TABLE 1. Mammalian faunal list for bed TB33, How Ledge Limestone, SW Headon Hill. Asterisk (*) indicates presence of species in the samples analysed herein. (L) indicates species larger than 1 kg, so not micromammals. NISP = number of identified specimens; MNI = minimum number of individuals. Artiodactyls are not typically micromammals, but tiny Mouillacitherium belongs in this category and is included.
Order | Family | Species | NISP | MNI | Family NISP |
Family MNI |
Didelphimorphia (primitive marsupials) |
Herpetotheriidae | * Amphiperatherium species A | 63 | 8 | 343 | 19 |
* Amphiperatherium species B | 67 | 7 | ||||
* Amphiperatherium species C | 29 | 3 | ||||
* Peratherium cuvieri | 1 | 1 | ||||
Rodentia | “Paramyidae” | * Plesiarctomys gervaisii (L) | 1 | 1 | 1 | 1 |
Gliridae | Gliravus daamsi | 0 | 0 | 179 | 13 | |
* Miniglis minor | 46 | 4 | ||||
* Bransatoglis bahloi | 11 | 3 | ||||
* Glamys priscus | 77 | 6 | ||||
Pseudosciuridae | * Sciuroides ehrensteinensis | 1 | 1 | 166 | 15 | |
* Suevosciurus bosmae | 94 | 8 | ||||
* Treposciurus mutabilis | 12 | 3 | ||||
Treposciurus gardneri | 0 | 0 | ||||
* Tarnomys quercyi vectisensis | 18 | 3 | ||||
Theridomyidae | * Thalerimys fordi | 782 | 77 | 1050 | 79 | |
* Isoptychus sp. | 13 | 2 | ||||
Lipotyphla | Talpidae | * Eotalpa anglica | 36 | 5 | 36 | 5 |
Amphilemuridae | * Gesneropithex latidens | 51 | 3 | 51 | 3 | |
Chiroptera | Hipposideridae | * Pseudorhinolophus sp. | 2 | 1 | 2 | 1 |
Palaeochiropterygidae | * Stehlinia sp. | 1 | 1 | 1 | 1 | |
Stem Euarchonta | Nyctitheriidae | * Saturninia gracilis | 64 | 8 | 663 | 44 |
* Cryptotopos woodi | 35 | 4 | ||||
* Cheilonyctia lawsoni | 221 | 20 | ||||
Scraeva hatherwoodensis | 0 | 0 | ||||
* Euronyctia curranti. | 92 | 12 | ||||
Primates | Adapidae | * Magnadapis sp. (L) | 1 | 1 | 1 | 1 |
Adapis stintoni (L) | 0 | 0 | ||||
Omomyidae | * Pseudoloris parvulus | 6 | 2 | 28 | 6 | |
* Vectipithex smithorum | 14 | 3 | ||||
* Microchoerus erinaceus | 5 | 1 | ||||
Pantolesta | Pantolestidae | * Cryptopithecus major | 32 | 3 | 32 | 3 |
Apatotheria | Apatemyidae | * Heterohyus nanus | 6 | 2 | 6 | 2 |
Heterohyus sp. | 0 | 0 | ||||
Carnivora | “Miacidae” | * Paramiacis sp. | 61 | 2 | 61 | 2 |
Artiodactyla | Anthracotheriidae | Diplopus aymardi (L) | 0 | 0 | 0 | 0 |
Xiphodontidae | Dichodon cervinus (L) | 0 | 0 | 0 | 0 | |
Dichobunidae | * Mouillacitherium sp. | 1 | 1 | 1 | 1 | |
Perissodactyla | Palaeotheriidae | Plagiolophus annectens (L) | 0 | 0 | 0 | 0 |
Palaeotherium muehlbergi (L) | 0 | 0 | ||||
Indet. | Indet. | Indet. | 377 | |||
Totals: | 16 in sample | 30 in sample | 1843 | 196 | 2998 | 196 |
TABLE 2. Quantification of surface modifications on mammalian skeletal elements by family and, in the case of Chiroptera, by order.
Theridomyidae | Pseudosciuridae | Gliridae | Herpetotheriidae | Nyctitheriidae | Omomyidae | Talpidae | Amphilemuridae | Apatemyidae | Pantolestidae | “Miacidae” | Chiroptera | Dichobunidae | |
Number of identified bones (NISP bones) | 2711 | 452 | 86 | 78 | 4 | 0 | 0 | 0 | 0 | 49 (4)5 |
0 | 0 | |
Number of identified teeth (NISP teeth) | 7333 | 1143 | 893 | 347 | 585 | 24 (6)5 |
35 | 18 | 6 | 32 | 15 (5)5 |
3 | 1 |
Minimum number of complete bones (N bones) | 2371 | 37 | 52 | 30 | 1 | 0 | 0 | 0 | 0 | 15 | 0 | 0 | |
Minimum number of complete teeth (N teeth) | 561 (733)3 |
1143 | 893 | 257 | 585 | 24 (6)5 |
25 | 18 | 3 | 32 | 15 (5)5 |
3 | 1 |
Skeletal average relative abundance4 | 2.81 | 2.9 | 2.6 | - | - | - | - | - | - | - | - | - | |
Total extent of bone fragmentation % | 571 | 49 | 68 | 62 | 75 | - | - | - | - | 69 | - | - | |
Percentage of etched teeth | 543 | 613 | 323 | 20 | 18 | 42 (92)5 |
20 | 18 | 67 | 22 | 40 | 33 | 100 |
Percentage of etched bones | 421 | 47 | 47 | 28 | 25 | 100 | - | - | - | 70 | - | - | |
Puncture marks (by Paramiacis) | yes | no | no | no | no | no | no | no | no | no | no | no | no |
Chipping | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | no |
Cracking | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | no |
Gnaw marks (by glirids) | yes | no | no | no | no | no | no | no | no | no | no | no | no |
Parallel grooves (by insects?) | yes | no | yes | no | yes | no | no | no | no | no | no | no | no |
Transverse grooves (by nyctitheres?) | yes | yes | yes | yes | yes | yes | no | yes | no | yes | yes | yes | no |
Percentage of weathered elements | 18 | 15 | 10 | 10 | 5 | 11 | 0 | 0 | 0 | 0 | 5 | 0 | 0 |
Fine striations (by trampling?) | yes | no | no | no | yes | no | no | no | no | no | yes | no | no |
Rounding or polishing | no | no | no | no | no | no | no | no | no | no | no | no | no |
1 Includes doubtful theridomyid bones, which may include Pseudosciuridae.
2 Includes doubtfully attributed to family.
3 Cheek teeth only, as some incisors of Theridomyidae and Pseudosciuridae may be confused; figure in parentheses is with the incisors included. Gliridae are treated similarly for consistency within Rodentia.
4 Average relative abundance is mean of relative abundances of all elements except teeth, incus and malleus.
5 Figures in parentheses are MNI, recognizing that nine of the omomyid teeth belong to one individual of Vectipithex smithorum and that 11 of the teeth and 46 of the bones of Paramiacis sp. (“Miacidae”) belong to a single individual.
TABLE 3. Counts of the minimum number of complete specimens (Ni) of all the skeletal elements and the Relative Abundances for Thalerimys fordi from bed TB33, How Ledge Limestone. Column 2: number of elements in a single theridomyid skeleton (based on Schmidt-Kittler and Storch, 1985); vertebral number includes the four sacrals and the phalangeal and metapodial number excludes the manual ray I, which is greatly reduced in Theridomyidae. Column 3: number of T. fordi specimens expected, based on the MNI of 77 for the species. Column 4: total Ni (Thalerimys + ? Thalerimys + ?? Thalerimys + “possible Theridomyidae”). Column 5: % Relative Abundance (Ri) of total in relation to T. fordi expected [(column4/column3)*100]; the average skeletal Relative Abundance appears in the table as “Total” Relative Abundance and equals the mean value of the abundances of all the skeletal elements (excluding teeth).
Skeletal element | N elements T. fordi |
N T. fordi expected |
Total Ni | % Ri of total T. fordi expected |
Dentary | 2 | 154 | 2 | 1.3% |
Maxilla | 2 | 154 | 6 | 3.9% |
Vertebrae | 53 | 4081 | 0 | 0% |
Ribs | 24 | 1848 | 0 | 0% |
Scapula | 2 | 154 | 1 | 0.6% |
Humerus | 2 | 154 | 12 | 7.8% |
Radius | 2 | 154 | 11 | 7.1% |
Ulna | 2 | 154 | 16 | 10.4% |
Carpals | 18 | 1386 | 2 | 0.1% |
Pelvis | 2 | 154 | 0 | 0% |
Femur | 2 | 154 | 5 | 3.2% |
Tibia | 2 | 154 | 3 | 1.9% |
Fibula | 2 | 154 | 1 | 0.6% |
Astragalus | 2 | 154 | 8 | 5.2% |
Calcaneum | 2 | 154 | 3 | 1.9% |
Other tarsals | 10 | 770 | 11 | 1.4% |
Metapodials | 18 | 1386 | 24 | 1.7% |
Phalanges | 52 | 4004 | 132 | 3.3% |
Total | 199 | 15323 | 237 | 2.8% |
TABLE 4. Counts and percentages of the total for each of the six limb bones in each breakage class for the total theridomyid and possible theridomyid material from bed TB33, How Ledge Limestone.
Total = Thalerimys + ? Thalerimys + ?? Thalerimys + possible Theridomyidae | |||||||||||||||||
Humerus | Ulna | Radius | Femur | Tibia | Fibula | ||||||||||||
Breakage class | N | % | N | % | N | % | N | % | N | % | N | % | |||||
Complete | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | |||||
Proximal | 0 | 0 | 3 | 19 | 11 | 100 | 5 | 100 | 0 | 0 | 0 | 0 | |||||
Distal | 11 | 100 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 100 | 1 | 100 | |||||
Shaft | 0 | 0 | 13 | 81 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | |||||
Total | 11 | 100 | 16 | 100 | 11 | 100 | 5 | 100 | 2 | 100 | 1 | 100 |
TABLE 5. Percentages of breakage types, including rounding by etching, of elongate bones for common micromammalian families in bed TB33, How Ledge Limestone. Numbers in parentheses have been calculated by dividing the percentages of each breakage category per family by the square root of the sample number in each category per family. They are used in the bars in Figures 3 and 10.
Gliridae | Herpetotheriidae | Theridomyidae | Nyctitheriidae | |
Total broken elongate bones (N) | 21 | 49 | 131 | 23 |
Spiral irregular breakage (N) % of broken modified when fresh |
7 33% (12.2%) |
25 51% (10.2%) |
65 50% (6.2%) |
6 26% (10.4%) |
Perpendicular smooth breakage-dry (N) % of broken modified when dry |
13 62% (17.2%) |
24 49% (10.0%) |
4 3% (1.5%) |
11 48% (14.6%) |
Broken elongate with rounded edges (N) % of broken with rounded edges |
6 29% (11.6%) |
13 27% (7.5%) |
35 27% (6.9%) |
4 17.4% (8.7%) |
TABLE 6. Percentages of broken flat, elongate and compact bones for three most common micromammalian families in bed TB33, How Ledge Limestone. Numbers in parentheses have been calculated as in Table 5.
Gliridae | Herpetotheriidae | Theridomyidae | |
Total flat | 0 | 9 | 9 |
Broken flat % broken flat |
0 0% (0%) |
9 100% (33.3%) |
9 100% (33.3%) |
Total elongate | 40 | 66 | 207 |
Broken elongate % broken elongate |
21 53% (11.5%) |
49 74% (10.6%) |
131 63% (5.5%) |
Total compact | 5 | 10 | 55 |
Broken compact % broken compact |
1 20% (20%) |
0 0% (0%) |
14 26% (7.0%) |
Total bone fragmentation degree | 49% (10.4%) | 68% (9.0%) | 57% (4.6%) |
TABLE 7. Percentage of etched teeth (including the incisors of the rodents) of Theridomyidae, Gliridae, Herpetotheriidae and Nyctitheriidae and of Theridomyidae + Herpetotheriidae (semiterrestrial) and Gliridae + Nyctitheriidae (scansorial) in bed O3 (“Os1” of Vasileiadou et al., 2007a, 2009), Osborne Member and bed TB33, How Ledge Limestone. Percentages in the last column have been calculated as in Table 5.
Total | Affected | % of total |
% sq root |
|
Theridomyidae | ||||
O3 | 785 | 505 | 64% | 2.8% |
TB33 | 608 | 327 | 54% | 3.0% |
Gliridae | ||||
O3 | 296 | 122 | 39% | 3.5% |
TB33 | 105 | 42 | 40% | 6.2% |
Herpetotheriidae | ||||
O3 | 747 | 344 | 46% | 2.5% |
TB33 | 257 | 51 | 20% | 2.8% |
Nyctitheriidae | ||||
O3 | 163 | 36 | 22% | 3.7% |
TB33 | 584 | 102 | 18% | 1.8% |
Theridomyidae + Herpetotheriidae | ||||
O3 | 1532 | 849 | 55% | 1.9% |
TB33 | 865 | 378 | 44% | 2.3% |
Gliridae + Nyctitheriidae | ||||
O3 | 459 | 158 | 34% | 2.7% |
TB33 | 689 | 144 | 21% | 1.8% |
Total | ||||
O3 | 1991 | 1007 | 51% | 1.6% |
TB33 | 1554 | 522 | 34% | 1.5% |
TABLE 8. Percentages of fresh breakage of elongate bones and etched (rounded) bones and teeth for rodents (theridomyids, pseudosciurids, and glirids) and insectivorous mammals (herpetotheriid marsupials and nyctitheres) in bed TB33, How Ledge Limestone. N.B., pseudosciurids are only represented by teeth. Percentages in the last column have been calculated as in Table 5.
Total | Affected | % of total |
% square root |
|
Breakage of elongate fresh bone | ||||
Rodents | 247 | 152 | 62% | 5.0% |
Insectivores | 97 | 72 | 74% | 8.7% |
Etching of bones | ||||
Rodents | 376 | 135 | 36% | 3.1% |
Insectivores | 164 | 62 | 38% | 4.8% |
Etching of teeth | ||||
Rodents | 713 | 369 | 52% | 2.7% |
Insectivores | 841 | 153 | 18% | 1.5% |
TABLE 9. Counts and percentages from each family represented in bed TB33, How Ledge Limestone by numerous postcranial elements, showing four weathering stages defined by Andrews (1990). Numbers in parentheses have been calculated as in Table 5.
Theridomyidae | Gliridae | Herpetotheriidae | Nyctitheriidae | |
Total N | 879 | 105 | 343 | 662 |
Weathering stage 0 | 722 | 95 | 310 | 630 |
Weathering stage 1 | 140 | 10 | 30 | 29 |
Weathering stage 2 | 17 | 0 | 2 | 3 |
Weathering stage 3 | 0 | 0 | 1 | 0 |
Total weathered | 157 | 10 | 33 | 32 |
% stage 0 | 82.1% (±3.1%) | 90.5% (±9.2%) | 90.4% (±5.1%) | 95.2% (±3.8) |
% stage 1 | 15.9% (±1.4%) | 9.5% (±3.0%) | 8.7% (±1.6%) | 4.4% (±0.8) |
% stage 2 | 2.0% (±0.5%) | 0% | 0.6% (±0.4%) | 0.4% (±0.3%) |
% stage 3 | 0% | 0% | 0.3% (±0.3%) | 0% |
% weathered | 17.0% (±1.4%) | 9.5% (±3.0%) | 9.6% (±1.7%) | 4.8% (±0.8%) |
TABLE 10. Counts of the bones of Paramiacis sp., belonging to a single individual from bed TB33, How Ledge Limestone, SW Headon Hill. The figure in parentheses represents the number of first phalanges when the broken proximal and distal ends are united with the rest of the bone in each case.
Skeletal element | Right | Left | Complete | Distal | Proximal | Shaft | % Fragmentation |
Petrosal | 1 | 1 | 2 | 0 | |||
Caudal vertebra | 2 | 1 | 50 | ||||
Metacarpal II | 1 | 0 | 0 | 0 | 1 | 0 | 100 |
Astragalus | 1 | 0 | 0 | 100 | |||
Calcaneum | 0 | 1 | 0 | 100 | |||
Entocuneiform | 0 | 1 | 1 | 0 | |||
Ectocuneiform | 0 | 1 | 1 | 0 | |||
Metatarsal II | 0 | 1 | 0 | 0 | 1 | 0 | 100 |
Metatarsal IV | 0 | 1 | 0 | 0 | 1 | 0 | 100 |
Unidentified M/P | 3 | 0 | 3 | 0 | 0 | 100 | |
Phalanx 1 | 16 (12) | 1 | 9 | 4 | 2 | 94 | |
Phalanx 2 | 10 | 1 | 7 | 2 | 0 | 90 | |
Ungual | 7 | 7 | 0 | ||||
Total bones | 47 | 14 | 19 | 9 | 2 | 64 |