = 71) | |||||||
---|---|---|---|---|---|---|---|---|---|---|
Max | Min | Mean | Max | Min | Mean | Max | Min | Mean | ||
GL/Bp | 6.09 | 7.70 | 6.76 | 7.29 | 7.52 | 6.06 | 7.01 | 6.56 | 4.55 | 5.61 |
GL/Dp | 7.81 | 9.89 | 8.15 | 9.07 | 9.95 | 7.96 | 9.43 | 9.62 | 6.36 | 7.67 |
GL/Bd | 9.25 | 12.38 | 10.26 | 11.37 | 14.49 | 9.01 | 11.31 | 10.84 | 7.20 | 8.79 |
GL/Dd | 12.94 | 16.21 | 12.75 | 14.13 | 16.79 | 10.94 | 14.50 | 12.77 | 7.69 | 10.78 |
A right ulna, DMR-KS-05-04-11-34; a right femur, DMR-KS-05-04-28-13.
DMR-KS-05-04-11-34 is a half proximal ulna preserving complete parts from the olecranon to the midshaft (Fig.
The right femur preserves a complete proximal part and broken shaft (Fig.
The proximal ulna of canids is characterized by a bilobed and laterally compressed olecranon process, well-developed anconeal and lateral coronoid processes, and a laterally compressed shaft. The proximal crest of the olecranon is grooved anteriorly, but enlarged and rounded posteriorly (
Measurements (in millimetres) of ulnae and femurs of Khok Sung and other extant and fossil canids. * indicates a subadult individual. Metrical data of fossil canids are from
Ulna | |||||||
Specimen no. | Taxa | Age | Locality | LO | DPA | SDO | BPC |
DMR-KS-05-04-11-34 | Cuon sp. | late Middle Pleistocene | Khok Sung, northeastern Thailand | 15.16 | 18.51 | 15.21 | 11.65 |
|
Canis lupus | Recent | Eastern India | 29.91 | 24.11 | 18.38 | 15.65 |
29.29 | 24.43 | 18.43 | 15.33 | ||||
|
Canis lupus | Late Pleistocene | Geographical Society Cave, Russia | – | 32.30 | 27.80 | – |
NHMP R5387 | Canis lupus | Late Pleistocene | Srbsko Chlum-Komín Cave, Czech Republic | – | 34.80 | 27.60 | – |
|
Cuon alpinus | Recent | Java, Indonesia | 19.23 | 19.37 | 16.36 | 14.43 |
19.74 | 19.29 | 16.33 | 14.07 | ||||
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 1 Cave, Southern Ossetia, Caucasus | – | – | – | 18.30 |
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 1 Cave, Southern Ossetia, Caucasus | – | 32.20 | – | 17.20 |
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 3 Cave, Southern Ossetia, Caucasus | – | 28.70 | 24.50 | 18.90 |
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 3 Cave, Southern Ossetia, Caucasus | – | 34.00 | 29.50 | 21.50 |
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 3 Cave, Southern Ossetia, Caucasus | – | 33.60 | 28.60 | 21.70 |
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 3 Cave, Southern Ossetia, Caucasus | – | 30.30 | 26.50 | 17.00 |
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 3 Cave, Southern Ossetia, Caucasus | – | 28.80 | – | 18.50 |
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 3 Cave, Southern Ossetia, Caucasus | – | – | – | 15.00 |
Femur | |||||||
Specimen no. | Taxa | Age | Locality | Bp | Dp | DC | SD |
DMR-KS-05-04-28-13 | Cuon sp. | late Middle Pleistocene | Khok Sung, northeastern Thailand | 35.69 | 17.90 | 16.58 | 11.34 |
|
Canis lupus | Recent | Eastern India | 35.05 | 16.70 | 16.75 | 10.43 |
35.57 | 16.82 | 16.73 | 10.52 | ||||
|
Cuon alpinus | Recent | Java, Indonesia | 31.03 | 15.95 | 16.62 | 11.66 |
31.58 | 16.08 | 16.38 | 11.79 | ||||
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 3 Cave, Southern Ossetia, Caucasus | 48.70 | – | 22.70 | – |
|
Cuon alpinus caucasicus | Late Pleistocene | Kudaro 3 Cave, Southern Ossetia, Caucasus | – | – | 21.70 | 15.20 |
Living canids generally show a typical morphology of the proximal femur, characterized by their relatively vertical intertrochanteric crests, prominent lesser trochanter with the sharp crest extending downward along the shaft, moderately-sized greater trochanter, and slender shaft (
Because the Khok Sung ulna and femur morphologically match better Cuon alpinus than Canis lupus, we identify these two postcranial specimens as belonging to Cuon sp.
A right DP4 (posterior part), DMR-KS-05-03-28-14; a left DP4 (anterior part), DMR-KS-05-03-19-7; a left M2, DMR-KS-05-03-29-1 (posterior part); a right M3, DMR-KS-05-03-22-19 (posterior part); a fragmentary tusk, DMR-KS-05-03-15-2; a left dp3 (anterior part), DMR-KS-05-04-01-8; two mandibles with m3—DMR-KS-05-03-08-1 (right) and DMR-KS-05-03-08-2 (left); a right humerus fragment (proximal part), DMR-KS-05-03-10-5; a left humerus, DMR-KS-05-03-10-6; two ulna fragments (proximal parts)—DMR-KS-05-03-09-7 and DMR-KS-05-03-10-2; a femoral head fragment, DMR-KS-05-03-10-3; a right femur, DMR-KS-05-03-10-4; a right tibia fragment (distal part), DMR-KS-05-03-10-3; a right fibula, DMR-KS-05-03-00-124; two pelvis fragments—DMR-KS-05-03-10-11 (right) and DMR-KS-05-03-10-12 (left); five vertebrae—DMR-KS-05-03-17-11, DMR-KS-05-03-10-7, DMR-KS-05-03-09-18, DMR-KS-05-03-10-1, and DMR-KS-05-03-28-20; a sacrum fragment, DMR-KS-05-03-10-8; two ribs—DMR-KS-05-03-10-13 and DMR-KS-05-03-10-14; three rib fragments—DMR-KS-05-03-09-6 (body), DMR-KS-05-03-09-45 (body), and DMR-KS-05-03-09-4 (head and neck).
Upper dentition: both fragments of DP4 (DMR-KS-05-03-28-14: Fig.
Dental remains of Stegodon cf. orientalis from Khok Sung: A–B DMR-KS-05-03-28-14, a right DP4 in occlusal (A) and buccal (B) views C DMR-KS-05-03-19-7, an anterior lobe of DP4 in occlusal view D DMR-KS-05-04-01-8, a left dp3 in occlusal view E–F DMR-KS-05-03-29-1, a left posterior fragment of M2 in occlusal (E) and buccal (F) views G–H DMR-KS-05-03-22-19, a right posterior fragment of M3 in occlusal (G) and buccal (H) views I–K DMR-KS-05-03-15-2, a fragmentary upper tusk in proximal (I–J) and dorsal (K) views L DMR-KS-05-03-08-1, a left mandible with m3 in occlusal view M DMR-KS-05-03-08-2, a right mandible with m3 in occlusal view.
Measurements (in millimeters) of cheek teeth of Khok Sung proboscideans, including a number of preserved ridges (NR), lengths (L), widths (W), heights (H), enamel thickness (ET), H/W indices (100 × H/W), and laminar frequencies (LF). The laminar frequencies are expressed as the following formula: LF=n*100/dl + n*100/db / 2, where “dl” and “db” are referred to distances at the lingual and buccal side of the tooth, respectively, and “n” is equivalent to the number of ridges between two measuring points (
Specimen no. | NR | L | W | H | ET | H/W index | LF | |
---|---|---|---|---|---|---|---|---|
Stegodon cf. orientalis | ||||||||
DMR-KS-05-03-28-14 | DP4 | 4 | 60.08 | 50.04 | 26.71 | 0.69–1.21 | 53.38–58.23 | 7.99 |
DMR-KS-05-03-19-7 | DP4 | 1 | 18.65 | 49.89* | 26.71 | 2.06 | 53.53 | – |
DMR-KS-05-03-29-1 | M2 | 3 | 70.14* | 78.83 | 55.18 | 1.62–3.06 | 70.00–73.34 | 4.61 |
DMR-KS-05-03-22-19 | M3 | 3 | 90.43* | 84.66 | 46.14 | 3.77–4.35 | 57.33–62.71 | 3.86 |
DMR-KS-05-04-01-8 | dp3 | 3 | 26.68* | 26.09 | 12.08 | 1.82 | 46.30–47.76 | 10.41 |
DMR-KS-05-03-08-1 | m3 | 8 | 245.86* | 95.66 | 41.50 | 3.41–6.87 | 43.38–51.77 | 3.91 |
DMR-KS-05-03-08-2 | m3 | 8 | 247.78* | 95.57 | 42.56 | 3.39–6.54 | 44.53–52.21 | 3.94 |
Elephas sp. | ||||||||
DMR-KS-05-03-17-12 | Lower molar | 2 | 41.04* | 66.77* | 108.94 | 2.48–3.30 | 163.16–165.18 | 10.61 |
DMR-KS-05-03-29-1 (M2) preserves three posterior ridges with a small cingulum (Fig.
DMR-KS-05-03-22-19 (M3) preserves only three posterior ridges with a cingulum (Fig.
A fragmentary tusk (DMR-KS-05-03-15-2) contains dentine (outer and inner layers), cementum, and a pulp cavity (Fig.
Lower dentition: DMR-KS-05-04-01-8 (dp3) is heavily worn and comprises three preserved ridges and an anterior cingulum (Fig.
Two hemi-mandibles of the same individual (DMR-KS-05-03-08-1 and DMR-KS-05-03-08-2) are moderately well-preserved (Tab.
Postcranial remains: postcranial elements include two humeri (Fig.
Postcranial remains of Stegodon cf. orientalis from Khok Sung: A–B DMR-KS-05-03-10-6, a left distal humerus in anterior (A) and distal (B) views C–D DMR-KS-05-03-10-4, a right femur posterior (C) and distal (D) views E DMR-KS-05-03-00-124, a right fibula in posterior view F DMR-KS-05-03-10-11, a right pelvis in dorsal view G DMR-KS-05-03-10-12, a left pelvis in lateral view H DMR-KS-05-03-09-18 and I DMR-KS-05-03-10-7, vertebrae in anterior view J DMR-KS-05-03-10-8, a sacrum in ventral view K DMR-KS-05-03-10-14 and L DMR-KS-05-03-10-13, ribs in anterior view.
The proboscidean cheek teeth from Khok Sung are assigned to Stegodon because there are more than five ridges or loph(id)s on molars, V-shaped valleys between ridges on molars , and step-like worn surface reliefs on the enamel layer (
The morphologies and ridge sizes of upper molars from Khok Sung are congruent with Chinese S. orientalis (Tabs
Ridge dimensions (lengths and widths in millimeters) of upper fourth deciduous premolars between Khok Sung Stegodon and Stegodon orientalis.
DP4 | Ridge (from anterior to posterior) | |||||
---|---|---|---|---|---|---|
1st | 2nd | 3rd | 4th | 5th | 6th | |
Stegodon cf. orientalis (Khok Sung) | ||||||
Length | 15.7 | – | > 10.7 | 12.4 | 13.5 | 13.5 |
Width | 49.9 | – | 49.9 | 50.0 | 49.7 | 48.4 |
Specimen measurements: DMR-KS-05-03-28-14 and DMR-KS-05-03-19-7 | ||||||
Stegodon orientalis (×6×) | ||||||
N | 3 | 3 | 3 | 3 | 3 | 3 |
Length | 12.3–16.2 | 15.3–19.7 | 14.3–20.4 | 13.3–18.4 | 12.6–16.2 | 11.1–16.5 |
Mean | 14.1 | 17.0 | 17.4 | 16.1 | 15.0 | 13.6 |
N | 3 | 3 | 2 | 3 | 3 | 3 |
Width | 43.7–54.1 | 49.2–63.1 | 51.8–63.3 | 51.2–60.2 | 50.0–57.2 | 45.8–52.2 |
Mean | 49.0 | 54.6 | 57.5 | 54.4 | 53.0 | 48.9 |
Specimen measurements: |
Ridge dimensions (lengths and widths in millimeters) of lower third deciduous premolars between Khok Sung Stegodon and Stegodon orientalis.
dp3 | Ridge (from anterior to posterior) | |||||
1st | 2nd | 3rd | 4th | 5th | 6th | |
Stegodon cf. orientalis (Khok Sung) | ||||||
Length | 9.3 | – | – | – | – | – |
Width | 25.5 | > 26.1 | – | – | – | – |
Stegodon orientalis (×5×) | ||||||
N | 7 | 7 | 7 | 7 | 7 | – |
Length | 7.2–10.8 | 6.5–10.3 | 9.6–12.9 | 10.9–12.0 | 10.0–14.0 | – |
Mean | 8.5 | 9.0 | 11.0 | 11.5 | 12.6 | – |
N | 7 | 7 | 7 | 7 | 7 | – |
Width | 19.5–32.3 | 24.8–27.9 | 27.3–31.9 | 32.6–37.8 | 36.2–42.3 | – |
Mean | 25.0 | 26.6 | 29.9 | 34.9 | 39.2 | – |
Specimen measurements: |
||||||
Stegodon orientalis (×6×) | ||||||
N | 5 | 5 | 5 | 5 | 5 | 5 |
Length | 8.6–13.1 | 7.0–11.8 | 10.1–12.8 | 10.6–13.0 | 10.6–13.4 | 8.5–12.5 |
Mean | 10.5 | 8.6 | 11.5 | 11.7 | 11.7 | 10.0 |
N | 4 | 5 | 5 | 5 | 5 | 5 |
Width | 23.7–31.1 | 26.8–32.1 | 29.1–34.7 | 33.1–41.1 | 36.7–47.3 | 36.0–52.4 |
Mean | 27.3 | 28.9 | 31.5 | 36.8 | 41.3 | 40.4 |
Specimen measurements: IVPP1799, |
Ridge dimensions (lengths and widths in millimeters) of upper second and third molars between Khok Sung Stegodon and Stegodon orientalis. The total ridge number of upper molars of Khok Sung stegodontids used for our comparisons follows that of Stegodon orientalis.
M2 and M3 | Ridge (from anterior to posterior) | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|
1st | 2nd | 3rd | 4th | 5th | 6th | 7th | 8th | 9th | 10th | Posterior halfridge | |
Stegodon cf. orientalis (Khok Sung) | |||||||||||
DMR-KS-05-03-29-1 (M2) | |||||||||||
Length | – | – | – | – | – | 28.2 | 23.9 | 19.7 | – | – | – |
Width | – | – | – | – | – | 78.8 | 76.9 | 63.3 | – | – | – |
DMR-KS-05-03-22-19 (M3) | |||||||||||
Length | – | – | – | – | – | – | – | 29.3 | 24.2 | 21.8 | 12.9 |
Width | – | – | – | – | – | – | – | 80.5 | 77.6 | 70.1 | 49.7 |
Stegodon orientalis (M2) (×8×) | |||||||||||
N | 1 | 1 | 1 | 1 | 1 | 2 | 2 | 2 | – | – | – |
Length | 23.4 | 25.0 | 30.7 | 25.4 | 22.1 | 20.3–22.5 | 20.5–22.0 | 15.4–17.7 | – | – | – |
Mean | – | – | – | – | – | 21.4 | 21.2 | 16.6 | – | – | – |
N | 1 | 1 | 1 | 1 | 1 | 2 | 2 | 2 | – | – | – |
Width | 77.8 | 80.4 | 83.1 | 83.0 | 81.6 | 76.4–78.4 | 73.0–73.8 | 63.2–69.1 | – | – | – |
Mean | – | – | – | – | – | 77.4 | 73.4 | 66.1 | – | – | – |
Specimen measurements: |
|||||||||||
Stegodon orientalis (M3) (×10×) | |||||||||||
N | 3 | 3 | 3 | 2 | 1 | 1 | 1 | 2 | 2 | 2 | 2 |
Length | 22.4–25.3 | 22.2–27.4 | 22.3–27.1 | 24.6–24.8 | 25.5 | 22.5 | 21.1 | 19.9–26.3 | 17.8–24.1 | 16.4–22.4 | 7.2–15.8 |
Mean | 23.7 | 25.6 | 24.5 | 24.7 | – | – | – | 23.1 | 20.9 | 19.4 | 11.5 |
N | 2 | 2 | 2 | 1 | – | 2 | 2 | 2 | 2 | 2 | 2 |
Width | 85.4–91.5 | 88.3–96.7 | 84.9–101.3 | 100.4 | – | 81.4–87.1 | 83.3–85.7 | 75.0–87.9 | 65.4–89.5 | 57.4–81.8 | 38.6–54.6 |
Mean | 88.4 | 92.5 | 93.1 | – | – | 84.3 | 84.5 | 81.4 | 77.5 | 69.6 | 46.6 |
Specimen measurements: |
Ridge dimensions (lengths and widths in millimeters) of lower third molars of derived Stegodon in Southeast Asia. The ridge formula of each taxon follows
Lower third molar | Ridge (from anterior to posterior) | ||||||||||||
1st | 2nd | 3rd | 4th | 5th | 6th | 7th | 8th | 9th | 10th | 11th | |||
Stegodon cf. orientalis (Khok Sung) ((×?2)9×) | |||||||||||||
N | – | – | – | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 | ||
Length | – | – | – | 29.8–32.4 | 28.8–30.6 | 31.8–32.8 | 28.2–34.1 | 28.9–32.3 | 24.5–30.7 | 23.3–26.9 | 16.6–22.4 | ||
Mean | – | – | – | 31.1 | 29.7 | 32.3 | 31.2 | 30.6 | 27.6 | 25.1 | 19.5 | ||
N | – | – | – | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 | ||
Width | – | – | – | 95.7–97.6 | 94.4–95.8 | 92.7–94.0 | 83.1–83.3 | 72.3–76.3 | 67.7–68.9 | 61.6–65.9 | 55.4–58.6 | ||
Mean | – | – | – | 96.7 | 95.1 | 93.4 | 83.2 | 74.3 | 68.3 | 63.8 | 57.0 | ||
Stegodon cf. orientalis (×11×) | |||||||||||||
N | – | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 | ||
Length | – | 26.0–31.7 | 26.4–33.8 | 23.3–34.4 | 25.5–33.1 | 28.6–31.7 | 24.8–39.4 | 26.8–36.4 | 25.6–31.0 | 21.1–24.3 | 15.4–15.9 | ||
Mean | – | 28.9 | 30.1 | 28.9 | 29.3 | 30.2 | 32.1 | 31.6 | 31.3 | 22.7 | 15.7 | ||
N | – | 1 | 2 | 1 | 1 | 2 | 2 | 2 | 2 | 2 | 2 | ||
Width | – | 82.43 | 68.0–86.1 | 88.34 | 90.11 | 72.4–93.1 | 72.0–91.9 | 71.2–88.3 | 64.6–80.1 | 54.8–63.6 | 41.8–43.1 | ||
Mean | – | – | 77.1 | – | – | 82.8 | 82.0 | 79.8 | 72.4 | 59.2 | 42.5 | ||
Specimen measurements: |
|||||||||||||
1st | 2nd | 3rd | 4th | 5th | 6th | 7th | 8th | 9th | 10th | 11th | 12th | ||
Stegodon insignis (×12×) | |||||||||||||
N | – | 1 | 2 | 3 | 2 | 2 | 2 | 3 | 3 | 2 | 2 | 2 | |
Length | – | 27.7 | 26.4–29.2 | 20.7–23.7 | 24.2 | 21.6–23.8 | 20.9–22.8 | 23.0–25.6 | 23.4–30.9 | 22.5–25.3 | 21.5–23.5 | 19.5–23.7 | |
Mean | – | – | 27.8 | 22.1 | 24.2 | 22.7 | 21.9 | 24.6 | 26 | 23.9 | 22.5 | 21.6 | |
N | – | 1 | 3 | 3 | 2 | 2 | 2 | 3 | 2 | 2 | 2 | 2 | |
Width | – | 79.3 | 83.9–92.6 | 81.7–91.2 | 92.7–94.8 | 90.7–98.5 | 89.5–89.9 | 84.6–88.0 | 73.7–77.7 | 66.6–68.8 | 61.6–64.0 | 47.1–52.5 | |
Mean | – | – | 87.4 | 88.0 | 93.8 | 94.6 | 89.7 | 85.9 | 75.7 | 67.7 | 62.8 | 49.8 | |
Specimen measurements: |
|||||||||||||
Stegodon orientalis (×12×) | |||||||||||||
N | 4 | 1 | 4 | 4 | 4 | 4 | 4 | 4 | 5 | 8 | 8 | 8 | |
Length | 17.4–25.1 | 20.4 | 25.5–31.7 | 25.9–33.8 | 23.3–34.42 | 25.5–35.0 | 28.6–35.3 | 24.8–39.4 | 26.8–36.4 | 25.6–37.0 | 16.2–31.5 | 13.4–22.6 | |
Mean | 21.1 | – | 28.0 | 28.8 | 30.3 | 30.3 | 31.1 | 32.3 | 31.1 | 29.4 | 23.0 | 16.8 | |
N | 4 | 1 | 2 | 3 | 3 | 3 | 4 | 4 | 4 | 6 | 8 | 8 | |
Width | 71.6–81.0 | 75.7 | 81.2–82.4 | 68.0–86.1 | 85.6–88.3 | 84.9–90.1 | 72.4–93.1 | 72.0–91.9 | 71.2–88.3 | 64.6–82.7 | 54.8–78.3 | 28.7–59.1 | |
Mean | 74.7 | – | 81.8 | 79.5 | 87.1 | 87.6 | 84.7 | 84.2 | 82.2 | 75.2 | 66.0 | 46.0 | |
Specimen measurements: |
|||||||||||||
1st | 2nd | 3rd | 4th | 5th | 6th | 7th | 8th | 9th | 10th | 11th | 12th | 13th | |
Stegodon trigonocephalus trigonocephalus (×13×) | |||||||||||||
N | – | – | – | 1 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | – |
Length | – | – | – | 16.8 | 20.4–25.0 | 22.1–26.0 | 24.4–26.0 | 23.7–24.9 | 21.4–24.5 | 21.1–24.0 | 16.6–24.2 | 18.4–19.4 | – |
Mean | – | – | – | – | 23.4 | 24.0 | 25.3 | 24.1 | 22.9 | 22.5 | 21.1 | 19.0 | – |
N | – | – | – | 1 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | – |
Width | – | – | – | 71.8 | 71.4–87.3 | 71.4–86.8 | 75.0–87.1 | 76.8–83.3 | 72.6–81.1 | 70.5–76.4 | 66.6–69.6 | 53.9–63.1 | – |
Mean | – | – | – | – | 80.2 | 80.4 | 80.8 | 80.4 | 77.0 | 72.6 | 68.0 | 58.4 | – |
Specimen measurements: |
|||||||||||||
1st | 2nd | 3rd | 4th | 5th | 6th | 7th | 8th | 9th | 10th | 11th | 12th | 13th | |
Stegodon florensis (×13×) | |||||||||||||
N | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 1 | 1 | 1 | 1 | – | – |
Length | 23.1–25.8 | 20.2–20.9 | 19.2–21.6 | 18.4–23.8 | 17.6–22.7 | 18.9–20.9 | 18.3–20.3 | 21.16 | 21.83 | 26.48 | 26.97 | – | – |
Mean | 24.5 | 20.6 | 20.4 | 21.1 | 20.2 | 19.9 | 19.3 | – | – | – | – | – | – |
N | 1 | 2 | 2 | 2 | 2 | 2 | 2 | 1 | 1 | 1 | 1 | – | – |
Width | 63.1 | 66.0 | 67.2–68.4 | 69.3–69.9 | 69.0–69.9 | 67.5–69.9 | 68.3–68.6 | 66.95 | 60.47 | 65.75 | 58.71 | – | – |
Mean | – | 66.0 | 67.8 | 69.6 | 69.5 | 68.7 | 68.5 | – | – | – | – | – | |
Specimen measurements: |
A fragmentary tusk, DMR-KS-05-03-22-1; a posterior fragment of a right lower molar, DMR-KS-05-03-17-12.
Upper tusk: DMR-KS-05-03-22-1 is a short fragmentary tusk. The dorsal side is partially broken away (Fig.
Lower molar: DMR-KS-05-03-17-12 preserves only two adjoining worn plates of a high-crowned molar, distinctly more hypsodont than that of Stegodon (Tab.
The fragmentary tusk (DMR-KS-05-03-22-1) is distinguished from DMR-KS-05-03-15-2 (S. orientalis) by a more rounded cross-section, a larger diameter, and a radiate fracture pattern with the development of concentric incremental lines (Fig.
A left P2, DMR-KS-05-03-00-128; a left P3, DMR-KS-05-03-22-17; a left M1, DMR-KS-05-03-00-129; a left M3, DMR-KS-05-03-00-127; a mandible with right (i2 and p2–m3) and left (p3–m3) tooth rows, DMR-KS-05-03-00-126; a partial mandible, DMR-KS-05-03-31-28; a fragmentary nasal bone, DMR-KS-05-03-00-56; a left scapula, DMR-KS-05-03-00-58; a left humerus, DMR-KS-05-03-31-3; a right metacarpus II, DMR-KS-05-03-28-29; a metacarpus III, DMR-KS-05-03-22-49; a right metacarpus IV, DMR-KS-05-04-05-15; a left tibia, DMR-KS-05-03-00-52; a right calcaneus, DMR-KS-05-04-27-19; a left astragalus, DMR-KS-05-03-26-23.
Upper dentition: P2 (DMR-KS-05-03-00-128: Fig.
Cranial, mandibular, dental remains of Rhinoceros sondaicus from Khok Sung: ADMR-05-03-00-128, a left P2 in occlusal view B DMR-KS-05-03-22-17, a left P3 in occlusal view C DMR-KS-05-03-00-129, a left M1 in occlusal view D DMR-KS-05-03-00-127, a left M3 in occlusal view E–G DMR-KS-05-03-00-126, a mandible in lateral (E), occlusal (F) and ventral (G) views H–I DMR-KS-05-03-31-28, a fragmentary mandible in occlusal (H) and lateral (I) views J–K DMR-KS-05-03-00-56, a nasal in dorsal (J) and lateral (K) views.
Measurements (in millimeters) of cheek teeth of Khok Sung rhinoceroses, Rhinoceros sondaicus and Rhinoceros unicornis, compared to recent specimens (data from
Rhinoceros sondaicus | Rhinoceros unicornis | ||||||
---|---|---|---|---|---|---|---|
Khok Sung | Recent | Khok Sung | Recent | ||||
Anterior | Posterior | Range | Anterior | Posterior | Range | ||
Upper cheek teeth | |||||||
P2 | L | 35.57 (i) | 30–38.5 | – | 37–45.5 | ||
W | 42.34 (i) | 41.24 (i) | 34.5–44 | – | – | 43–48 | |
P3 | L | 42.00 (i) | 36.5–50 | – | 43–50 | ||
W | 55.36 (i) | 53.70 (i) | 42–55 | – | – | 55.5–60.5 | |
P4 | L | – | 41–47.5 | – | 42–51 | ||
W | – | – | 52–59 | – | – | 59–69.5 | |
M1 | L | 51.38 (i) | 46–51 | 47.95 (i) | 48–58 | ||
W | 63.53 (i) | 58.67 (i) | 52.5–60 | 70.48 (i) | 58.80 (i) | 62–72.5 | |
M2 | L | – | 44.5–55 | – | 53–62 | ||
W | – | – | 53–62.5 | – | – | 64.5–76 | |
M3 | L | 55.65 (i) | 44.5–61.5 | – | 59–65 | ||
W | 55.92 (i) | 43.5–57 | – | – | 56–68.5 | ||
Lower cheek teeth | |||||||
p2 | L | – | 25–29.5 | > 30.80 (i) | 31–32 | ||
W | – | – | 15.5–21 | 18.15 (i) | 22.39 (i) | 21.5–24.5 | |
p3 | L | 42.83 (m) | 33–39 | 40.24 (m) | 38–42 | ||
W | 26.58 (m) | 29.92 (m) | 22–27.5 | – | – | 27–32 | |
p4 | L | 43.03 (m) | 36.5–42.5 | 48.13 (m) | 41–46 | ||
W | 27.71 (m) | 33.42 (m) | 24–29 | – | – | 29–34 | |
m1 | L | 41.45 (m) | 41–46.5 | 42.57 (m) | 46–48 | ||
W | 28.8–29.67 (m) | 30.88 (m) | 26–32 | – | – | 28–32.5 | |
m2 | L | 44.83–48.87 (m) | 40.5–51 | 50.74 (m) | 52–56.5 | ||
W | 29.65 (m) | 30.78–31.79 (m) | 27–32.5 | – | – | 31–36 | |
m3 | L | 54.90 (m) | 41–53 | 55.48 (m) | 49.5–60 | ||
W | 32.54 (m) | 25.11* (m) | 24.5–29.5 | – | – | 29–35 | |
Lower tooth rows | |||||||
DMR–KS–05–03–00–126 | Recent | DMR-KS–05–03–17–13 | Recent | ||||
Molar row length | 133 (right) | 126.5–147 | 158 | 147.5–161 | |||
Tooth row length | > 238 | 211.5–257 | – | 242–276 |
Measurements (lengths and widths in millimeters) of cheek teeth of Khok Sung Sus barbatus compared to the recent and fossil species. The number of specimens is given within the parentheses. The measured specimens of recent Sus scrofa include three subspecies: S. s. scrofa, S. s. vittatus, and S. s. attila.
Khok Sung | Recent | Java (Pleistocene) | ||||||
---|---|---|---|---|---|---|---|---|
S. barbatus | S. scrofa | S. barbatus | S. verrucosus | S. celebensis | S. brachygnathus | S. macrognathus | ||
P3 | L | 16.75 | 12.33–14.41 (16) | 13.17–14.98 (12) | 11.87–13.77 (8) | 9.37 | 11.09–12.29 (7) | 12.47–13.86 (2) |
W | 14.42 | 10.12–12.19 (16) | 10.06–13.22 (12) | 9.71–12.64 (8) | 7.35 | 9.60–11.54 (7) | 10.75–13.43 (2) | |
P4 | L | 15.06 | 11.41–14.61 (16) | 12.56–14.81 (12) | 11.85–13.97 (8) | 8.96–9.44 (3) | 10.35–11.61 (7) | 11.89–12.39 (3) |
W | 18.59 | 12.77–15.23 (16) | 13.51–16.00 (12) | 13.23–14.82 (8) | 10.68–11.01 (3) | 11.05–13.80 (7) | 13.72–15.68 (3) | |
M1 | L | 20.68 | 14.01–17.88 (16) | 16.71–19.24 (12) | 14.36–16.13 (8) | 13.44–13.76 (3) | 13.89–14.94 (6) | 13.62–17.17 (3) |
W | 17.17 | 13.59–17.57 (16) | 13.59–15.85 (12) | 13.32–15.78 (8) | 10.59–11.49 (3) | 12.68–14.36 (6) | 12.57–15.54 (3) | |
M2 | L | 29.35–29.49 (2) | 20.08–24.78 (16) | 22.60–24.60 (12) | 20.53–22.39 (8) | 16.89–17.98 (4) | 19.81–24.26 (7) | 17.17–24.38 (4) |
W | 21.37–23.40 (3) | 16.43–20.82 (16) | 17.45–19.89 (12) | 16.82–19.74 (8) | 13.33–14.96 (4) | 16.09–17.97 (7) | 15.54–21.06 (4) | |
M3 | L | 37.36 | 29.09–39.01 (16) | 30.31–36.50 (12) | 31.75–37.13 (8) | 21.59–24.81 (3) | 27.27–33.26 (8) | 31.44–40.89 (60 |
W | 21.46–24.97 (2) | 19.68–23.76 (16) | 17.44–24.94 (12) | 18.73–20.59 (8) | 14.88–16.18 (3) | 18.08–20.37 (8) | 19.95–24.30 (6) | |
p1 | L | 7.32–7.71 (2) | 7.03–9.13 (8) | 7.25–9.51 (8) | 5.42–7.81 (3) | ? | 6.32–9.98 (6) | ? |
W | 4.16–4.35 (2) | 3.56–4.17 (8) | 3.33–4.09 (8) | 3.22–3.88 (3) | ? | 3.50–5.15 (6) | ? | |
p2 | L | 11.71–13.17 (4) | 10.42–13.21 (16) | 12.10–14.80 (12) | 10.77–11.89 (8) | ? | 9.96–12.02 (6) | ? |
W | 5.55–6.66 (4) | 4.48–6.49 (16) | 4.78–6.61 (12) | 5.84–6.43 (8) | ? | 4.87–5.46 (6) | ? | |
p3 | L | 13.17–14.31 (4) | 13.09–15.75 (16) | 14.01–16.07 (12) | 12.91–14.85 (8) | 10.31 | 11.94–14.59 (7) | 12.14–13.84 (2) |
W | 7.84–8.60 (4) | 6.32–9.10 (16) | 6.51–8.53 (12) | 6.49–7.80 (8) | 6.57 | 6.56–7.38 (7) | 7.44–7.46 (2) | |
p4 | L | 13.87–15.01 (3) | 13.40–16.05 (16) | 14.57–17.29 (12) | 14.44–16.10 (8) | 10.11–10.22 (2) | 12.75–14.30 (8) | 15.41–15.75 (2) |
W | 10.13–11.68 (3) | 8.78–11.44 (16) | 9.18–10.60 (12) | 8.79–11.28 (8) | 7.46–8.34 (2) | 8.84–10.60 (8) | 9.56–10.48 (2) | |
m1 | L | 14.32–18.47 (2) | 14.64–18.75 (16) | 15.94–19.60 (12) | 12.90–14.95 (8) | 12.34–12.61 (3) | 13.77–14.83 (8) | 15.81–17.94 (2) |
W | 13.11–13.8 (2) | 11.55–13.94 (16) | 10.84–13.22 (12) | 11.04–13.56 (8) | 8.55–9.92 (3) | 10.80–12.07 (8) | 11.79–12.11 (2) | |
m2 | L | 19.96–23.38 (2) | 19.66–24.24 (16) | 21.84–23.97 (12) | 19.88–21.22 (8) | 15.35–16.01 (4) | 17.19–20.84 (8) | 21.31–25.00 (3) |
W | 17.65–18.06 (2) | 14.61–17.39 (16) | 14.61–16.56 (12) | 14.14–15.95 (8) | 10.77–13.25 (4) | 12.96–14.45 (8) | 14.15–16.30 (3) | |
m3 | L | 40.92 | 32.92–41.27 (16) | 35.60–43.02 (12) | 37.45–40.27 (8) | 21.68–24.44 (3) | 30.56–39.84 (7) | 40.72–46.37 (4) |
W | 19.89 | 16.71–19.32 (16) | 16.24–19.74 (12) | 15.92–17.84 (8) | 12.16–13.38 (3) | 16.06–21.44 (7) | 15.84–18.15 (4) |
Mandibles and lower dentition: a mandible (DMR-KS-05-03-00-126) preserves both sides of cheek tooth rows (right p2–m3 and left p3–m3), but most of its symphysis and entire ramus are broken off (Fig.
Nasal: a nasal bone (DMR-KS-05-03-00-56) is short and robust, bending downward and narrowing anteriorly towards the tip (Fig.
Postcranial remains: postcranial elements include a scapula (Fig.
Postcranial remains of Rhinoceros sondaicus from Khok Sung: A–B DMR-KS-05-03-00-58, a left scapula in lateral (A) and distal (B) views C–E DMR-KS-05-03-31-3, a left humerus in anterior (C), proximal (D), and distal (E) views F–H DMR-KS-05-04-05-15, a right metacarpus IV in posterior (F), proximal (G), and distal (H) views I DMR-KS-05-04-27-19, a right calcaneus in lateral view J DMR-KS-05-03-26-23, a left astragalus in dorsal view.
Four isolated cheek teeth (P2, P3, M1, and M3) assigned to R. sondaicus are characterized by the following morphological features: a presence of the moderately developed crochet, sinuosity of the ectoloph, distinct parastyle fold, and deeper median valley compared to the posterior valley, and the absences of an antecrochet, protocone fold, and metacone bulge on M3. All of these characters coincide with the upper molars of R. sondaicus (
Large tusk-like incisors (i2) are notably typical of Asian rhinoceroses. The two small alveoli corresponding to the lost central incisors are autapomorphic of Rhinoceros (
Isolated lower molars of rhinoceroses from Khok Sung are difficult to assign to either R. unicornis or R. sondaicus due to heavy wear. In addition, there is a significant size overlap between these two species (
A left mandible with p3–m3, DMR-KS-05-03-17-13; a left p2, DMR-KS-05-03-19-4; a right M1, KS-05-03-18-X; a left femur, DMR-KS-05-03-00-63; a left astragalus, DMR-KS-05-03-00-67.
Upper dentition: a relatively worn M1 (DMR-KS-05-03-18-X) is nearly square in outline and displays a flattened ectoloph and a well developed crochet, medifossette, and posterior fossette (Fig.
Remains of Rhinoceros unicornis from Khok Sung: A DMR-KS-05-03-18-X, a right M1 in occlusal view B DMR-KS-05-03-19-4, a left p2 in occlusal view C–E DMR-KS-05-03-17-13, a left mandible in occlusal (C), medial (D), and lateral (E) views F–G DMR-KS-05-03-00-63, a left femur in posterior (F) and distal (G) views.
Mandible and lower dentition: a hemi-mandible (DMR-KS-05-03-17-13) is strongly compressed laterally and preserves a partial mandibular ramus and body with worn cheek teeth, except for the m3 which is unbroken (Fig.
Postcranial remains: an isolated femur (Fig.
We assign the M1 (DMR-KS-05-03-18-X) to R. unicornis according to the presence of the flattened ectoloph and enclosed medifossette (on a worn specimen), as well as its larger size than that of R. sondaicus. These upper molar features are characteristic of R. unicornis (
A left maxillary fragment with P3–M2, DMR-KS-05-04-19-2; two left M2—DMR-KS-05-04-19-5 and DMR-KS-05-03-18-23 (posterior portion); two right M3—DMR-KS-05-04-03-4 and DMR-KS-05-04-19-4 (anterior portion); two mandible with two tooth rows—DMR-KS-05-03-15-1 (right: i1, i2, c1, p2, and p3 and left: i1, i2, c1, and p2–m2) and DMR-KS-05-04-19-1 (right: i1, i2, c1, and p1–m3 and left: i1, i2, c1, and p1–p4); a left posterior fragment of m3, DMR-KS-05-04-19-3; a right humerus, DMR-KS-05-03-26-8.
Upper dentition: DMR-KS-05-04-19-2 is a maxillary tooth row preserving a slightly worn P3 to M2 (Fig.
Remains of Sus barbatus from Khok Sung: A DMR-KS-05-04-19-2, a left upper cheek tooth row in occlusal view B DMR-KS-05-04-19-5, a left M2 C DMR-KS-05-03-18-23, a left fragmentary M2 D DMR-KS-05-04-03-4, a right M3; (E) DMR-KS-05-04-19-4, a right M3 F–G DMR-KS-05-03-15-1, a mandible in occlusal (F) and lateral (G) views H–I DMR-KS-05-04-19-1, a mandible in occlusal (H) and lateral (I) views J DMR-KS-05-04-19-3, a left fragmentary m3 K–N DMR-KS-05-03-26-8, a right humerus in proximal (K), posterior (L), anterior (M), and distal (N) views. Cross-sections of canines are given. All isolated teeth are shown in occlusal view.
Mandible and lower dentition: DMR-KS-05-03-15-1 is incomplete, lacking the body and ascending ramus, broken posterior to the right p3 and to the left m2 (Fig.
Lower incisors show a chisel-like appearance with long roots. The i2 is larger than the i1. Lower canines are slender and pointed, and curve backward. The lower canines of the mandible DMR-KS-05-03-15-1 belong to a male individual because of a more sharply triangular section (
Postcranial bone: DMR-KS-05-03-26-8 is a complete humerus (Fig.
We compare our material to some Pleistocene Southeast Asian suid species, although only two distinct suid species, S. scrofa and S. barbatus, are known from many Pleistocene localities of mainland Southeast Asia. The sizes of the Khok Sung material are obviously larger than those of Pleistocene and extant Indonesian suids (S. brachygnathus, S. macrognathus, S. verrucosus, and S. celebensis) (Tab.
However, it is difficult to distinguish S. scrofa from S. barbatus only based on the cheek teeth because both species overlap in size (Tab.
Measurements (in millimeters) of lower canines of Khok Sung Sus barbatus. The canine index is expressed by the following formula: labial surface*100/posterior surface (
Specimen no. | Widths | Canine index | |||
---|---|---|---|---|---|
anterolingual surface | posterior surface | labial surface | |||
DMR-KS-05-03-15-1 (male) | right c1 | 13.61 | 8.54 | 11.46 | 134.2 |
left c1 | 13.88 | 9.05 | 11.57 | 127.8 | |
DMR-KS-05-04-19-1 (female) | right c1 | 13.18 | 11.92 | 10.32 | 86.6 |
left c1 | 13.30 | 12.21 | 10.54 | 86.3 |
The female mandible (DMR-KS-05-04-19-1) and other isolated teeth are assigned to S. barbatus according to those described features. We also suggest that Pleistocene Sus barbatus probably shows evidence of sexual size dimorphism because the female specimen DMR-KS-05-04-19-1 is markedly smaller than the male specimen DMR-KS-05-03-15-1, as seen in the recent population.
Four crania—DMR-KS-05-04-18-50 (with two antlers), DMR-KS-05-03-00-30 (with left partial and right broken antlers), DMR-KS-05-03-18-X9 (with pedicles), and DMR-KS-05-03-27-1 (with pedicles); two right complete antlers—DMR-KS-05-03-31-30 and DMR-KS-05-03-22-4; a nearly complete left antler, DMR-KS-05-04-4-1; five right fragmentary antlers—DMR-KS-05-03-18-21, DMR-KS-05-03-19-82, DMR-KS-05-03-28-22, DMR-KS-05-06-22-2, and DMR-KS-05-03-28-1; eight left fragmentary antlers—DMR-KS-05-03-00-12, DMR-KS-05-03-19-81, DMR-KS-05-03-22-2, DMR-KS-05-03-24-1, DMR-KS-05-04-09-1, DMR-KS-05-03-19-13, DMR-KS-05-03-26-21, and DMR-KS-05-03-08-17; two left fragmentary maxilla—DMR-KS-05-03-28-6 (with M1–M3) and DMR-KS-05-03-08-31 (with P3, P4, and M1 root); a right P4, DMR-KS-05-04-01-3; a left M1, DMR-KS-05-04-28-5; a left M2, DMR-KS-05-03-14-5; thirteen right mandibles—DMR-KS-05-03-14-2 (with m3), DMR-KS-05-03-20-1 (with p4–m3), DMR-KS-05-03-20-2 (with m2 and m3), DMR-KS-05-03-22-7 (with m2 and m3), DMR-KS-05-04-03-1 (with p2–m3), and DMR-KS-05-03-27-3 (with m2 and m3), DMR-KS-05-03-19-1 (with p2–m3), DMR-KS-05-03-22-8 (with m2 and m3), DMR-KS-05-04-01-1 (with p2–m3), DMR-KS-05-03-24-4 (with m2), DMR-KS-05-03-26-12 (with m2 and m3), DMR-KS-05-04-7-10 (with p3, m1, and m2), and DMR-KS-05-03-26-10 (with p2–m1); eight left mandibles—DMR-KS-05-03-18-22 (with p2), DMR-KS-05-03-22-6 (with m1–m3), DMR-KS-05-03-27-22 (with p3-m2 sockets and broken m3), DMR-KS-05-04-09-2 (with p3, p4, m1 and m2 sockets, and m3), DMR-KS-05-03-00-102 (with p4 and m1), DMR-KS-05-03-19-2 (with m1–m3), DMR-KS-05-03-23-1 (with p2 and p3 roots and p4–m3), and DMR-KS-05-03-29-1 (with p2-m3); a left m1, DMR-KS-05-04-28-6; three m2—DMR-KS-05-03-25-4 (right), DMR-KS-05-03-00-104 (left), and DMR-KS-05-03-22-11 (left); four left m3—DMR-KS-05-04-9-4, DMR-KS-05-03-22-9, DMR-KS-05-04-01-2, and DMR-KS-05-03-08-33; three right fragmentary humeri (distal part)—DMR-KS-05-03-13-4, DMR-KS-05-04-11-32, and DMR-KS-05-03-17-17; six metacarpi—DMR-KS-05-03-18-2 (right), DMR-KS-05-03-19-3 (right), DMR-KS-05-03-22-28 (right), DMR-KS-05-03-08-2 (right), DMR-KS-05-04-30-20 (right proximal fragment), and DMR-KS-05-03-19-37 (left); a right fragmentary femur, DMR-KS-05-03-27-4 (distal part); three metatarsi—DMR-KS-05-03-26-3 (right), DMR-KS-05-03-29-30 (left), and DMR-KS-05-03-15-14 (left).
Crania and upper dentition: four crania are almost complete, lacking only the anterior portions (e.g., nasal, jugal, palatine, and maxilla) (Fig.
Cranial remains of Axis axis from Khok Sung: A–B DMR-KS-05-04-18-50, a cranium with nearly complete antlers in dorsal (A) and ventral (B) views C–D DMR-KS-05-03-00-30, a cranium in lateral (C) amd ventral (D) views E DMR-KS-05-03-18-X9, a cranium in anterior view F–G DMR-KS-05-03-27-1 a cranium in dorsal (F) and ventral (G) views H DMR-KS-05-03-31-30, a right antler in anterior view; (I) DMR-KS-05-03-22-4, a right antler in lateral view J DMR-KS-05-03-18-21, a left antler fragment in lateral view KDMR-05-03-22-2, a left antler fragment in lateral view L DMR-KS-05-03-19-81, a left antler fragment in medial view.
P3 and P4 are similar to recent Axis, characterized by well-developed styles, medial cristae (more distinct on the P4), and posterolingual fossettes (Fig.
Dental remains of Axis axis from Khok Sung: A DMR-KS-05-03-08-31, an upper left P3 and P4 in occlusal view B DMR-KS-05-03-28-6, a left upper molar row in occlusal view C DMR-KS-05-04-01-3, a right P4 in occlusal view D DMR-KS-05-04-28-5, a left M1 in occlusal view E DMR-KS-05-03-14-5, a left M2 in occlusal view F–G DMR-KS-05-03-29-1, a left mandible in occlusal (F) and lateral (G) views H–I DMR-KS-05-03-26-10, a right mandibular fragment in occlusal (H) and medial (I) views J–K DMR-KS-05-04-03-1, a right mandible in occlusal (J) and lateral (K) views L–M DMR-KS-05-03-20-1, a right mandible in occlusal (L) and lateral (M) views N–O DMR-KS-05-03-22-7, a right mandible in occlusal (N) and lateral (O) views P–Q DMR-KS-05-03-08-33, a left m3 in occlusal (P) and buccal (Q) views.
Measurements (lengths and widths in millimeters) of cervid teeth from Khok Sung. N=number of specimens.
Length | Width | |||||
---|---|---|---|---|---|---|
N | Range | Mean | N | Range | Mean | |
Axis axis | ||||||
P3 | 1 | 12.40 | – | 1 | 13.60 | – |
P4 | 2 | 10.04–11.29 | 10.67 | 2 | 12.19–14.28 | 13.24 |
M1 | 2 | 13.32–15.19 | 14.26 | 2 | 15.60–15.93 | 15.77 |
M2 | 2 | 18.07–18.08 | 18.08 | 2 | 17.41–17.84 | 17.63 |
M3 | 1 | 17.53 | – | 1 | 16.42 | – |
p2 | 6 | 7.93–9.54 | 8.72 | 6 | 5.44–6.89 | 5.93 |
p3 | 7 | 9.17–12.11 | 10.67 | 7 | 6.53–7.14 | 6.88 |
p4 | 8 | 10.64–13.62 | 11.65 | 10 | 6.77–8.13 | 7.39 |
m1 | 9 | 11.81–18.20 | 14.2 | 13 | 8.27–10.29 | 9.59 |
m2 | 18 | 15.94–21.42 | 17.91 | 19 | 8.56–11.67 | 10.56 |
m3 | 18 | 21.69–25.78 | 24.1 | 20 | 8.87–11.89 | 10.74 |
Panolia eldii | ||||||
P2 | 1 | 11.09 | – | 1 | 13.97 | – |
M1 | 2 | 12.07–14.95 | 13.51 | 2 | 16.52–17.77 | 17.15 |
M2 | 5 | 16.67–20.48 | 19.35 | 6 | 17.85–19.35 | 18.56 |
M3 | 5 | 18.80–21.39 | 19.96 | 5 | 16.99–19.50 | 18.30 |
i1 | 1 | 12.86 | – | 1 | 6.31 | – |
p2 | 2 | 9.97–11.33 | 10.65 | 2 | 7.03–7.44 | 7.24 |
p3 | 2 | 13.04–13.67 | 13.36 | 2 | 8.33–8.56 | 8.45 |
p4 | 2 | 13.65–14.05 | 13.85 | 2 | 8.94–9.33 | 9.14 |
m1 | 2 | 14.67–15.67 | 15.17 | 2 | 11.23–12.25 | 11.74 |
m2 | 2 | 17.73–19.36 | 18.55 | 2 | 12.63–13.26 | 12.95 |
m3 | 1 | 23.61 | 1 | 12.84 | ||
Rusa unicolor | ||||||
M1 | 1 | 17.15 | – | 1 | 20.10 | – |
M2 | 2 | 20.67–22.88 | 21.78 | 2 | 23.06–27.07 | 25.07 |
M3 | 1 | 25.37 | – | 1 | 24.97 | – |
p3 | 1 | 17.29 | – | 1 | 9.26 | – |
p4 | 1 | 17.71 | – | 2 | 10.34–13.35 | 11.85 |
m1 | 2 | 18.64–20.84 | 19.74 | 2 | 14.39–14.59 | 14.49 |
m2 | 3 | 22.77–23.82 | 23.33 | 3 | 15.37–15.61 | 15.46 |
m3 | 3 | 30.78–34.57 | 32.67 | 3 | 15.49–17.85 | 16.79 |
Mandibles and lower dentition: twenty one mandibles range from fragmentary (preserving only the broken corpus) to nearly complete (lacking only the ascending ramus and coronoid process) individuals (Fig.
Lower third and fourth premolars exhibit a well developed metaconid which projects obliquely in occlusal view, posterior to the entoconid (Fig.
Postcranial remains: postcranial bones include isolated humeri (Fig.
Postcranial remains of Axis axis from Khok Sung: A–B DMR-KS-05-04-11-32, a right distal humerus in anterior (A) and distal (B) views C–E DMR-KS-05-03-18-2, a right metacarpus in proximal (C), anterior (D), and distal (E) views F–H DMR-KS-05-03-19-37, a left metacarpus in proximal (F), anterior (G), and distal (H) views I–J DMR-KS-05-03-27-4, a right distal femur in posterior (I) and distal (J) views K–M DMR-KS-05-03-26-3, a right metatarsus in proximal (K), anterior (L), and distal (M) views.
Proportional indices of postcranial remains of identified ruminant taxa from Khok Sung.
Scapula | ||||||||||||
Specimen | Taxa | HS/Ld | DHA/Ld | Ld/SLC | LG/BG | GLP/LG | SLC/BG | |||||
DMR-KS-05-03-26-2 | Bubalus arnee | 1.50 | 1.28 | 3.89 | 1.20 | 1.30 | 1.12 | |||||
DMR-KS-05-02-20-4 | Bubalus arnee | 1.39 | 1.42 | 4.09 | 1.23 | 1.26 | 0.96 | |||||
DMR-KS-05-06-24-4 | Panolia eldii | 1.95 | 1.90 | 4.62 | 1.10 | 1.27 | 0.74 | |||||
Humerus | ||||||||||||
Specimen | Taxa | GL/Bp | GL/Dp | GL/Bd | GL/Dd | Bp/Bd | Dp/Dd | Bp/Dp | Bd/Dd | Bd/BT | ||
DMR-KS-05-03-20-2(1) | Bos sauveli | – | – | – | – | – | – | – | 0.99 | 1.04 | ||
DMR-KS-05-03-00-62 | Bos gaurus | – | – | 3.41 | 3.66 | – | – | – | 1.07 | 1.06 | ||
DMR-KS-05-05-1-1 | Bos gaurus | 2.91 | 2.74 | 3.44 | 3.67 | 1.18 | 1.34 | 0.94 | 1.07 | 1.05 | ||
DMR-KS-05-03-31-1 | Bubalus arnee | 3.57 | 3.25 | 4.30 | 4.77 | 1.21 | 1.47 | 0.91 | 1.11 | 1.05 | ||
DMR-KS-05-03-31-8 | Bubalus arnee | 3.54 | 3.29 | 4.25 | 4.74 | 1.20 | 1.44 | 0.93 | 1.11 | 1.03 | ||
DMR-KS-05-03-13-4 | Axis axis | – | – | – | – | – | – | – | 1.02 | 1.09 | ||
DMR-KS-05-04-11-32 | Axis axis | – | – | – | – | – | – | – | 1.06 | 1.07 | ||
DMR-KS-05-03-17-17 | Axis axis | – | – | – | – | – | – | – | 1.12 | 1.04 | ||
DMR-KS-05-04-11-35 | Panolia eldii | – | – | – | – | – | – | – | 1.12 | 1.13 | ||
DMR-KS-05-03-18-1 | Panolia eldii | – | – | – | – | – | – | 0.82 | – | – | ||
DMR-KS-05-03-15-43 | Rusa unicolor | – | – | – | – | – | – | – | 1.14 | 1.12 | ||
Ulna and radius | ||||||||||||
Specimen | Taxa | PL/Bp | PL/Dp | PL/Bd | PL/Dd | Bp/Bd | Dd/Dp | Bp/Dp | Bd/Dd | Bp/BFp | Bd/BFd | GL/LO |
DMR-KS-05-03-00-61 | Bubalus arnee | 2.87 | 5.76 | 3.04 | 4.63 | 1.06 | 1.24 | 2.00 | 1.52 | 1.15 | 1.11 | 3.86 |
DMR-KS-05-03-31-2 | Bubalus arnee | 3.15 | 5.85 | 3.25 | 4.61 | 1.03 | 1.27 | 1.86 | 1.42 | 1.09 | 1.12 | 3.48 |
DMR-KS-05-03-31-9 | Bubalus arnee | 3.09 | 5.88 | 3.24 | 4.55 | 1.05 | 1.29 | 1.90 | 1.40 | 1.10 | 1.12 | 3.45 |
DMR-KS-05-03-31-10 | Panolia eldii | 5.06 | 9.51 | 5.35 | 9.32 | 1.06 | 1.02 | 1.88 | 1.74 | 1.07 | 1.14 | – |
DMR-KS-05-04-11-3 | Panolia eldii | 4.83 | 9.09 | 5.54 | 8.70 | 1.15 | 1.04 | 1.88 | 1.57 | 1.11 | 1.06 | – |
DMR-KS-05-03-19-16 | Panolia eldii | 4.93 | 8.93 | 4.87 | 6.62 | 0.99 | 1.35 | 1.81 | 1.36 | 1.22 | 1.04 | – |
DMR-KS-05-03-25-9 | Rusa unicolor | – | – | – | – | – | – | 1.90 | – | 1.03 | – | – |
DMR-KS-05-03-19-14 | Rusa unicolor | – | – | – | – | – | – | 1.70 | – | 1.04 | – | – |
DMR-KS-05-03-26-19 | Rusa unicolor | – | – | – | – | – | – | – | 1.34 | – | 1.05 | – |
Femur | ||||||||||||
Specimen | Taxa | GL/Bp | GL/Dp | GL/Bd | GL/Dd | Bp/Bd | Dd/Dp | Bp/Dp | Dd/Bd | |||
DMR-KS-05-03-9-2 | Bos gaurus | 3.37 | 6.29 | 3.92 | 3.03 | 1.17 | 2.07 | 1.87 | 1.29 | |||
DMR-KS-05-04-1-1 | Bubalus arnee | 2.79 | 5.54 | 3.48 | 2.85 | 1.25 | 1.95 | 1.99 | 1.22 | |||
DMR-KS-05-04-1-2 | Bubalus arnee | 2.67 | 5.26 | 3.38 | 2.82 | 1.27 | 1.86 | 1.97 | 1.20 | |||
DMR-KS-05-03-20-8 | Bubalus arnee | – | – | – | – | – | – | – | 1.46 | |||
DMR-KS-05-03-27-4 | Axis axis | – | – | – | – | – | – | – | 1.37 | |||
DMR-KS-05-03-27-11 | Panolia eldii | – | – | – | – | 1.26 | 2.23 | 2.11 | 1.33 | |||
DMR-KS-05-03-17-36 | Panolia eldii | – | – | – | – | 1.21 | 2.06 | 1.93 | 1.29 | |||
DMR-KS-05-03-28-20 | Panolia eldii | – | – | – | – | – | – | – | 1.34 | |||
DMR-KS-05-04-05-38 | Panolia eldii | – | – | – | – | – | – | 1.92 | – | |||
DMR-KS-05-03-00-119 | Panolia eldii | – | – | – | – | – | – | – | 1.38 | |||
DMR-KS-05-03-19-2 | Panolia eldii | – | – | – | – | – | – | – | 1.41 | |||
DMR-KS-05-08-16-1 | Panolia eldii | – | – | – | – | – | – | 1.84 | – | |||
DMR-KS-05-04-11-2 | Rusa unicolor | – | – | – | – | – | – | – | 1.27 | |||
DMR-KS-05-03-19-7 | Rusa unicolor | – | – | – | – | – | – | 1.51 | – | |||
DMR-KS-05-03-12-2* | Rusa unicolor | – | – | – | – | – | – | 1.52 | – | |||
DMR-KS-05-04-30-9 | Rusa unicolor | – | – | – | – | – | – | – | 1.27 | |||
DMR-KS-05-04-19-10 | Rusa unicolor | – | – | – | – | – | – | – | 1.11 | |||
Tibia | ||||||||||||
Specimen | Taxa | GL/Bp | GL/Dp | GL/Bd | GL/Dd | Bp/Bd | Dp/Dd | Bp/Dp | Bd/Dd | |||
DMR-KS-05-04-1-11 | Bubalus arnee | 3.24 | 3.43 | 4.82 | 6.03 | 1.49 | 1.76 | 1.06 | 1.25 | |||
DMR-KS-05-04-1-3 | Bubalus arnee | 3.31 | 3.50 | 5.01 | 6.29 | 1.51 | 1.80 | 1.06 | 1.25 | |||
DMR-KS-05-03-20-9 | Bubalus arnee | 3.21 | 3.83 | 4.60 | 6.29 | 1.43 | 1.64 | 1.19 | 1.37 | |||
DMR-KS-05-03-28-16 | Rusa unicolor | 4.00 | 4.38 | 6.68 | 8.48 | 1.67 | 1.94 | 1.10 | 1.27 | |||
Metacarpus | ||||||||||||
Specimen | Taxa | GL/Bp | GL/Dp | GL/Bd | GL/Dd | Bp/Bd | Dp/Dd | Bp/Dp | Bd/Dd | |||
DMR-KS-05-03-26-27 | Bos gaurus | 3.66 | 5.57 | 3.96 | 7.66 | 1.08 | 1.37 | 1.52 | 1.93 | |||
DMR-KS-05-03-26-3(1) | Bubalus arnee | 2.68 | 4.17 | 2.64 | 4.87 | 0.98 | 1.17 | 1.55 | 1.85 | |||
DMR-KS-05-03-18-2 | Axis axis | 6.50 | 9.99 | 6.69 | 10.55 | 1.03 | 1.06 | 1.54 | 1.58 | |||
DMR-KS-05-03-22-28 | Axis axis | – | 9.59 | 6.81 | 10.36 | – | 1.08 | – | 1.52 | |||
DMR-KS-05-03-08-2 | Axis axis | 6.36 | 8.79 | 6.18 | 10.18 | 0.97 | 1.16 | 1.38 | 1.65 | |||
DMR-KS-05-03-19-3 | Axis axis | 6.58 | 9.06 | 6.30 | 10.42 | 0.96 | 1.15 | 1.38 | 1.65 | |||
DMR-KS-05-03-19-37 | Axis axis | 7.14 | 11.05 | 6.84 | 10.75 | 0.96 | 0.97 | 1.55 | 1.57 | |||
DMR-KS-05-04-30-20 | Axis axis | 6.87 | 10.36 | – | – | – | – | 1.51 | – | |||
DMR-KS-05-03-24-2 | Panolia eldii | 6.39 | 8.99 | 6.57 | 10.41 | 1.03 | 1.16 | 1.41 | 1.58 | |||
DMR-KS-05-03-17-26 | Rusa unicolor | 5.97 | 7.57 | 6.06 | 9.10 | 1.02 | 1.20 | 1.27 | 1.50 | |||
Metatarsus | ||||||||||||
Specimen | Taxa | GL/Bp | GL/Dp | GL/Bd | GL/Dd | Bp/Bd | Dp/Dd | Bp/Dp | Bd/Dd | |||
DMR-KS-05-04-1-8 | Bubalus arnee | 3.80 | 4.59 | 3.17 | 5.54 | 0.83 | 1.21 | 1.21 | 1.75 | |||
DMR-KS-05-04-1-6 | Bubalus arnee | 3.88 | 4.39 | 3.11 | 5.67 | 0.80 | 1.29 | 1.13 | 1.82 | |||
DMR-KS-05-03-28-30 | Bubalus arnee | 4.25 | 4.28 | 3.40 | 6.38 | 0.80 | 1.49 | 1.01 | 1.88 | |||
DMR-KS-05-03-26-3 | Axis axis | 7.21 | 6.91 | 6.99 | 9.16 | 0.97 | 1.33 | 0.96 | 1.31 | |||
DMR-KS-05-03-15-14 | Axis axis | 6.84 | 7.37 | 6.15 | 9.22 | 0.90 | 1.25 | 1.08 | 1.50 | |||
DMR-KS-05-03-29-30 | Axis axis | 6.91 | 6.82 | 6.52 | 8.58 | 0.94 | 1.26 | 0.99 | 1.32 | |||
DMR-KS-05-03-28-17 | Panolia eldii | 8.05 | 7.71 | 7.73 | 11.69 | 0.96 | 1.52 | 0.96 | 1.51 | |||
DMR-KS-05-03-25-8 | Panolia eldii | 7.81 | 7.47 | 7.22 | 11.57 | 0.92 | 1.55 | 0.96 | 1.60 | |||
DMR-KS-05-03-15-15 | Panolia eldii | 8.08 | 7.44 | 7.37 | 11.29 | 0.91 | 1.52 | 0.92 | 1.53 | |||
DMR-KS-05-03-19-11 | Rusa unicolor | 6.64 | 6.86 | 6.49 | 9.20 | 0.98 | 1.34 | 1.03 | 1.42 |
The antlers are useful to distinguish among the cervids, whereas the morphologies of lower cheek teeth are identical among Axis. The skulls, antlers, and teeth from Khok Sung are morphologically similar to those observed from recent A. axis. This suggests a morphological stasis in the evolution of antlers and teeth for this species.
Based on our observation on the extant comparative material of A. axis (e.g., the specimens
Body mass prediction of Khok Sung ruminants using second molar variables, compared to relative sizes of the recent population (
Body mass (kg) | ||||
---|---|---|---|---|
Cervidae | Khok Sung | Recent | ||
Taxa | N | Range | Mean | Range |
Axis axis | 17 | 67.6–127.6 | 90.8 | 75–100 |
Panolia eldii | 7 | 99.1–157.6 | 133.5 | 95–150 |
Rusa unicolor | 5 | 215.6–332.3 | 255.4 | 100–350 |
Bovidae | Khok Sung | Recent | ||
Taxa | N | Range | Mean | Range |
Bos sauveli | 3 | 660.8–756.0 | 720.5 | 700–900 |
Bos gaurus | 3 | 808.5–940.8 | 873.2 | 700–1000 |
Bubalus arnee | 12 | 694.5–1243.0 | 944.7 | 700–1200 |
Compared to other Pleistocene cervid species, the cheek teeth of A. axis from Khok Sung are smaller than those of A. shansius from Anhui and Yunnan (China) and of A. javanicus from Ngandong and Buitenzorg in Java and Carnul Cave in India, but are larger than those of A. lydekkeri from Trinil H. K. (Java) (Figs
A cranium with a right partial antler, DMR-KS-05-04-20-4; a right P2, DMR-KS-05-03-15-11; two left M1—DMR-KS-05-03-00-24 and DMR-KS-05-03-00-25; six M2—DMR-KS-05-03-00-23 (right), DMR-KS-05-03-30-5 (right), DMR-KS-05-04-3-4 (right), DMR-KS-05-03-30-6 (left posterior lobe), DMR-KS-05-03-27-7 (left), and DMR-KS-05-04-3-5 (left); five M3—DMR-KS-05-03-27-6 (right), DMR-KS-05-04-9-1 (right), DMR-KS-05-04-8-3 (right), DMR-KS-05-03-00-22 (left), and DMR-KS-05-04-9-2 (left); two left mandibles—DMR-KS-05-03-27-2 (with p2–m3) and DMR-KS-05-04-9-5 (with p2–m2); a right i1, DMR-KS-05-03-29-2; a right scapula, DMR-KS-05-06-24-4; a left humerus, DMR-KS-05-04-11-35; a right fragmentary humerus, DMR-KS-05-03-18-1 (proximal part); three radii—DMR-KS-05-03-31-10 (right), DMR-KS-05-04-11-3 (right), and DMR-KS-05-03-19-16 (left); a right metacarpus, DMR-KS-05-03-24-2; two right femora—DMR-KS-05-03-27-11 and DMR-KS-05-03-17-36; five fragmentary femora—DMR-KS-05-04-05-38 (right proximal part), DMR-KS-05-03-28-20 (right distal part), DMR-KS-05-03-00-119 (right distal part), DMR-KS-05-03-19-2 (right distal part), and DMR-KS-05-08-16-1 (left proximal part); three left metatarsi—DMR-KS-05-03-25-8, DMR-KS-05-03-28-17, and DMR-KS-05-03-15-15.
Cranium and upper dentition: DMR-KS-05-04-20-4 is an incomplete cranium, lacking the whole anterior parts (nasal, jugal, palatine, and maxilla) (Fig.
Remains of Panolia eldii from Khok Sung: A–C DMR-KS-05-04-20-4, a cranium in dorsal (A), lateral (B), and ventral (C) views D DMR-KS-05-03-15-11, a right P2 E DMR-KS-05-03-00-24, a left M1 F DMR-KS-05-03-00-23, a right M2 G DMR-KS-05-03-27-6, a left M3 H DMR-KS-05-04-9-2, a left M3 I DMR-KS-05-03-29-2, a right i1 in lingual view J–K DMR-KS-05-03-27-2, a left mandible in lateral (J) and occlusal (K) views L–M DMR-KS-05-04-9-5, a left mandible in occlusal (L) and lateral (M) views. All teeth are shown in occlusal view.
P2 exhibits a prominent medial crista which divides the fossette into two islands (Fig.
Mandibles and lower dentition: Two mandibles (DMR-KS-05-03-27-2: Fig.
An isolated i1 is spatulate (Fig.
Postcranial remains: postcranial bones include a scapula (Fig.
Postcranial remains of Panolia eldii from Khok Sung: A–B DMR-KS-05-06-24-4, a right scapula in lateral (A) and distal (B) views C–E DMR-KS-05-03-18-1, a right proximal humerus in proximal (C), anterior (D), and posterior (E) views F–H DMR-KS-05-03-31-10, a right radius in proximal (F), anterior (G), distal (H) views I–K DMR-KS-05-03-24-2, a right metacarpus in proximal (I), anterior (J), and distal (K) views L–N DMR-KS-05-03-25-8, a left metatarsus in proximal (L), anterior (M), distal (N) views O–Q DMR-KS-05-03-17-36, a right femur in proximal (O), posterior (P), distal (Q) views.
Several authors consider Eld’s deer as belonging to either the genus Cervus (e.g.,
The shed antler of the Eld’s deer, Panolia eldii, is characterized by bow- or lyre-like shapes, long, noticeable, and laterally bending-main beams with a distal portion curving medially, and small ornamented branches of brow tines. The cheek teeth of P. eldii differ from those of A. axis in having a larger size, a more complex wear pattern of the mesolingual conids on the p3, more developed anterior cingulids on the lower molars, and a posterior ectostylid on the m3. The Khok Sung specimens assigned to P. eldii are similar in morphology to the extant specimens. As demonstrated by the body mass estimation (Tab.
Scatter diagrams of upper cheek tooth (P2, M1, M2, and M3) lengths and widths of some recent and fossil large cervids. The measurements of fossil cervids from the caves of Phnom Loang, Thum Wiman Nakin, and Ma U’Oi are obtained from
Scatter diagrams of lower cheek tooth (p2–m3) lengths and widths of some recent and fossil large cervids. The measurements of fossil cervids from the caves of Thum Wiman Nakin, Thum Prakai Phet, and Ma U’Oi are obtained from
Three right antlers—DMR-KS-05-03-20-11 (nearly complete specimen), DMR-KS-05-03-26-2 (fragment), and DMR-KS-05-03-28-23 (fragment); a right M1, DMR-KS-05-03-22-10; two left M2—DMR-KS-05-04-9-3 and DMR-KS-05-04-3-3; a left M3, DMR-KS-05-03-31-1; two right mandibles—DMR-KS-05-03-31-2 (with m2) and DMR-KS-05-03-13 (with p4–m3); two left mandibles—DMR-KS-05-03-00-101 (with p3–m3) and DMR-KS-05-03-27-4 (with m3); a right m1, DMR-KS-05-03-00-5; a left fragmentary humerus, DMR-KS-05-03-15-43 (distal part); three right fragmentary radii—DMR-KS-05-03-25-9 (proximal part), DMR-KS-05-03-19-14 (proximal part), and DMR-KS-05-03-26-19 (distal part); a left metacarpus, DMR-KS-05-03-17-26; six fragmentary femora—DMR-KS-05-03-19-7 (right proximal part), DMR-KS-05-03-12-2 (right proximal part), DMR-KS-05-04-11-2 (right distal part), DMR-KS-05-03-26-5 (left proximal part), DMR-KS-05-04-30-9 (left distal part), and DMR-KS-05-04-19-10 (left distal part); a right tibia, DMR-KS-05-03-28-16; a right metatarsus, DMR-KS-05-03-19-11
Antlers: DMR-KS-05-03-20-11 is a nearly complete antler, slightly broken at the middle part of the main beam (Fig.
Remains of Rusa unicolor from Khok Sung: A DMR-KS-05-03-20-11, a right antler in lateral view B DMR-KS-05-03-26-2, a right antler fragment in lateral view C DMR-KS-05-03-28-23, a right antler fragment in medial view D DMR-KS-05-04-9-3, a left M2 E DMR-KS-05-03-31-1, a left M3 F–G DMR-KS-05-03-13, a right mandible in lateral (F) and occlusal (G) views H–I DMR-KS-05-03-00-101, a left mandible in lateral (H) and occlusal (I) views J DMR-KS-05-03-00-5, a right m1 K DMR-KS-05-03-27-4, a left m3. All isolated teeth are shown in occlusal view.
Upper dentition: upper molars are robust (Tab.
Mandibles and lower dentition: four mandibles are incomplete (for measurements, see Appendix
Postcranial remains: postcranial elements include a humerus (Fig.
Postcranial remains of Rusa unicolor from Khok Sung: A–C DMR-KS-05-03-15-43, a left humerus in anterior (A), posterior (B), and distal (C) views D–E DMR-KS-05-03-19-14, a right proximal radius in proximal (D) and anterior (E) views F–G DMR-KS-05-03-26-19, a right distal radius in anterior (F) and distal (G) views H–J DMR-KS-05-03-17-26, a left metacarpus in proximal (H), anterior (I), and distal (J) views K–L DMR-KS-05-03-19-7, a right proximal femur in proximal (K) and anterior (L) views M–N DMR-KS-05-04-30-9, a left distal femur in posterior (M) and distal (N) views O–Q DMR-KS-05-03-28-16, a right tibia in proximal (O), anterior (P), and distal (Q) views R–T DMR-KS-05-03-19-11, a right metatarsus in proximal (R), anterior (S), and distal (T) views.
According to Leslie (2011), we regard here Rusa as a separate genus within the family Cervidae. Four species are currently recognized: R. unicolor (sambar), R. marianna (Philippine deer), R. timorensis (rusa), and R. alfredi (Prince Alfred’s deer).
Antlers of R. unicolor are characterized by its typical three tines and forked beams at the tip, similar in shape to those of Axis porcinus but much more robust. The sambar deer shares a similar dental morphology with the Eld’s deer. But it differs from P. eldii as well as A. axis in being larger-sized and in having more developed anterior cingulids on lower molars. The sambar deer is much larger than A. axis (Figs
A left DP3, DMR-KS-05-03-29-8; a left P3, DMR-KS-05-04-01-4; a left fragmentary M1 or M2 (posterior portion), DMR-KS-05-03-23-2; a right M3, DMR-KS-05-03-29-6; a right mandible with m1–m3, DMR-KS-05-03-9-1; two left mandibles—DMR-KS-05-04-9-1 (with p2, p4, and m1–m3) and DMR-KS-05-04-29-1 (with m3); a left i2, DMR-KS-05-03-15-12; a right i3, DMR-KS-05-03-23-4; a right p2, DMR-KS-05-04-01-6; a right m1, DMR-KS-05-03-15-10; a right m2, DMR-KS-05-03-29-7; two m3—DMR-KS-05-04-28-4 (right broken posterior lobe) and DMR-KS-05-03-24-5 (left); a left humerus, DMR-KS-05-03-20-2(1).
Upper dentition: DP3 (DMR-KS-05-03-29-8) is molariform and elongated, characterized by well-developed anterior and posterior cingula, buccal styles, and medial fossettes, a slightly-developed entostyle, and a reduction of the anterior lobe width and height compared to the posterior lobe (Fig.
Remains of Bos sauveli from Khok Sung: A DMR-KS-05-03-29-8, a left DP3 B DMR-KS-05-04-01-4, a left P3 C–D DMR-KS-05-03-23-2, a left M1 or M2 in occlusal (C) and lingual (D) views E DMR-KS-05-03-29-6, a right M3 F DMR-KS-05-04-01-6, a right p2 G DMR-KS-05-04-28-4, a broken right m3 H–I DMR-KS-05-04-9-1, a left mandible in occlusal (H) and lateral (I) views J–K DMR-KS-05-03-9-1, a right mandible in occlusal (J) and lateral (K) views L–N DMR-KS-05-03-20-2(1), a left humerus in anterior (L), posterior (M), and distal (N) views. All isolated teeth are shown in occlusal view.
Measurements (lengths and widths in millimeters) of large bovid teeth from Khok Sung. N=number of specimens.
Length | Width | |||||
---|---|---|---|---|---|---|
N | Range | Mean | N | Range | Mean | |
Bos sauveli | ||||||
DP3 | 1 | 27.39 | – | 1 | 14.91 | – |
P3 | 1 | 17.57 | – | 1 | 19.71 | – |
M1 or M2 | – | – | – | 1 | 25.63 | – |
M3 | 1 | 35.46 | – | 1 | 23.55 | – |
i2 | 1 | 13.67 | – | 1 | 11.53 | – |
i3 | 1 | 13.68 | – | 1 | 8.68 | – |
p2 | 2 | 14.13–14.77 | 14.45 | 2 | 8.52–10.39 | 9.46 |
p4 | 1 | 23.39 | – | 1 | 12.91 | – |
m1 | 3 | 27.24–27.96 | 27.72 | 3 | 17.21–18.26 | 17.73 |
m2 | 3 | 29.70–32.47 | 30.11 | 3 | 17.87–18.79 | 18.29 |
m3 | 3 | 40.60–47.60 | 43.78 | 5 | 17.09–19.91 | 18.37 |
Bos gaurus | ||||||
DP2 | 1 | 22.28 | – | 1 | 10.67 | – |
P2 | 2 | 19.42–20.79 | 20.11 | 2 | 13.55–15.58 | 14.57 |
DP3 | 1 | 28.73 | – | 1 | 18.97 | – |
DP4 | 1 | 29.75 | – | 1 | 22.55 | – |
M1 | 1 | 26.33 | – | 1 | 29.95 | – |
M3 | 1 | 36.96 | – | 1 | 26.94 | – |
i1 | 1 | 20.30 | – | 1 | 11.35 | – |
p2 | 1 | 13.77 | – | 1 | 8.56 | – |
p3 | 1 | 21.58 | – | 1 | 11.92 | – |
p4 | 1 | 21.11 | – | 1 | 12.72 | – |
m1 | 2 | 25.29–28.67 | 26.98 | 2 | 18.25–19.28 | 18.77 |
m2 | 3 | 30.36–35.09 | 32.82 | 3 | 19.00–20.07 | 19.45 |
m3 | 2 | 42.56–46.23 | 44.40 | 2 | 18.72–18.79 | 18.76 |
Bubalus arnee | ||||||
P2 | 3 | 22.30–26.78 | 24.04 | 3 | 14.47–17.26 | 15.76 |
DP3 | 1 | 31.92 | – | 1 | 19.75 | – |
P3 | 7 | 17.85–25.03 | 21.58 | 7 | 15.56–21.93 | 20.32 |
DP4 | 1 | 31.60 | – | 1 | 23.36 | – |
P4 | 7 | 17.76–23.55 | 20.46 | 7 | 21.01–23.20 | 22.34 |
M1 | 9 | 25.73–33.16 | 28.61 | 8 | 26.01–29.79 | 27.30 |
M2 | 8 | 30.45–36.18 | 33.11 | 7 | 26.09–29.23 | 27.23 |
M3 | 6 | 33.74–37.40 | 36.07 | 6 | 25.26–27.30 | 26.29 |
i1 | 1 | 21.21 | – | 1 | 10.31 | – |
i2 | 1 | 16.17 | – | 1 | 11.94 | – |
i3 | 1 | 16.61 | – | 1 | 11.63 | – |
i4 | 1 | 15.82 | – | 1 | 8.80 | – |
p2 | 4 | 13.56–16.24 | 15.05 | 4 | 8.01–9.80 | 8.87 |
dp3 | 2 | 21.59–23.20 | 22.40 | 2 | 8.65–9.90 | 9.28 |
p3 | 3 | 21.88–23.09 | 22.30 | 3 | 10.23–13.09 | 11.80 |
dp4 | 3 | 37.25–42.59 | 40.74 | 3 | 13.34–15.24 | 14.39 |
p4 | 2 | 23.81–24.97 | 24.39 | 3 | 11.93–13.26 | 12.76 |
m1 | 9 | 30.49–36.77 | 32.66 | 6 | 17.67–20.36 | 18.94 |
m2 | 6 | 32.13–39.20 | 36.03 | 5 | 19.00–21.22 | 20.18 |
m3 | 3 | 46.52–48.33 | 47.29 | 4 | 17.64–20.72 | 19.66 |
Mandible and lower dentition: two mandibles, DMR-KS-05-03-9-1 (Fig.
The i2 (DMR-KS-05-03-22-15) and i3 (DMR-KS-05-03-23-4) are spatulate and small, compared to other species of Bos (for measurements, see Tab.
Postcranial remains: a humerus, DMR-KS-05-03-20-2(1), preserves the shaft and distal part (Fig. L–N). We attribute this humerus to B. sauveli according to the proportional correlation with the extant specimens (Tab.
Southeast Asian large bovids are accurately identified by differences in cranial features (especially horn cores), although they show sexual and ontogenetic variation in morphology. Lacking the cranial remains, it is difficult to make a distinction within the species of Bos. Due to the lack of cranial remains of koupreys (B. sauveli) collected from Khok Sung, we identify these fossils on the basis of dental features.
Based on our comparisons with some extant specimens (
Scatter diagrams of lower cheek tooth (p2–m3) widths of recent and fossil large bovids. Fossil data from Phnom Loang, Lang Trang, Thum Wiman Nakin, Thum Prakai Phet, Duoi U’Oi, and Tam Hang South are from
According to the molecular phylogenetic analyses, the kouprey may have been domesticated in Cambodia (
A left horn core, DMR-KS-05-03-26-22; a right DP2, DMR-KS-05-03-20-4; two right P2—DMR-KS-05-03-19-27 and DMR-KS-05-04-03-3; a right DP3, DMR-KS-05-03-20-3; a right DP4, DMR-KS-05-03-17-3; a right M1, DMR-KS-05-03-00-20; a right M3, DMR-KS-05-03-17-1; a right mandible with m1–m3, DMR-KS-05-03-00-1; a left mandible with p2–m3, DMR-KS-05-04-3-1; a left i1, DMR-KS-05-03-00-27; two left m2—DMR-KS-05-03-19-26 and DMR-KS-05-03-16-1; two humeri—DMR-KS-05-05-1-1 (right) and DMR-KS-05-03-00-62 (left); a right metacarpus, DMR-KS-05-03-26-27; two left femora—DMR-KS-05-03-9-2 and DMR-KS-05-04-30-1 (proximal part).
Horn core: a single horn core (DMR-KS-05-03-26-22) is small, curved upward (Fig.
Remains of Bos gaurus from Khok Sung: A–B DMR-KS-05-03-26-22, a left horn core in posterior (A) and anterior (B) view C DMR-KS-05-03-20-1, a right DP2 D DMR-KS-05-03-19-27, a right P2 E DMR-KS-05-04-03-03, a right P2 F DMR-KS-05-03-20-3, a right DP3 G–HDMR-05-03-17-3, a right DP4 in occlusal (G) and lingual (H) views IDMR-05-03-17-1, a right M3 JDMR-05-03-19-26, a left m2 K–L DMR-KS-05-04-3-1, a left mandible in and occlusal (K) and lateral (L) views M DMR-KS-05-03-00-1, a fragmentary mandible in occlusal view. All isolated teeth are shown in occlusal view.
Upper dentition: DP2 (DMR-KS-05-03-20-4) is small and elongated, characterized by three main cones (anterior cone, paracone, and metacone) and a well-developed metastyle (Fig.
Mandibles and lower dentition: DMR-KS-05-04-3-1 is complete, posterior to the p2, with the exception of a small part of the angular region (Fig.
Postcranial remains: postcranial elements include humeri (Fig.
Postcranial remains of Bos gaurus from Khok Sung: A–D DMR-KS-05-05-1-1, a right humerus in proximal (A), posterior (B), anterior (C), and distal (D) views E–G DMR-KS-05-03-26-27, a right metacarpus in proximal (E), anterior (F), and distal (G) views H–J DMR-KS-05-03-9-2, a left femur in proximal (H), distal (I), and anterior (J) views.
According to
We assign the juvenile horn core (DMR-KS-05-03-26-22) to B. gaurus because the horn cores of gaurs are different from all other Bos species. They grow outward and curve upward, similar to those of Bubalus arnee, but their apical portion curves inward and slightly forward (
Mandibles and isolated teeth of B. gaurus are also observed. The cheek teeth of B. gaurus are distinguished from B. sauveli and B. javanicus by having two separate fossettes on the P2, more developed metaconids and more anteroposteriorly constricted postprotocristids on the p3 and p4, and more robust cheek teeth (Figs
A nearly complete cranium associated with a right mandible, DMR-KS-05-03-20-1; a cranium with a right tooth row (P3–M3), DMR-KS-05-03-21-1; a partial cranium with two tooth rows (P3–M1), DMR-KS-05-03-16-3; a partial cranium with a right tooth row (P3–M3), DMR-KS-05-03-11-1; three horn cores—DMR-KS-05-03-16-2 (right), DMR-KS-05-03-31-6 (right), and DMR-KS-05-03-19-28 (left); a left P2, DMR-KS-05-03-18-14; a left DP3, DMR-KS-05-03-00-103; two right P3—DMR-KS-05-03-22-14 and DMR-KS-05-04-05-3; a right DP4, DMR-KS-05-04-29-8 (broken anterior lobe); two P4—DMR-KS-05-03-18-13 (right) and DMR-KS-05-03-18-9 (left); four M1—DMR-KS-05-03-31-5 (right), DMR-KS-05-03-18-12 (right), DMR-KS-05-03-18-6 (left), and DMR-KS-05-03-22-13 (left); five M2—DMR-KS-05-03-00-2 (right), DMR-KS-05-03-25-21 (right), DMR-KS-05-03-18-5 (right), DMR-KS-05-03-16-2(1) (left), and DMR-KS-05-03-18-7 (left); four M3—DMR-KS-05-03-00-7 (right), DMR-KS-05-03-22-12 (left), DMR-KS-05-03-14-1 (left), and DMR-KS-05-03-18-10 (left); a right mandible with p2–m1, DMR-KS-05-03-20-2; three left mandibles—DMR-KS-05-03-10-3 (with p2–m3), DMR-KS-05-03-20-10 (with p2–m1), and DMR-KS-05-03-20-20 (with m1 and m2); a right i1, DMR-KS-05-03-18-8; a right i2, DMR-KS-05-03-22-15; a left i3, DMR-KS-05-03-00-106; a right i4, DMR-KS-05-03-16-3; a right p3, DMR-KS-05-03-14-4; a left dp4, DMR-KS-05-03-00-4; a right p4, DMR-KS-05-03-19-6; four m1—DMR-KS-05-03-25-3 (right), DMR-KS-05-03-18-18 (right), DMR-KS-05-03-00-105 (left), and DMR-KS-05-03-00-3 (left); two m2—DMR-KS-05-03-27-12 (right) and DMR-KS-05-03-25-2 (left); two m3—DMR-KS-05-03-18-11 and DMR-KS-05-04-29-2 (left posterior lobe); eleven thoracic vertebrae—DMR-KS-05-04-1-11 (T3), DMR-KS-05-04-1-26 (T4), DMR-KS-05-04-1-13 (T5), DMR-KS-05-04-1-14 (T6), DMR-KS-05-04-1-15 (T7), DMR-KS-05-04-1-16 (T8), DMR-KS-05-04-1-12 (T9), DMR-KS-05-04-1-17 (T10), DMR-KS-05-04-1-18 (T11), DMR-KS-05-04-1-19 (T12), and DMR-KS-05-04-1-20 (T13); four lumbar vertebrae—DMR-KS-05-04-1-24 (L1), DMR-KS-05-04-1-23 (L2), DMR-KS-05-04-1-22 (L3), and DMR-KS-05-04-1-21 (L4); two humeri—DMR-KS-05-03-31-1 (right) and DMR-KS-05-03-31-8 (left); two scapulae—DMR-KS-05-03-26-2 (right) and DMR-KS-05-02-20-4 (left); three ulnae and radii—DMR-KS-05-03-00-61 (right), DMR-KS-05-03-31-2 (right) and DMR-KS-05-03-31-9 (left); a right metacarpus, DMR-KS-05-03-26-3(1); a pelvis, DMR-KS-05-04-1-25; two femora—DMR-KS-05-04-1-1 (right) and DMR-KS-05-04-1-2 (left); a right fragmentary femur, DMR-KS-05-03-20-8 (distal part); three tibiae—DMR-KS-05-4-1-11 (right), DMR-KS-05-04-1-3 (left), and DMR-KS-05-03-20-9 (left); two fourth tarsal bones—DMR-KS-05-04-1-7 (right) and DMR-KS-05-04-1-5 (left); three metatarsi—DMR-KS-05-04-1-8 (right), DMR-KS-05-04-1-6 (left), and DMR-KS-05-03-28-30 (left); a left astragalus, DMR-KS-05-04-1-4; a left phalanx I, DMR-KS-05-04-1-9; a left phalanx II, DMR-KS-05-04-1-10.
Crania and upper dentition: DMR-KS-05-03-20-1 is undeformed and nearly complete (for measurements, see Appendix
Cranial and upper dental remains of Bubalus arnee from Khok Sung: A–C DMR-KS-05-03-20-1, a cranium in dorsal (A), ventral (B), and lateral (C) views and D–E DMR-KS-05-03-21-1, a cranium in dorsal (D) and ventral (E) views F–G DMR-KS-05-03-11-1, a right upper jaw in lateral (F) and occlusal (G) views H–I DMR-KS-05-03-16-3, a partial cranium in ventral view (H) with a right tooth row (I) J DMR-KS-05-03-16-2, a right horn core in dorsal view K DMR-KS-05-03-18-14, a left P2 L DMR-KS-05-03-00-103, a left DP3 M DMR-KS-05-04-29-8, a right DP4 N DMR-KS-05-03-00-7, a right M3. Cross-sections of basal horn cores are given. All isolated teeth are shown in occlusal view.
DMR-KS-05-03-21-1, a juvenile cranium, is incomplete but slightly deformed. The posterior part of the skull is almost complete but the anterior part is broken (Fig.
DMR-KS-05-03-11-1 preserves the right zygomatic bone and the premaxilla and maxilla with a nearly complete tooth row (P3–M3) (Fig.
Three isolated horn cores (DMR-KS-05-03-16-2: Fig.
Upper cheek teeth of Bubalus arnee are more robust, compared to those of Bos. P2 (DMR-KS-05-03-18-14: Fig.
Upper molars display Bos-like patterns (e.g., the degree of the hypsodonty and selenodonty and the presence of distinct styles) but are more robust than most species of Bos (e.g., B. sauveli and B. javanicus) (Tab.
Mandibles and lower dentition: five mandibles: DMR-KS-05-03-20-1 (Fig.
Mandibular and lower dental remains of Bubalus arnee from Khok Sung: A–B DMR-KS-05-03-20-1, a right mandible in lateral (A) and occlusal (B) views C–D DMR-KS-05-03-10-3, a left mandible in mesial (C) and occlusal (D) views E–FDMR-05-03-20-2, a right mandible in lateral (E) and occlusal (F) views G–HDMR-05-03-20-10, a left mandible in lateral (G) and occlusal (H) views I DMR-KS-05-03-20-20, a left fragmentary mandible with m1 and m2 in occlusal view J DMR-KS-05-03-18-8, a right i1 in lingual view; (K) DMR-KS-05-03-00-106, a left i3 in lingual view L DMR-KS-05-03-16-3, a right i4 M DMR-KS-05-03-00-4, a left dp4 in occlusal view N–O DMR-KS-05-03-00-105, a left m1 in occlusal (N) and buccal (O) views.
All lower cheek teeth are robust. All lingual stylids are distinct. The p2 has a well-developed postentocristid and posthypocristid (Fig.
Lower molars have well-developed stylids and conids. The metastylid is most developed on the unworn to slightly worn specimens (Fig.
Postcranial remains: postcranial elements include scapulae (Fig.
Articulated postcranial skeletons of Bubalus arnee from Khok Sung: A thoracic (abbreviated as “T”) vertebrae in lateral view: DMR-KS-05-04-1-11 (T3), DMR-KS-05-04-1-26 (T4), DMR-KS-05-04-1-13 (T5), DMR-KS-05-04-1-14 (T6), DMR-KS-05-04-1-15 (T7), DMR-KS-05-04-1-16 (T8), DMR-KS-05-04-1-12 (T9), DMR-KS-05-04-1-17 (T10), DMR-KS-05-04-1-18 (T11), DMR-KS-05-04-1-19 (T12), and DMR-KS-05-04-1-20, (T13) B lumbar (L) vertebrae in dorsal view: DMR-KS-05-04-1-24 (L1), DMR-KS-05-04-1-23 (L2), DMR-KS-05-04-1-22 (L3), and DMR-KS-05-04-1-21 (L4) C–E a left forelimb in anterior view: (C) DMR-KS-05-02-20-4, a scapula in lateral and distal views D DMR-KS-05-03-31-8, a humerus in proximal and distal views E DMR-KS-05-03-31-9, an ulna and a radius in proximal and distal views F DMR-KS-05-03-26-3(1), a right metacarpus in proximal, anterior, and distal views G DMR-KS-05-04-1-25, a pelvis in ventral view H–R hindlimbs in anterior view: H DMR-KS-05-04-1-1, a right femur in proximal and distal views I DMR-KS-05-4-1-11, a right tibia in proximal and distal views; (J) DMR-KS-05-04-1-7, a right 4th tarsal bone in dorsal view K DMR-KS-05-04-1-8, a right metatarsus in proximal and distal views L DMR-KS-05-04-1-2, a left femur M DMR-KS-05-04-1-3, a left tibia N DMR-KS-05-04-1-4, a left astragalus in plantar view O DMR-KS-05-04-1-5, a left 4th tarsal bone P DMR-KS-05-04-1-6, a left metatarsus Q DMR-KS-05-04-1-9, a left phalanx I in lateral view R DMR-KS-05-04-1-10, a left phalanx II in lateral view.
According to
Although the cheek teeth of Bos and Bubalus are almost morphologically identical and often show highly variable occlusal morphologies in relation to the wear stages, they are distinguishable based on the dental morphology.
As demonstrated by the scatter diagrams (Figs
A left M2, DMR-KS-05-03-18-16; three m3—DMR-KS-05-04-05-4 (right), DMR-KS-05-03-27-5 (left), and DMR-KS-05-03-28-10 (left posterior fragment).
Isolated teeth are almost complete (for measurements, see Tab.
Dental remains of Capricornis sumatraensis from Khok Sung: A–B DMR-KS-05-03-18-16, a left M2 in occlusal (A) and lingual (B) views C–D DMR-KS-05-04-05-4, a right m3 in occlusal (C) and buccal (D) views E–F DMR-KS-05-03-27-5, a left m3 in occlusal (E) and buccal (F) views G–H DMR-KS-05-03-28-10 in occlusal (G) and buccal (H) views.
We assign these isolated teeth from Khok Sung to Capricornis sumatraensis (Sumatran serow) because they are comparable in size and morphology to the extant specimens (Fig.
Scatter diagrams of M2 and m3 lengths and widths of recent and fossil serows and gorals. The measurements of fossil specimens from Lang Trang, Thum Wiman Nakin, Thum Prakai Phet, and Tam Hang South are from
Compared to other fossil records, C. sumatraensis from Khok Sung is smaller than that from the Late Pleistocene of Lang Trang in Vietnam (
A fragmentary cranium, DMR-KS-05-03-30-30; a dentary fragment with one tooth, DMR-KS-05-03-21-1; five isolated teeth—DMR-KS-05-03-00-19, DMR-KS-05-03-14-3, DMR-KS-05-03-22-22, DMR-KS-05-04-06-3, and DMR-KS-05-04-29-10; three osteoderms—DMR-KS-05-03-29-57, DMR-KS-05-03-29-58, and DMR-KS-05-03-27-25.
Skull and dentition: DMR-KS-05-03-30-30 is a slightly deformed skull preserving a nearly complete premaxilla, maxilla, nasal, and palatine process (Fig.
Remains of non-mammalian vertebrates from Khok Sung: Crocodylus cf. siamensis— A–B DMR-KS-05-03-30-30, a cranium in dorsal (A) and ventral (B) views C DMR-KS-05-03-21-1, a tooth in lingual view D–E DMR-KS-05-03-29-57 and F–G DMR-KS-05-03-27-25, osteoderms in dorsal (D, F) and ventral (E, G) views; Python sp.— H–I DMR-KS-05-03-00-16, a trunk vertebra in anterior (H) and ventral (I) views; Varanus sp. J–K DMR-KS-05-03-08-36, a trunk vertebra in anterior (J) and ventral (K) views; Galliformes indet.— L–M DMR-KS-05-04-05-40, a cervical vertebra fragment in dorsal (L) and ventral (M) views; Siluridae indet.— N–O DMR-KS-05-03-22-76, a vertebra in anterior (N) and lateral (O) views P DMR-KS-05-04-11-20, a pectoral spine in dorsal view Q DMR-KS-05-04-05-25, a pectoral spine in medial view. Anatomical abbreviations: al, alveolus; pmx, premaxilla; en, external naris; n, nasal; mx, maxilla; pal, palatine; palp, palatine process.
Osteoderms: two nearly complete specimens (Fig.
The specimen DMR-KS-05-03-30-30 is a crocodilian cranium with a possible maximum length up to 50 cm. All morphological characters of the Khok Sung crocodiles are congruent with the extant fresh water crocodile, Crocodylus siamensis, as well as with its fossils recovered from the Early and Middle Pleistocene of Java (Trinil H. K., Kedung Brubus, and Kedung Lumbu) (
Four trunk vertebrae—DMR-KS-05-03-00-21, DMR-KS-05-03-00-16 (two attached vertebrae), and DMR-KS-05-04-28-12.
Vertebrae are almost complete and represent a large-sized snake (for measurements, see Tab.
Measurements (in millimeters) of vertebrae of Python and Varanus from Khok Sung. Abbreviations: CL, centrum length (measured at the ventral midline); H, maximum height (measured from the tip of the neural spine to the ventral rim of the cotyle); WPP, width between pre- and postzygapophyseal processes; Wpre, width across zygapophyseal processes; Wpost, width across postzygapophyseal processes; Wcd, width of the condyle; Hcd, height of the condyle (measured from the dorsal to ventral rim); Wct, width of the cotyle; Hct, height of the cotyle. + refers to the measurement of two attached vertebrae and * indicates an incomplete preservation.
CL | H | WPP | Wpre | Wpost | WCd | Hcd | WCd | Hct | |
---|---|---|---|---|---|---|---|---|---|
Python sp. | |||||||||
DMR-KS-05-03-00-21 | 20.85 | 40.36 | 22.47 | 36.48 | 15.27 | 14.03 | 13.95 | 12.87 | 15.12 |
DMR-KS-05-03-00-16 | 28.75+ | 26.35* | 23.82 | 35.23 | 13.73 | – | 12.04 | – | 16.44 |
DMR-KS-05-04-28-12 | 14.06 | 17.69* | 17.18 | 20.46 | 23.64 | 6.50 | 6.80 | 6.62 | 7.82 |
Varanus sp. | |||||||||
DMR-KS-05-03-29-36 | 24.98 | 25.39* | 27.90 | 21.91 | 34.98 | 7.18 | 9.21 | 18.96 | 22.09 |
DMR-KS-05-03-08-36 | 31.73 | 28.56 | 34.11 | 36.21 | 35.60 | 7.82 | 12.68 | 18.27 | 21.91 |
These four vertebrae are attributed to the family Boidae because of the following characters: a short, wide, and massive vertebral body (i.e., the widths of the centra are greater than the lengths, sensu
Two trunk vertebrae—DMR-KS-05-03-08-36 and DMR-KS-05-03-29-36.
The vertebra DMR-KS-05-03-08-36 is more complete than the specimen DMR-KS-05-03-29-36 (for measurements, see Tab.
We assign these two vertebrae to the the family Varanidae due to the following morphological characters: a centrum tapering posteriorly, a precondylar constriction, a ventrally facing cotyle, and a large and flared condyle (
Nine taxa: seven Testudines, an extinct gharial (Gavialisbengawanicus), and a spotted hyaena (Crocuta crocuta ultima), have been previously described from Khok Sung by
Mammalia | ||
Proboscidea | ||
Stegodontidae | ||
Stegodon cf. orientalis | ||
Elephantidae | ||
Elephas sp. | ||
Perissodactyla | ||
Rhinocerotidae | ||
Rhinoceros sondaicus | ||
Rhinoceros unicornis | ||
Artiodactyla | ||
Bovidae | ||
Bos sauveli | ||
Bos gaurus | ||
Bubalus arnee | ||
Capricornis sumatraensis | ||
Cervidae | ||
Axis axis | ||
Panolia eldii | ||
Rusa unicolor | ||
Suidae | ||
Sus barbatus | ||
Primates | ||
Cercopithecidae | ||
Macaca sp. | ||
Carnivora | ||
Hyaenidae | ||
Crocuta crocuta ultima (identified by |
||
Canidae | ||
Cuon sp. | ||
Reptilia | ||
Testudines (identified by |
||
Geoemydidae | ||
Batagur cf. trivittata | ||
Heosemys annandalii | ||
Heosemys cf. grandis | ||
Malayemys sp. | ||
Trionychidae | ||
Chitra sp. | ||
cf. Amyda sp. | ||
Crocodilia | ||
Gavialidae | ||
Gavialis cf. bengawanicus (identified by |
||
Crocodylidae | ||
Crocodylus cf. siamensis | ||
Squamata | ||
Varanidae | ||
Varanus sp. | ||
Boidae | ||
Python sp. | ||
Actinopterygii | ||
Siluridae indet. | ||
Aves | ||
Galliformes indet. |
According to the fact that Khok Sung yields only large mammals (> 8 kg), the absence of medium- and small-sized mammal remains is likely due to taphonomic conditions and/or fossil collecting methods. Similarly to most of the Middle and Late Pleistocene fossil sites in Southeast Asia, the biodiversity of Khok Sung large mammals is likely greater than that of present-day faunas (see Appendices
Past records and recent distribution patterns of large mammalian species present in Khok Sung are revealed in this work. Paleontological sites in Southeast Asia as well as South China are examined for the Early, Middle, and Late Pleistocene, compared with the modern distribution patterns. We only focus on mammalian taxa assigned to the species-level, including Stegodon orientalis and its co-occuring species, Rhinoceros sondaicus, Rhinoceros unicornis, Sus barbatus, Axis axis, Panolia eldii, Rusa unicolor, Bos sauveli, Bos gaurus, Bubalus arnee, and Capricornis sumatraensis.
The earliest records of derived Stegodon (e.g., Stegodon orientalis from Dayakou (
A fossil species of Palaeoloxodon is reported from several Middle Pleistocene localities in mainland Southeast Asia (Fig.
The Middle (red) and Late (yellow) Pleistocene records of stegodontids and relative fossil elephants, and the current distribution (green) of Elephas maximus (Indian elephant). Stars indicate the co-occurrence of sympatric proboscideans. The current distribution of Indian elephants is compiled from
Elephas maximus is known from the late Middle Pleistocene of Thum Wiman Nakin (northeastern Thailand) (
The Early Pleistocene records of Asian rhinoceroses are poorly documented in Southeast Asia. Only R. sondaicus is reported from the upper part of the Irrawaddy Formation, near Pauk Township in central Myanmar (
The Early (blue circle), Middle (red circle), and Late (yellow circle) Pleistocene records and the current distribution (green) of Rhinoceros sondaicus (Javan rhinoceros). The current distribution of the species is compiled from
The Middle Pleistocene record, especially the late Middle Pleistocene, includes numerous reports of Asian rhinoceroses (Figs
The Middle (red circle) and Late (yellow circle) Pleistocene records and the current distribution (green) of Rhinoceros unicornis (Indian rhinoceros). “?” indicates the possible record of R. unicornis according to
During the late Pleistocene, Javan and Indian rhinoceroses were widespread in Indochinese subregion (Figs
Nowadays, the Indian rhinoceros is locally extinct from the Thai territory and several other countries in Southeast Asia. The species is restricted to Nepal and India and some parts of northernmost Myanmar (
During the Middle Pleistocene, Sus barbatus (bearded pig) is known from the caves of Thum Wiman Nakin and Thum Prakai Phet (
In the late Pleistocene, S. barbatus is well-documented from many localities, extending its geographic distribution across Sumatra, Borneo, and Java. This species is likely more widespread in the late Pleistocene than the Middle Pleistocene (Fig.
Today S. barbatus is restricted to Peninsular Malaysia, Sumatra, and Borneo (
Fossils of Axis axis (chital) have never been previously recorded from Thailand but were present in mainland Southeast Asia, at least in Khok Sung, during the late Middle Pleistocene (Fig.
Nowadays Axis axis is restricted to the Indian Subcontinent (India, Nepal, Sikkim, and Sri Lanka) (Fig.
The Eld’s deer is known from the Middle Pleistocene of Thailand. Fossils of P. eldii are collected from the caves of Thum Wiman Nakin and Kao Pah Nam (
During the Late Pleistocene, the Eld’s and sambar deer co-occurred in the Cave of the Monk (Ban Fa Suai), northern Thailand (
Nowadays, Panolia eldii is restricted to the Indochinese province (Fig.
Large bovids in Southeast Asia currently comprise four wild species: Bos sauveli (kouprey), Bos javanicus (banteng), Bos gaurus (gaur), and Bubalus arnee (wild water buffalo). Bantengs, gaurs, and koupreys presumably shared a common ancestor at 2.6 Ma (Plio-Pleistocene) and their lineages split in a short period of time (i.e., between 200 and 300 ka) based on the molecular estimations of divergence times (
During the Late Pleistocene, the locality of the Cave of the Monk (Ban Fa Suai) yielded remains of these bovid species (cf.) (
The historical distribution of koupreys during the last century is restricted to Cambodia, southern Laos, southeastern Thailand, and western Vietnam (
Overall, the Pleistocene large bovid species in Southeast Asia is more widespread than the modern population. The anthropogenic impacts on the environments and landscapes seem to have caused the reduction of large bovid population in several areas during the past decade. The koupreys is more widely distributed during the Pleistocene than today (Fig.
The possible earliest records of Capricornis sumatraensis are from the middle Early Pleistocene site of Gongwangling (
The Sumatran serow is a widespread species, native to mountain forests on the Himalayan range (northern India, Sikkim, and Nepal) of the Indochinese subregion (Southern China, Myanmar, Thailand, Laos, Cambodia, Vietnam, and Peninsular Malaysia) and on the island of Sumatra (
For the comparisons of vertebrate faunas between Khok Sung and other Pleistocene sites, we focus only on large mammals (for the mammalian fauna lists of the Middle to Late Pleistocene of Southeast Asian sites, see Appendices
The Khok Sung assemblage shares at least one similar archaic mammal taxon such as Crocuta crocuta ultima and Stegodon orientalis, with these faunas. Crocuta crocuta is also recorded from Thum Wiman Nakin, Thum Prakai Phet, and Nam Lot, whereas Stegodon orientalis is reported from two Laotian sites: Nam Lot and Tam Hang South. By the way, most of forest dwelling and carnivorous taxa that are representatives of Middle Pleistocene mammalian assemblages such as Ailuropoda melanoleuca (giant panda), Ursus thibetanus (Asiatic black bear), Pongo pygmaeus (orang-utan), Muntiacus muntjak (Southern red muntjac), and Tapirus indicus (Malayan tapir) are absent in Khok Sung. The paleoenvironments of Khok Sung corresponded to a floodplain near the river channel (
The degree of the faunal similarity also depends on the number of identified taxa for each site. We further analyse the relationships between the geographic regions and faunas in Southeast Asia, using the Simpson coefficient of faunal similarity (Tab.
Similarity matrix based on the Simpson coefficients. Locality abbreviations: YCK, Yenchingkou; KLS, Koloshan; DX, Daxin; HJ, Hejiang; GX, Ganxian; PXDD, Panxian Dadong; WY, Wuyun; MB, Maba; HST, Hoshantung; KS, Khok Sung; TWN ; Thum Wiman Nakin; TPKP, Thum Phra Khai Phet; TK, Tham Khuyen; TO, Tham Om; BDB, Boh Dambang; KDBB, Kedung Brubus; TNHK, Trinil Hauptknochenschicht; ND, Ngandong.
YCK | KLS | DX | HJ | GX | PXDD | WY | MB | HST | KS | TWN | TPKP | TK | TO | BDB | KDBB | TNHK | ND | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
YCK | 1.00 | |||||||||||||||||
KLS | 0.38 | 1.00 | ||||||||||||||||
DX | 0.54 | 0.31 | 1.00 | |||||||||||||||
HJ | 0.55 | 0.27 | 0.45 | 1.00 | ||||||||||||||
GX | 0.50 | 0.20 | 0.40 | 0.40 | 1.00 | |||||||||||||
PXDD | 0.53 | 0.46 | 0.46 | 0.55 | 0.30 | 1.00 | ||||||||||||
WY | 0.53 | 0.23 | 0.38 | 0.45 | 0.70 | 0.33 | 1.00 | |||||||||||
MB | 0.94 | 0.38 | 0.62 | 0.55 | 0.50 | 0.44 | 0.47 | 1.00 | ||||||||||
HST | 0.50 | 0.30 | 0.20 | 0.30 | 0.20 | 0.40 | 0.40 | 0.50 | 1.00 | |||||||||
KS | 0.50 | 0.00 | 0.08 | 0.18 | 0.10 | 0.25 | 0.17 | 0.25 | 0.10 | 1.00 | ||||||||
TWN | 0.48 | 0.15 | 0.31 | 0.27 | 0.60 | 0.24 | 0.40 | 0.50 | 0.30 | 0.83 | 1.00 | |||||||
TPKP | 0.58 | 0.17 | 0.17 | 0.36 | 0.30 | 0.33 | 0.33 | 0.50 | 0.30 | 0.50 | 0.92 | 1.00 | ||||||
TK | 0.63 | 0.31 | 0.54 | 0.55 | 0.60 | 0.41 | 0.40 | 0.63 | 0.40 | 0.33 | 0.53 | 0.42 | 1.00 | |||||
TO | 0.94 | 0.38 | 0.54 | 0.55 | 0.50 | 0.50 | 0.47 | 0.81 | 0.40 | 0.42 | 0.63 | 0.50 | 0.75 | 1.00 | ||||
BDB | 0.80 | 0.10 | 0.20 | 0.20 | 0.30 | 0.40 | 0.50 | 0.60 | 0.20 | 0.40 | 0.80 | 0.60 | 0.60 | 0.60 | 1.00 | |||
KDBB | 0.11 | 0.08 | 0.00 | 0.09 | 0.00 | 0.06 | 0.07 | 0.13 | 0.10 | 0.17 | 0.22 | 0.17 | 0.06 | 0.13 | 0.10 | 1.00 | ||
TNHK | 0.21 | 0.08 | 0.08 | 0.09 | 0.00 | 0.14 | 0.14 | 0.21 | 0.10 | 0.08 | 0.14 | 0.17 | 0.07 | 0.14 | 0.30 | 0.64 | 1.00 | |
ND | 0.10 | 0.10 | 0.00 | 0.10 | 0.00 | 0.10 | 0.10 | 0.10 | 0.10 | 0.00 | 0.10 | 0.00 | 0.00 | 0.10 | 0.00 | 0.90 | 0.60 | 1.00 |
As a result, the Middle Pleistocene Southeast Asian taxa reveal two distinct associations (Javanese and mainland Southeast Asian faunas) (Fig.
Overall, this analysis suggests initially that the differences in species composition and distribution do not follow a trend of the latitudinal gradient north to south, but show spatial and time variability of large mammalian fauna in Southeast Asia. The main problems of mammalian fauna comparisons in Southeast Asia are likely due to the poorly-known species diversity and/or the imprecisely chronological determination in several localities.
According to the similarity analysis of the fauna, the mammalian fauna composition of Khok Sung is considerably different from the Early to early Middle Pleistocene assemblage of Java. This suggests an inconsistent age of the Early Pleistocene for Khok Sung. The Khok Sung assemblage is highly comparable in composition to three late Middle Pleistocene faunas: Thum Wiman Nakin (> 169 ka,
On the basis of the previous paleomagnetic data analysed by
Relationships of the Khok Sung vertebrate fauna for dispersal events from India to Java has been first proposed by
However, in terms of faunal age, this scenario is no longer consistent because the Khok Sung fauna is recently attributed to a late Middle Pleistocene age (
Based on the occurrence of mammalian taxa in Khok Sung, we suggest an alternative relevance of this fauna for the “Sino-malayan” dispersal events from mainland Southeast Asia to Java (Fig.
On the other hand, the Khok Sung fauna lacks any evidence of taxa originating from Java. But the possible presence of Duboisia santeng in Tambun site (Peninsular Malaysia) may indicate the faunal exchange from Indonesia to the mainland Southeast Asia (
The Khok Sung mammalian assemblage supports that Thailand was a biogeographic gateway of the Sino-Malayan migration event as the mainland forested faunal association replaced the earlier Siva-Malayan fauna (Stegodon-Homo erectus complex) subsequently in Java (
We would like to thank Joséphine Lesur (
Measurements (in millimeters) of postcranial remains of identified mammal taxa from Khok Sung. * indicates a juvenile individual.
Scapula | |||||||||||||||
Specimen | Taxa | GLP | LG | SLC | BG | Ld | DHA | HS | |||||||
DMRKS-05-03-00-58 | Rhinoceros sondaicus | 87.49 | 78.81 | 62.97 | 55.33 | 144.77 | – | 325.57 | |||||||
DMR-KS-05-03-26-2 | Bubalus arnee | 106.72 | 81.99 | 76.45 | 68.40 | 297.28 | 381.27 | 445.28 | |||||||
DMR-KS-05-02-20-4 | Bubalus arnee | 84.00 | 66.89 | 52.13 | 54.50 | 213.22 | 301.79 | 296.17 | |||||||
DMR-KS-05-06-24-4 | Panolia eldii | 44.00 | 34.56 | 23.41 | 31.54 | 108.17 | 206.05 | 211.37 | |||||||
Humerus | |||||||||||||||
Specimen | Taxa | Bp | Dp | Bd | BT | Dd | SD | GLC | GLl | GL | |||||
DMR-KS-05-03-10-6 | Stegodon cf. orientalis | – | – | 195.27 | 173.98 | 162.73 | – | – | – | – | |||||
DMR-KS-05-03-31-3 | Rhinoceros sondaicus | 174.58 | 142.51 | 155.86 | 94.39 | 115.77 | 66.09 | 377.04 | 422.18 | 437.23 | |||||
DMR-KS-05-03-26-8 | Sus barbatus | 65.67 | 76.44 | 57.39 | 47.06 | 45.11 | 20.72 | 224.71 | 236.13 | 238.28 | |||||
DMR-KS-05-03-20-2(1) | Bos sauveli | > 99.80 | – | 88.14 | 84.92 | 88.80 | 38.86 | – | – | – | |||||
DMR-KS-05-03-00-62 | Bos gaurus | > 99.45 | > 98.85 | 103.69 | 98.14 | 96.59 | 50.36 | 328.18 | 347.28 | 353.87 | |||||
DMR-KS-05-05-1-1 | Bos gaurus | 122.37 | 130.12 | 103.54 | 98.77 | 97.16 | 56.37 | 317.26 | 354.28 | 356.28 | |||||
DMR-KS-05-03-31-1 | Bubalus arnee | 125.53 | 135.07 | 104.40 | 101.70 | 93.72 | 52.29 | 343.74 | 413.22 | 444.12 | |||||
DMR-KS-05-03-31-8 | Bubalus arnee | 125.79 | 138.19 | 104.38 | 99.09 | 94.07 | 52.92 | 347.41 | 424.77 | 449.11 | |||||
DMR-KS-05-03-13-4 | Axis axis | – | – | 34.75 | 31.96 | 34.12 | < 17.02 | – | – | – | |||||
DMR-KS-05-04-11-32 | Axis axis | – | – | 35.98 | 33.77 | 34.05 | 17.37 | – | – | – | |||||
DMR-KS-05-03-17-17 | Axis axis | – | – | 36.86 | 35.57 | 32.77 | 16.10 | – | – | – | |||||
DMR-KS-05-04-11-35 | Panolia eldii | – | > 59.10 | 46.10 | 40.85 | 41.18 | 23.3 | 187.53 | – | – | |||||
DMR-KS-05-03-18-1 | Panolia eldii | 53.34 | 65.34 | – | – | – | 21.45 | – | – | – | |||||
DMR-KS-05-03-15-43 | Rusa unicolor | – | – | 54.89 | 49.04 | 48.30 | < 27.70 | – | – | – | |||||
Ulna and radius | |||||||||||||||
Specimen | Taxa | Bp/BPC | BFp | Dp | Bd | BFd | Dd | LO | DPA | SDO | SD | PL | Ll | GLl | GL |
DMR-KS-05-03-00-61 | Bubalus arnee | 106.16 | 92.17 | 52.99 | 100.36 | 90.17 | 65.85 | 125.53 | 93.64 | 72.61 | 55.54 | 305.18 | 308.75 | 476.61 | 484.72 |
DMR-KS-05-03-31-2 | Bubalus arnee | 108.45 | 98.91 | 57.03 | 103.37 | 92.25 | 73.75 | 131.15 | 98.37 | 76.03 | 50.74 | 335.29 | 343.51 | 427.78 | 452.17 |
DMR-KS-05-03-31-9 | Bubalus arnee | 106.69 | 97.99 | 57.44 | 103.49 | 92.06 | 72.92 | 128.89 | 99.00 | 76.54 | 47.66 | 335.93 | 344.77 | 424.77 | 449.11 |
DMR-KS-05-03-31-10 | Panolia eldii | 39.40 | 36.72 | 20.95 | 37.22 | 32.67 | 21.37 | – | – | – | 22.52 | 199.23 | 198.21 | – | 197.52 |
DMR-KS-05-04-11-3 | Panolia eldii | 42.30 | 38.22 | 22.47 | 36.84 | 34.64 | 23.46 | – | – | – | 22.61 | 204.18 | 203.71 | – | 215.70 |
DMR-KS-05-03-19-16 | Panolia eldii | 40.16 | 33.03 | 22.16 | 40.62 | 39.21 | 29.89 | – | – | – | 23.58 | 197.89 | 193.72 | – | 204.53 |
DMR-KS-05-03-25-9 | Rusa unicolor | 55.06 | 53.58 | 28.92 | – | – | – | – | – | – | – | – | – | – | – |
DMR-KS-05-03-19-14 | Rusa unicolor | 52.37 | 50.16 | 30.83 | – | – | – | – | – | – | 28.12 | – | – | – | – |
DMR-KS-05-03-26-19 | Rusa unicolor | – | – | – | 43.12 | 41.08 | 32.20 | – | – | – | 24.83 | – | – | – | – |
Pelvis | |||||||||||||||
Specimen | Taxa | GL | LA | LS | SH | SB | SC | LFo | GBTc | GBA | GBTi | SBI | |||
DMR-KS-05-03-10-11 | Stegodon cf. orientalis | > 855.79 | 148.93 | – | 139.18 | 66.29 | 399.46 | 206.77 | – | – | – | – | |||
DMR-KS-05-03-10-12 | Stegodon cf. orientalis | > 496.75 | 140.75 | – | 145.17 | 59.60 | 389.92 | 201.25 | – | – | – | – | |||
DMR-KS-05-04-1-25 | Bubalus arnee | 494.85 | 96.57 | 149.23 | 70.55 | 35.81 | 256.67 | 103.24 | 517.48 | 303.98 | 319.25 | 204.37 | |||
Femur | |||||||||||||||
Specimen | Taxa | Bp | Dp | DC | Bd | Dd | SD | GLC | GL | ||||||