PREDATORS CATEGORIZATION BASED ON TAPHONOMIC

ANALYSIS OF MICROMAMMAL BONES: A COMPARISON TO

PROPOSED MODELS
Gustavo N. Gmez*
* INCUAPA, Facultad de Ciencias Sociales (UNCPBA), Avda. del Valle 5737, B7400JWI, Olavarra, Buenos Aires, Argentina. E-mail:
ggomez@soc.unicen.edu.ar

ABSTRACT
The aim of this paper is to present the results of an experiment which was conducted in order to study the ingestion and
digestion effects that some predators from South America, both wild and in captivity, produce on their prey bones. This
taphonomic analysis is based on the theoretical and methodological framework proposed by Andrews (1990) and aims to
classify predators from the Pampean region according to the modifications that they produce on the bones of their prey. The
archaeological record from the Interserrana Bonaerense (Pampean region, Argentina) area is rich in small mammal bones.
However, very few taphonomic studies on micromammals have been carried out in this area and there is little knowledge
about their role in the archaeological record.

RESUMEN
En este trabajo se presentan los resultados de un experimento realizado a fin de estudiar los efectos de la ingestin y
digestin que determinados predadores de Amrica del Sur en cautiverio y en estado salvaje producen sobre sus presas. El
anlisis tafonmico se realiz bajo los conceptos tericos y metodolgicos diseados por Andrews (1990) con el propsito
de realizar una categorizacin de predadores de la regin Pampeana a travs de las modificaciones que ellos producen sobre
los huesos de sus presas. El registro arqueolgico del rea Interserrana Bonaerense (regin Pampeana, Argentina) posee
una gran riqueza de restos de pequeos mamferos, de los cuales escasos estudios tafonmicos han sido realizados y poco
se conoce acerca de su ingreso en el registro arqueolgico.

archaeologists, since many of these remains were deposited

in the sites due to diverse taphonomic agents (i.e., fluvial or
eolian action, predators, natural or catastrophic deaths). In
this context, this experiment aims to provide criteria to identify
the intervention of possible predators in archaeological
sites, based on Andrews (1990) methodological proposal.

INTRODUCTION
Archaeological research in the Interserrana Bonaerense area
(Pampean region, Argentina) has identified a large quantity
of bone remains from different micromammal species; many
of them related to diverse cultural activities that range from
the end of the Pleistocene to historical times (Politis and
Madrid 1988; Quintana 2004, 2005; Quintana and Mazzanti
2001). It is also necessary to remark, the abundance in general
terms, of these kinds of bone remains, which have been
used as environmental indicators (Crivelli Montero et al.
1987-1988; Pardias 1991).
The objective of this paper is to present and discuss the
results of an experiment carried out with different predator
species from the Pampean region and other areas of
Argentina, in order to evaluate their ingestion and digestion
effects on microfaunal bones.

Taphonomic analysis was used to evaluate possible

grouping agents for these micromammal bones in
archaeological sites. Several research projects carried out
on this topic have recognized the action of predators as
producers of associations of microvertebrate fossils
(Andrews 1990; Bochenski et al. 1997; Dauphin et al. 1988;
Denys 1985, 1986; Denys et al., 1995, 1996; Fernndez Jalvo
1992, 1995, 1996; Fernndez Jalvo et al. 1998; Laudet et al.
1996, 1997). These authors suggest that predators are
important agents of deposit and formation of microfaunal
assemblages (Denys et al. 1995).

In some cases, micromammal remains found in archaeological

sites have produced contextual interpretation problems for

The studies carried out by Andrews (1990); Fernndez Jalvo

(1992, 1996); Andrews and Fernndez Jalvo (1992); Denys

et al. (1997); Fernndez Jalvo (1996) establish several

predator categories using a) anatomical elements retrieved
from pellets and scats, b) digestion traits, and c) fracture
patterns. These predator categories are based on European,
African and some North American fauna. However, similar
studies from Interserrana Bonaerense area have not been
done yet and, therefore, it is necessary to taphonomicaly
correlate predator categories identified in previous studies
with potential South American predator species found in
archaeological and paleontological sites. To obtain
identification criteria of possible predators in these sites
would therefore help to interpret the origin of fossils
associations and possible bias produced by paleobiological
agents.

MATERIALS AND METHODS

Most of the predators involved in this experiment were
animals that were held in captivity at La Mxima County
Zoo in the city of Olavarra, Buenos Aires province. Other
samples, including Tyto alba, Asio flammeus, Circus buffoni,
and Oncifelis geoffroyi come from wild individuals whose
pellets and scats were collected from different parts of
Buenos Aires province.
Some species used for this study were fed on Mus musculus
during seven days in order to prepare them for this new
diet. In the case of other big animals, such as foxes, pumas,
jaguaroundis and wild cats, their diets were complemented
with meat with bones. In all cases one individual of each
predator species was used. After 8 days on this diet, the
collection of samples was started and lasted for the following
8 days. Predators were fed with 4 mice daily. Once recovered,
the pellets and scats were carefully desiccated during 7 days
under sunlight or incubator with an average temperature of
39 C. After being desiccated, the samples were kept in paper
bags and labelled. Number of pellet or scat, species and
recovery date was recorded in each case (Table 1).

cleaned and Mus musculus bones were recovered. Samples

were afterwards counted and taphonomicaly analysed.
The micromammal bones recovered were analysed taking into
account three different perspectives: 1) numerical data; 2)
description of the recorded features; and 3) comparative study.

1. Numerical data
This analysis comprises the accounting of relevant traits,
for example, the anatomic representation, taphonomic
features and digestion traces.
The analysis of variations on the representation between
postcranial and cranial elements was done applying two
kinds of rates: pc/c (Andrews 1990), which compares the
number of all major postcranial elements (i.e., femur, tibia,
humerus, radius and ulna) with cranial elements (mandible,
maxilla and isolated molars). The second rate is f+h/md+mx
(Andrews 1990), which compares the number of proximal
limb elements (femur and humerus) with the number of
mandibles and maxillae. These rates give a clear view of the
conservation state of cranial elements, from which it is
possible to make inferences about post-depositional effects.
Another rate that was used is that which measures the limbs
preferential distal element loss (t+r)/(f+h) (tibia + radius) /
(femur + humerus).

2. Description of the recorded features

The breakage of the material was quantified according to
Andrews (1990), who considered femur, humerus, ulna and
tibia. The elements that were analysed, depending on their
breakage level, were complete elements (in spite of the lack of
a fragment of the epiphyses), proximal and distal epiphyses,
shafts, and distal and proximal ends with part of the shaft.

As regards cranial breakages, the following categories were

considered: complete skull, broken skull with intact zygomatic
The analysis of the samples was carried out at the
process, and maxilla fragment lacking the zygomatic process.
Paleobiology Department of Natural Sciences National
This last category is considered to be a diagnostic element of
Museum (Madrid, Spain), where the scats and pellets were
breakage (Fernndez Jalvo 1992) due to
the fact of it being a delicate element.
Family
Species
Common name
Wild sample Zoo sample
Furthermore, a comparison between the
Tytonidae
Tyto alba
Barn owl
4 pellets
Strigidae
Bubo virginianus
Great Horned owl
9 pellets
empty alveoli as well as molars in situ was
Strigidae
Athene cunicularia
Burrowing owl
11 pellets
considered
together with maxillary tooth
Strigidae
Asio flammeus
Short-eared owl
1 pellet
loss.
Accipitridae
Circus buffoni
Long winged harrier
4 pellets
Accipitridae
Falconidae
Canidae
Felidae
Felidae
Felidae
Mustelidae
Didelphidae

Milvago chimango
Polyborus plancus
Canis (Pseudalopex) gimnocercus
Puma concolor
Oncifelis geoffroyi
Herpailurus yagouaroundi
Conepatus chinga
Didelphis albiventris

Chimango
Crested caracara
Pampas fox
Puma
Geoffroys cat
Otter cat
Skunk
Opossum

Table 1. Actual predator species analysed in this paper.

2 pellets
2 pellets
2 scats
2 scats
2 scats
3 scats
2 scats
2 scats

In the process of analyzing broken

mandibles, it is necessary to take into
account that these elements are denser
than skulls and therefore less likely to be
broken. The following categories were
considered: complete mandible,
ascending ramus broken and ascending

entire exposed surface altered. Moreover, the incisors that

are in the maxillas present a more remarkable alteration than
those that are in mandibles, due to the fact that maxillas are
more porous than mandibles and can therefore affect the
parts protected by the bone.

ramus missing and inferior border broken (Andrews 1990).

As well as maxillas, the presence or absence of incisors and
molars was considered.
An important rate is the quotient between isolated teeth
from the sample and the number of empty alveoli from the
mandibles and maxillas, which is multiplied by 100. The teeth
that are free are considered as belonging to mandibles and
maxillas that have been destroyed.

Incisor roots are not considered to be a digestion diagnostic

element because their appearance is not as homogeneous as
the rest of the surface. Their aspect is due to the fact that
rodents have incisors with continuous growth, and they have
a greater mineral concentration in the roots, being therefore
more liable to be dissolved than other skeletal parts.

2.1 Postcranial digestion

Digestion affects both extremes and salient and inner parts

of the elements, as well as enamel and mineralized parts (like
roots), dentine and bone (Denys et al. 1997). Four digestion
degrees have been established by Andrews (1990):

Predators produce physical breakage as well as chemical

modifications on postcranial skeletons. Chemical
modifications are exclusively produced by predators and
cannot be caused by other post-depositional events. The
digestion of cranial and postcranial elements is determined
by the pH of the stomach. Depending on each type of
predator, the pH level varies, and so do the acidity and its
influence on the bones.

Light: affects the entire enamel surface that shows a lightly

pecked aspect. In some cases, digestion is concentrated on the
teeths extra mandible part, where the enamel is partially or
completely missing, (it means that the tooth was kept in the
mandible during digestion).

Digestion evidence in postcranial elements is sometimes

difficult to distinguish from the effects of other diagenetic
agents (including soil corrosion, etc.), mainly when the
digestion is not strong. Andrews (1990) and Fernndez Jalvo
(1992) take into consideration the alterations that appear on
the proximal epiphyses of femur and distal epiphyses of
humerus, mainly in the articulation joints. The digestion
effects performed in the inner part of the elements produce
a rounding of the extremes or of the breakage edge.

Moderate: Enamel is more affected, as well as dentine that

shows an undulated surface. Enamel is on the tooth, except in
the occlusion part.
Strong: Digestion evidence appears on the incisors as well as
on dentine, where an undulating surface is present on dentine.
The enamel appears like islands.
Extreme: The alteration is present all over the tooth. In some
teeth there is no enamel and there is a bit of dentine that has a
much undulated texture. In these cases, everything is dentine
with some enamel islands. The dentine can be so affected that
it may collapse and it is therefore extremely difficult to see the
incisors shape.

The presence of alterations in postcranial elements is

considered by Fernndez Jalvo (1992) as a criterion to
distinguish concentrations produced by diurnal birds and
small mammal carnivores. This author incorporates the
postcranial elements within the four categories:

2.3 Molars digestion

Light: the digestion affects the extremes of articulation joints

of major limb bone elements, which present a corroded
appearance.

The percentage of teeth alteration due to digestion degrees

is usually higher in rodent incisors than in molars. This is
due to the fact that digestion usually polishes, smoothes
and erodes the salient surfaces and the extremes. In the
case of incisors, the salient surfaces are the occlusion
extremes; while in molars the alteration differs depending
on the taxa and the molar shape.

Moderate: The surface of the shafts presents a polished

appearance and rounded breakage edges.
Strong: The surface of the shafts is undulating and the
fragments edges of the breakages are more rounded and present
a fine flange.
Extreme: the shaft surfaces are more undulating and the
breakage edges are more rounded than the others, although the
flange is heavier than those described in the previous category.

Digestion alteration in molars is more variable than in incisors

due to morphological differences among different taxa.
Andrews (1990) and Andrews and Fernndez Jalvo (1992)
have differentiated the following intensity degrees in
digestion corrosion:

2.2 Incisor digestion

Light: the enamel presents a homogeneous light pecked surface

and the apex relief is lightly attenuated.

The digestion effects on incisors are different depending

on its occurrence in the mandibles and maxillas or elsewhere.
Separated incisors have a more generalized alteration on
their surface than those that are in situ, which have the

Moderate: the enamel surface appears more pecked than in

the last case, the enamel vanishes from the contact surface
edges and from the occlusion apex.

Strong: there are some areas where the enamel has vanished
but the dentine has not been affected. The surfaces that
previously appeared pecked are at present more evident.

Tyto alba
Pellets: 4
Maxilla
Mandible
Scapulae
Humerus
Radius
Ulna
Pelvis
Femur
Tibia
Fibula
Vertebrae
Incisor
Molar
Calcaneus
Talus
Rib
Metapodial
Phalange
Total
MNI
Pc/c
f+h/md+mx
t+r/f+h
Isolated
molar
Isolated
incisor
Relative
Abundance

Extreme: The alteration is stronger. The enamel has

disappeared on most of the tooth.

Considering the different types of modifications on the

diverse skeletal parts, Andrews (1990) has established a
predators classification according to the modifications that
they produce:
Little modification: Tyto alba; Nyctea scandiaca; Bubo lacteus;
Strix nebulosa; Asio flammeus.
Intermediate modification: Bubo bubo, Bubo africanus, Strix
aluco.
Moderate modification: Athene noctua; Falco tinnunculus
(also Falco peregrinus).
Great modification: Circus cyaneus; Nyctea scandiaca;
Ichneumia albicauda; Canis latrans; Genetta genetta; Otocyon
megalopis (also Milvus milvus, Buteo buteo, and Aegypius
occipitalis).
Extreme modification: Martes martes; Canis latrans; Vulpes
vulpes; Alopex lagopus.

3. Comparative study
Images from SEM, belonging to Dr. Fernndez Jalvo and
from Dr. Andrewss collection were used in order to identify
different taphonomic traits and their origin. These images
belong to several collections that have different
proveniences, and are formed by present-day and fossil
samples. Is it important to remark that none of these images
refer to materials collected in South America.

MNE

44
45
40
39
37
41
57
50
46
37
618
23
86
27
20
240
129
180
1,759
29
195
100
93

75.9
77.6
69.0
67.2
63.8
70.7
98.3
86.2
79.3
63.8
59.2
19.8
24.7
46.6
34.5
34.5
22.2
11.1

Asio flammeus
Pellets: 1
Maxilla
Mandible
Scapulae
Humerus
Radius
Ulna
Pelvis
Femur
Tibia
Fbula
Vertebrae
Incisor
Molar
Calcaneus
Talus
Rib
Metapodial
Phalange
Total
MNI
pc/c
F+h/md+mx
T+r/f+h
Isolated molar

MNE

4
6
5
4
23
4
6
5
7
5
79
2
11
4
3
32
15
59
274
12
328
90
333
79

16,7
25,0
20,8
16,7
95,8
16,7
25,0
20,8
29,2
20,8
18,3
4,2
7,6
16,7
12,5
11,1
6,3
8,8

42
67

Isolated incisor
58

22,6

Relative abundance
59.9

Table 2. Cranial and postcranial elements from Tyto alba and Asio
flammeus pellets.

great losses. The graphic type obtained is a profile

characterized by Andrews (1990) as owl model (Figure 1).
The Tyto alba samples analysed present a pc/c index that
agrees with the expectations stated by Andrews (1990), as
well as the number of isolated teeth. The f+h/md+mx index is
similar in both the postcranial and cranial material (Table 2).

The analysis focused on the recovered skeletal parts, the

frequency and the absence of cranial parts, breakage,
digestion evidence observed on femoral proximal ends, distal
humerus ends, and digestion degrees on
120
incisors and molars.
100

Tyto alba (Barn owl, Lechuza del campanario)

%

60

40

20

Figure 1. Percentages of skeletal parts recovered from Tyto albas pellets.

"

phalanges

metapodials

molars

incisors

vertebrae

fibula

tibia

femur

pelvis

ulna

radius

humerus

scapula

mandible

maxillas

Some elements recovered from their pellets

present rates that show equality that suggests
a minimal loss. MNI index was obtained through
the pelvises. Table 2 shows the skeletal parts
percentages recovered in Tyto albas pellets. It
can be observed that there is no great variation
between cranial and postcranial elements
percentages that therefore do not produce a
serrated curve. With the exception of incisors,
molars and phalanges, the proportions of which
are minor when compared with the other
elements, the other skeletal parts do not present

80

The cranial material seems to be complete. Maxillas maintain

the zygomatic process. Nevertheless, there are an irregular
percentage of lost molars reaching up to 85% in maxillas
and a loss of incisors that is also high (55%). As regards
mandibles, they are complete, in general, with a low
percentage of broken ascending ramus and inferior border.
There is a high loss of molars, but it is low if compared to
maxillas. The index of isolated molars is too high for this
species (42%), as the index for isolated incisors that is 58%.
This elevated separation of teeth from their alveoli seems to
be a sample problem. In any case, there is no evidence of
destruction of maxillas/mandibles due to the low breakage
rate of these anatomical elements and the low rates of isolated
teeth that are always lower than 100 (Table 3).

is higher than that present in previous samples. The distal

elements, with respect to the proximal ones, are close to the
unit and there is a good relative anatomical representation
of these elements (Table 6). The rates of isolated teeth are
contradictory, while incisors suggest a light destruction of
maxillas and mandibles; molars are separated from their
alveoli in a high percentage. A curved line that is more
serrated than in other cases, which indicates different
percentage variations in the recovered elements, represents
the relative abundance of elements.
In spite of having recorded many complete postcranial
elements (between 70% and 57%) the breakage level is higher
than in those recorded in previous cases (Table 5). None of
the maxillas, despite the fact that they have the zygomatic
process, maintain molars or incisors in situ. With respect to
mandibles, some of them are complete and with relatively
few signals of breakage. However, a total lost of incisors
and 94% of the molars have been recorded. There are a 25%
of broken incisors, all of them are isolated, and an 8% of
broken molars that are also isolated (Table 3).

A small amount of material has digestion traces. In all cases

it is inferior to 50% of the total. The largest percentage is
recorded in the proximal ends of femurs (35% have signals
of digestion); while isolated molars have the smallest
percentage of digestion traces. As regards teeth, the
tendency is to light digestion. There are signals of moderate
digestion only in isolated incisors and molars (Table 4).

The postcranial material has a light degree of digestion.

There are no recorded molars digested in situ. The isolated
incisors show light and moderate degrees of digestion, while
isolated molars reach extreme degrees (13% moderate, 5%
great, and 11% extreme) (Table 4).

Asio flammeus (Short-eared owl, Lechuzn campestre)

The elements recovered from Asio flammeus pellets show
little abundance of material, and a slight majority of
postcranial elements and mandibles. The MNE of Asio
flammeuss pellets is 274, with a relative abundance of 22.6
and a MNI of 12 (calculated as from the radius). The pc/c
rate shows a great abundance of postcranial elements (Table
2) however the relation gets closer to the unit when the radii
are not considered. As regards breakage in tibias, there are
broken distal epiphyses and isolated diaphyses (Table 5).

Athene cunicularia (Burrowing owl, Lechuza de las

vizcacheras)
The Athene cunicularias pellets studied show a great
abundance of skeletal parts. Relative abundance is low
(35.45%). Only 1047 elements out of 5984 that correspond
to the 32 individuals that were swallowed (17.6%) have been
recovered.

Cranial material is scarce. In maxillas, only a 50% of incisors

are lost, while in molars only a 17% and all maxillas keep the
zygomatic process suggesting a low breakage rate. The
mandibles are complete, with an absence of 17% of incisors
and a 67% of molars missing from their alveoli (Table 3).
The digested material is scarce; most
of it shows a light degree of digestion
(Table 4). The major percentage of
digested elements is in incisors in situ
(57%). Isolated molars have the minor
percentage, 18% of them present a
light degree of digestion.

Bubo virginianus (Great Horned owl,

acurut)
The pc/c rate shows an abundance of
postcranial material. This suggests a
light level of cranial destruction, which

Mwz
P
Mml
Mil
mc
Arb
Ibb
mml
mil
Imb
Iib

The pc/c rate is higher than 100 (equality), which shows a

great abundance of postcranial over cranial parts. The same
happens with the f+h/md+mx rate that is 115 (see Table 6). A

Ta Ta% Af Af% Bv Bv% Ac

44
100
4 100 13 100 19
9
20
2
50
112
85
2
17 39 100 57
24
55
2
50 13 100 19
42
93
6 100 3
25
11
2
4
4
33
13
1
2
5
42
3
94
70
12 67 34
94
85
16
36
1
17 12 100 24
-

%
100
100
100
38
45
10
98
83
-

Cb
20
52
22
16
19
46
20
-

%
91
79
100
80
95
77
100
-

Hj
4
12
4
1
1
3
1
-

%
100
100
100
100
100
100
100
-

Cch
4
12
4
1
11
4
3
25

%
100
100
100
25
92
100
11
89

Da
6
18
4
1
3
2
14
6
19

%
100
100
67
17
50
33
78
100
70

Table 3. Cranial elements obtained from the pellets and scats. Ta: Tyto alba; Af: Asio flammeus;
Bv: Bubo virginianus; Ac: Athene cunicularia; Cb: Circus buffoni; Hj: Herpailurus jaguaroundi;
Cch: Conepatus chinga; Da: Didelphis albiventris; Mwz: Maxillas with zygomatic; P: Palate;
Mml: Maxilla molars lost; Mil: Maxilla incisors lost; mc: mandible complete; Arb: Ascending
ramus broken; Ibb: Inferior border broken; mml: mandible molars lost; mil: mandible incisors
lost; Imb: Isolated molars broken; Iib: Isolated incisors broken.

Tyto alba
Femur proximal
Humero distal
Isolated incisor
Molar in situ
Isolated molar
Asio flammeus
Femur proximal
Humerus distal
Incisor in situ
Isolated molar
Bubo virginianus
Femur proximal
Humerus distal
Isolated incisor
Isolated molar
Athene cunicularia
Femur proximal
Humerus distal
Incisor in situ
Isolated incisor
Isolated molar
Circus buffoni
Femur proximal
Humerus distal
Isolated incisor
Molar in situ
Isolated molar
Milvago chimango
Isolated incisor
Isolated molar
Oncifelis geoffroyi
Femur proximal
Isolated incisor
Isolated molar
Herpailurus jaguaroundi
Humerus distal
Isolated incisor
Isolated molar
Conepatus chinga
Femur proximal
Humerus distal
Isolated incisor
Didelphis albiventris
Femur proximal
Humerus distal
Isolated incisor
Isolated molar

N
49
38
23
61
86

digested
17
6
4
5
5

%
35
16
17
8
6

Light
17
6
2
5
4

%
35
16
9
8
5

Moderate
1
1

5
4
7
11

1
1
4
2

20
25
57
18

1
1
4
2

20
25
57
18

% Great % Extreme %
4
1
-

16
11
32
38

5
4
8
11

31
36
25
29

5
4
4
-

31
36
13
-

4
5

13
13

11

24
25
5
54
66

10
11
4
5
42

42
44
80
9
64

10
11
4
1
33

42
44
80
2
50

4
9

7
14

4
2
42
30
65

2
2
33
21
46

50
100
79
70
71

2
2
5
-

50
100
12
-

12
10
24

29
33
37

12
8
12

29
27
18

4
3
10

10
10
15

1
2

1
1

100
50

50

1
6
12

1
6
7

100
100
58

1
-

100
-

3
3

50
25

2
4

33
33

1
-

17
-

1
6
11

1
6
11

100
100
100

1
-

100
-

3
3

50
27

2
4

33
36

1
-

17
-

5
6
28

2
4
13

40
67
46

2
4
5

40
67
18

21

7
7
27
34

2
4
20
20

29
57
74
59

2
4
7
-

29
57
26
-

7
8

26
24

5
10

19
29

1
2

4
6

Table 4. Digested cranial and postcranial elements obtained from pellets and scats.

Tyto alba
Complete
Proximal
Distal
Proximal + Shaft
Distal + Shaft
Shaft
Asio flammeus
Complete
Proximal
Distal
Proximal + Shaft
Distal + Shaft
Shaft
Bubo virginianus
Complete
Proximal
Distal
Proximal + Shaft
Distal + Shaft
Shaft
Athene cunicularia
Complete
Proximal
Distal
Proximal + Shaft
Distal + Shaft
Shaft
Circus buffoni
Complete
Proximal
Distal
Proximal + Shaft
Distal + Shaft
Shaft
Conepatus chinga
Complete
Proximal
Distal
Proximal + Shaft
Distal + Shaft
Shaft
Didelphis albiventris
Complete
Proximal
Distal
Proximal + Shaft
Distal + Shaft
Shaft

Femuri
49
1
-

Humeri
36
1
1
2
-

Tibia
45
1
-

Ulna
41
-

% Femuri
98
2
-

% Humeri
92.3
2.6
2.6
5.1
-

% Tibia
97.8
2.2
-

% Ulna
100
-

5
-

4
-

4
2
1

4
-

100
-

100
-

57.1
28.5
14.2

100
-

12
3
1
1

8
3
1
2
-

13
1
5
1
-

8
2
1
1

70.6
17.6
5.9
5.9

57.1
21.4
7.1
14.3
-

65
5
25
5
-

66.6
16.6
8.3
8.3

23
1
1
-

19
1
1
5
3
1

26
3
2
4

12
7
2
2

92
4
4
-

63.3
3.3
3.3
16.7
10
3.3

74.3
8.6
5.7
11.4

52.2
30.4
8.7
8.7

2
2

2
1

1
2

1
1
2

50
50

66.7
33.3

33.3
66.7

33.3
33.3
66.7

1
4
1
-

8
6
2
1
-

5
8
2
3
6

3
4
-

16.7
66.7
16.7
-

47.1
35.3
11.8
5.9
-

20.8
33.3
8.3
12.5
25

42.9
57.1
-

3
4
1
-

4
2
3
2
-

2
2
1
1
4

4
2
1
1

37.5
50
12.5
-

36.4
18.2
27.3
18.2
-

20
20
10
10
40

50
25
12.5
12.5

Table 5. Postcranial elements obtained from pellets and scats of nocturnal and diurnal birds and small mammals.

MNE

13
12
12
14
13
12
27
17
20
12
17
32
38
11
7
46
55
79
437
14
193
124
106
52
128

46.4
42.9
42.9
50
46.4
42.9
96.4
60.7
71.4
42.9
3.4
57.1
22.6
39.3
25
13.7
19.6
10.1

40.9

Athene
cunicularia
Pellets: 11
Maxilla
Mandible
Scapulae
Humerus
Radius
Ulna
Pelvis
Femur
Tibia
Fibula
Vertebrae
Incisor
Molar
Calcaneus
Talus
Ribs
Metapodial
Phalange
Total
MNI
pc/c
f+h/md+mx
t+r/f+h
Isolated molar
Isolated incisor
Relative
Abundance

MNE

19
29
13
30
21
23
62
25
35
22
299
54
66
17
17
116
82
117
1,047
31
188
115
102
46
126

30.6
46.8
21
48.4
33.9
37.1
100
40.3
56.5
35.5
26.8
43.5
17.7
27.4
27.4
15.6
13.8
6.7

low, with a high percentage of large bones and complete

mandibles and maxillas with zygomatic process. The traces
of digestion in incisors can enter Categories 1 or 2, like the
entire European birds prey studied by Andrews (1990).
There is no record of complete maxillas, although the remains
of them still have the zygomatic process. Complete mandibles
have been detected, although the mandibles with broken
ascending ramus are more numerous and there are a low
proportion of mandibles with a broken inferior border (Table
3). The percentage of molars and incisors missing from the
mandible alveoli is high, 83% in incisors and 98% in molars.
The complete postcranial elements are well represented
(Table 5). The number of isolated ends (proximal and distal)
is scarce. The same happens with isolated shafts. The
proximal ends that are associated to shafts are more
numerous. There are a small number of distal ends, as well
as those that are associated to the shafts. There are a high
proportion of elements with signs of digestion (Table 4).
Sixty four percent of the total amount of molars and 15% of
the incisors show signs of digestion. As regards postcranial
elements, there is a 42% in femora, and a 44% in humeri, that
is to say that half of the elements are affected by digestion.
The digestion degree observed in incisors in situ is light,
while in isolated incisors; there are four that have a moderate
degree and only one that has a light degree of digestion. As
regards molars, 14% of the isolated ones present corrosion
in a moderate degree, and 50% present a light degree.

35.45

Table 6. Cranial and postcranial elements obtained from Bubo

virginianus and Athene cunicularias pellets.

low abundance of cranial material is observed in incisors

and isolated molars. A 46% (<100) of molars are missing,
perhaps due to the sample processing. On the other hand,
the percentage of incisors is 126% (>100), showing that
there were more maxillas and mandibles in the original sample,
from where the isolated incisors accounted for in the sample
come from and that they exceed the number of empty alveoli
in a 26%.

Table 7 shows the skeletal elements of Circus buffonis

pellets, which show the scarcity of postcranial elements.
The isolated incisors and molars index give contradictory
values with respect to the rates of molars and incisors. There
is great destruction; the majority of the fragments are isolated
shafts (Table 5).

$
#
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!




m
ax
ill
ar
s
m
an
di
bl
es
sc
ap
ul
as
hu
m
er
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ra
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ou
s

Figure 2 shows the percentages of relative

abundance of Mus musculus skeletal
elements taken from the pellets. It can be
observed that there is a serrated disposition
in the elements percentage curve. Between
40 and 50% of them are mandibles, humeri,
tibias and incisors, followed by maxillas, radii,
ulnae and femora. The others elements have
lower relative abundance values (inferior to
20%). The curve of Figure 2 shows little
rough changes between recovered elements
percentages, which differ from other graphic
such as those of Tyto alba. On general
grounds, the profile of relative abundance
of Athene cunicularia can be included in
Category 1. The breakage and destruction
level of their prey remains is, in general, very

Circus buffoni (Long winged harrier, Gaviln planeador)

ul
na
s
fe
m
or
i
tib
ia
s
ve
rte
br
ae
in
ci
so
rs
m
ol
ar
m
s
et
ap
od
ia
ls
ph
al
an
ge
s

Bubo
virginianus
Pellets: 9
Maxilla
Mandible
Scapulae
Humerus
Radius
Ulna
Pelvis
Femur
Tibia
Fibula
Vertebrae
Incisor
Molar
Calcaneus
Talus
Rib
Metapodial
Phalange
Total
MNI
pc/c
F+h/md+mx
T+r/f+h
Isolated molar
Isolated incisor
Relative
Abundance

Figure 2. Percentages of skeletal elements of Mus musculus in Athene cunicularia

pellets.

&

Pellets: 4
Maxilla
Mandible
Scapulae
Humerus
Radius
Ulna
Pelvis
Femur
Tibia
Fibula
Vertebrae
Incisor
Molar
Calcaneus
Talus
Rib
Metapodial
Phalange
Total
MNI
Pc/c
F+h/md+mx
T+r/f+h
Isolated molar
Isolated incisor
Relative Abundance

MNE
22
20
0
3
1
3
1
4
3
0
34
42
65
1
1
2
32
55
289
11
21
17
57
68
100
18.8

remains and consumes the meat. This behaviour is similar

to that of the Milvago chimango and their range overlaps,
especially the way in which they consume their prey is similar
and the subsequent scarcity of bones in the pellets. Only
two isolated molars, one maxilla and one vertebra were
recovered from two pellets of Poliborus planctus (see Tables
3, 4, 5 and 8)

%
100
90
0
13.6
4.5
13.6
4.5
18.2
13.6
0
8.6
95.5
49.2
4.5
4.5
0.8
14.5
8.9

Canis (Pseudalopex) gimnocercus (Pampas fox, Zorro de

las Pampas)
Unfortunately, the recovered fox scats material does not
allow calculating the pc/c and f+h/md+mx indexes, because
there is no record of femora and humeri and the cranial parts
are very scarce. The percentage of isolated molars reaches
up to 33% and the relative abundance is 7.1%. The only
isolated incisor recorded has a moderate digestion degree.
The calculated MNI is 1. (Table 8)

Puma concolor (Cougar, Puma)

The recovered material from Puma concolor is scarce (MNE:
6; MNI: 1). There are no cranial parts and the postcranial
elements are highly broken and digested. The value obtained
from the distal limbs loss is low, but this value is the result
of a sample size problem as only one element of each
postcranial part that was recorded was recovered. (Table 8)

Oncifelis geoffroyi (Geoffroys cat, Gato monts)

Table 7. Cranial and postcranial elements obtained from Circus

buffonis pellets.

Oncifelis geoffroyi has provided a small amount of material

(MNE: 34; MNI: 2). The pc/c index suggests a great
abundance of cranial elements represented by isolated teeth
without the presence of maxillas bones. No postcranial
element is complete; there are only isolated epiphyses of
femur and tibia (Table 8).

With respect to the cranial material, all the incisors are missing
from their alveoli (Table 3). No mandible is complete, and
the fragments have a broken inferior border and there is a
high frequency of broken ascending ramus. Isolated broken
molars are scarce and there is no record of in situ broken
molars. A 50% of the isolated incisors are broken.

The femur obtained is completely digested and the isolated

incisors present different degrees of digestion: moderate
(50%), great (33%) and extreme (17%). Fifty eight percent of
the isolated molars have signals of digestion, 33% great
digestion and 25% moderate digestion (Table 4).

The small amount of postcranial elements has signs of

digestion (Table 4). The dental material, in general, including
isolated incisors, in situ and isolated molars, shows a
moderate or great degree of digestion. Still, there are some
elements that have an extreme degree of digestion. A high
percentage of incisors (79%) and molars (70%) present signs
of digestion.

Herpailurus yagouaroundi (Otter cat, Jaguaroundi)

This material is scarce; the pc/c and f+h/md+mx indexes
show that cranial elements are more numerous than
postcranial elements. There is only one humerus which is
complete and has a light digestion degree.

Poliborus plancus (Crested caracara, Carancho)

The scarcity of material from its prey in the pellets of this
diurnal bird does not allow taphonomic analysis. This
scarcity is due to the fact that the Poliborus plancus is
above all a scavenger and, when hunting, it destroys all the
bones pulling out the flesh. Probably it rejects the bone

A large amount of incisors and molars are missing from their

alveoli in the recorded cranial material. There are no complete
maxillas. The only recovered mandible is not complete, its
ascending ramus and the inferior border are broken. There

'

Milvago chimango
Pellets: 2
Mandible
Femur
Incisor
Molar
Phalange
Total
Poliborus planctus
Pellets: 2
Maxilla
Vertebrae
Molar
Total
Canis (Pseudalopex)
gimnocercus Scats: 2
Maxilla
Scapulae
Vertebrae
Incisor
Total
Puma concolor
Scats: 2
Femur
Tibia
Rib
Metapodial
Phalange
Total
Oncifelis geoffroyi
Scats: 2
Femur
Tibia
Vertebrae
Incisor
Molar
Metapodial
Total
Herpailurus
jaguaroundi Scats: 3
Maxilla
Mandible
Humerus
Vertebrae
Incisor
Molar
Talus
Rib
Metapodial

MNE

1
1
1
2
3
8

50
50
25
16.7
5.4

MNI
pc/c
f+h/md+mx
Isolated molar
Isolated incisor
Relative Abundance

Conepatus chinga (Skunk, Zorrino)

Among the scats of Conepatus chinga there is a high

proportion of tibias, from which the MNI was calculated.
There are many postcranial elements such as humerus,
pelvis and radius. Among cranial elements, isolated incisors
have the highest proportion (Table 9).

53
100
67
100
6.5

The material obtained from the scats is mainly postcranial

according to the pc/c and f+h/md+mx indexes. Perhaps this
phenomenon is due to the Conepatus chinga way of chewing
and to the fragility of the cranial elements of their prey. At
the same time, postcranial material is broken with a low
percentage of complete elements, and fragments of distal
and proximal ends (Table 5). The cranial material is scarce,
and the incisors and molar appear isolated. The recovered
elements represent the entire skeleton, but their relative
abundance is variable. The percentage of breakage is high,
mainly in isolated incisors: 89% and it is low in isolated
molars: 11% (Table 3).

1
67
3.3

1
1
2
4

50 Isolated molar
2.8 Relative Abundance
16.7

1
1
5
1
8

50 Isolated incisor
50 Relative Abundance
13.9
25

1
1
1
1
2
6

50
50
4.2
5
3.6

pc/c
t+r/f+h
Relative Abundance

1
3.2
1
7.05

2
1
6
6
12
1
34

50
25
8.3
75
50
2.5

pc/c
t+r/f+h
Relative Abundance

2
40
50
5.7

4
1
1
7
6
11
1
2
5

100
25
25
9.7
75
45.8
25
25
12.5

pc/c
f+h/md+mx
Isolated molar
Isolated incisor
Relative Abundance

1
33
7.1

There are a high percentage of postcranial elements with signs

of digestion (see Figures 3 and 4). In isolated incisors the
degree of digestion is moderate (21%), although there are
some incisors with a light degree (18%) and with a great degree
of digestion (7%). Regarding isolated molars the highest
percentage is among those with great degree of digestion
(26%), moderate (22%), and extreme (15%) (Table 4).
Conepatus
chinga Scats: 2
Maxilla
Mandible
Scapulae
Humerus
Radius
Ulna
Pelvis
Femur
Tibia
Fibula
Vertebrae
Incisor
Molar
Calcaneus
Talus
Rib
Metapodial
Phalange
Total
MNI
pc/c
F+h/md+mx
T+r/f+h
Isolated molar
Isolated incisor
Relative
Abundance

2
10
20
73
120
13.4

Table 8. Cranial and postcranial elements obtained from the pellets

and scats of Milvago chimango, Poliborus planctus, Canis
(Pseudalopex) gimnocercus, Puma concolor, Oncifelis geoffroyi,
and Herpailurus jaguaroundi.

are a 100% of incisors and molars missing from their alveoli.

The molars, both in situ and isolated, do not present signals
of breakage.
The whole set of material presents traces of digestion. Fifty
percent of the incisors present a moderate digestion degree,
33% of them a great digestion and the remaining 17% an
extreme digestion degree. All the molars have evidence of
digestion but none of them presents an extreme degree of
digestion (Tables 3, 4 and 8).

MNE

4
4
6
17
12
7
17
6
24
4
114
28
27
10
10
73
68
188
619
12
302
288
157
117
350

16.7
16.7
25
70.8
50
29.2
70.8
25
100
16.7
26.4
58.3
18.8
41.7
41.7
25.3
28.3
28

38.3

Didelphis
albiventris Scats: 2
Maxilla
Mandible
Scapulae
Humerus
Radius
Ulna
Pelvis
Femur
Tibia
Fibula
Vertebrae
Incisor
Molar
Calcaneus
Talus
Rib
Metapodial
Phalange
Total
MNI
Pc/c
F+h/md+mx
T+r/f+h
Isolated molar
Isolated incisor
Relative
Abundance

MNE

6
6
10
11
5
8
18
8
10
4
123
27
34
6
5
38
58
74
451
9
146
158
79
106
270

33.3
33.3
55.6
61.1
27.8
44.4
100
44.4
55.6
22.2
38
75
31.5
33.3
27.8
17.6
32.2
14.7

40.1

Table 9. Cranial and postcranial elements obtained from Conepatus

chinga and Didelphis albiventris scats.



The material recovered from Didelphis

albiventris has a MNE: 451 and MNI: 9.
The relative abundance is 40. The pc/c and
f+h/md+mx indexes show a higher
representation of postcranial material than
of cranial one (Table 9). There is a high level
of breakage of postcranial material,
although there are some complete elements
(Table 5).
The breakage of long bones from the
sample is versatile and variable due, to the
strong destruction produced by the
chewing of this predator. This is the only
marsupial included in this experiment.

Figure 3. SEM image from a proximal tibia recovered from Conepatus chinga scats. This
element shows a great digestion degree associated to breakage.

With respect to the digested material,

postcranial elements have a light degree of
digestion: 29% in femora and 57% in humeri.
Isolated incisors also have light and
moderate degrees of digestion (Figure 5),
while most isolated molars have a great
degree of digestion (29%), and a moderate
one (24%). There is no material with a light
degree of digestion, but there is a 6% with
an extreme digestion degree (Table 4).

DISCUSSION
Following Andrews methodology (1990),
it is interesting to compare the different
samples from owls, diurnal prey birds and
carnivore mammals to have a wider view of
the behaviour of South American species
in the dispersion graphics and to therefore
establish which category they are in. It must
be remarked, however, that this analysis was
limited to the sample obtained from the
Pampean region. In Figure 6 there are spots
with the postcranial/cranial proportions and
their relative abundances.

Figure 4. SEM image shows a Mus musculus molar with a moderate

produced by Conepatus chingas gastric juices.

Didelphis albiventris (Opossum, Comadreja overa)

There are a low proportion of cranial elements. The MNI
was obtained from the pelvises. The proportion of
postcranial elements is very similar among all of them.

The majority of the owls have the highest

amounts of postcranial elements recovered
from their pellets. Athene cunicularia (pc/
c = 188) has nocturnal habits but is one of
the most diurnal prey birds among
strigiforms. Its prey record has mandibles
digestion degree and maxillas without molars therefore, it is
in a percentage balance, 100, which points
out a light predominance of postcranial elements. Among
mammals, Andrews (1990) considers that mustelids like
Martes martes and viverrids (Genetta genetta) scats have a
high percentage of postcranial elements. In the experimental
sample there is a mustelid that has a high proportion of



and cranial elements as Andrews (1990)

examined. A different case is represented
by the sample from Puma concolor that is
situated in the postcranial skeleton
predominance; this may be as a
consequence of the small sample obtained.

Figure 5. SEM image from a Mus musculus incisor recovered from Didelphis albiventris
scats. This element shows a moderate digestion degree.

CONCLUSIONS

350
12

300

pc/c

250
4

200

150
7
100
9

50

5
11

10

10

20

30

40

50

60

70

Relative abundance

Figure 6. Plot graphic between relative abundante and pc/c values obtained from South
american predators. 1: Tyto alba; 2: Asio flammeus; 3: Bubo virginianus; 4: Athene
cunicularia; 5: Circus buffoni; 6: Milvago chimango; 7: Didelphis albiventris; 8: Conepatus
chinga; 9: Oncifelis geoffroyi; 10: Canis (Pseudalopex) gimnocercus; 11: Herpailurus
yaguaroundi; 12: Puma concolor.

Below 100 (with a higher proportion of

cranial elements) there are diurnal birds like
Circus buffoni, which can be due to the
fact that this species does not behead their
prey but destroys the other parts of the
skeleton. As well, in this zone there are also
felidae and canids, whose chewing can
produce a high destruction of postcranial

In order to state the modification in the

predator categories from the Pampean
region, the methodological analysis
proposed by Andrews (1990) was followed.
Samples from the same species and genus
were analysed. Tyto alba and Asio flammeus
samples present the same behaviour and
are similar to the samples analysed by
Andrews (1990), in spite of having
differences in a few features. For example,
the inclusion of Tyto alba in category 3,
the loss of teeth from a mandible, could
have been the result of an aggressive
treatment during the processing and
cleaning stages. Andrews (1990) argues

350

7
300

f + h / md. + mx.

postcranial elements (Conepatus chinga)

that agrees with Andrewss statement.
However, there are no viverrids in South
America although there are carnivore
marsupials like Didelphis albiventris,
whose samples present a slight
predominance of postcranial elements in
their scats.

Figure 7 shows other indexes of

representation of cranial and postcranial
proportions on the Y-axis. It is different from
the previous graphic because the majority
of the owls are close to the equality limit
(100). The samples that came from
Conepatus chinga and Didelphis
albiventris maintain a great abundance of
postcranial elements with values over 100.
Below that value are on one side a diurnal
bird (Circus buffoni) and, on the other,
mammal predators (mainly felidae).

250
200

6
150

100

50

10

0
0

8
10

5
20

30

40

50

60

70

Relative abundance
Figure 7. Plot graphic between relative abundante and f+h/md+mx values obtained from
South american predators. 1: Tyto alba; 2: Asio flammeus; 3: Bubo virginianus; 4: Athene
cunicularia; 5: Circus buffoni; 6: Didelphis albiventris; 7: Conepatus chinga; 8: Oncifelis
geoffroyi; 9: Herpailurus yaguaroundi; 10: Canis (Pseudalopex) gimnocercus.

that there is a variability which causes differences in the

interpretation of incisors and molars missing from their alveoli
that depends on the prey. This is because the shape of their
roots supplies a stronger or softer grip of the teeth in the
alveoli. This was observed in species with molars from the
Octodontidae family.

diurnal prey birds which destroy the distal ends, as well as

postcranial elements with a high digestion degree. Another
species to point out is Bubo virginianus whose modification
features are similar to Bubo africanus and Bubo bubo, with
a high digestion degree. The relative abundance is high but
the breakage level is not so.

The samples from Poliborus planctus, Canis (Pseudalopex)

gymnocercus, and Puma concolor, despite the scarcity of
elements, have provided some data that allow establishing
their category. Some samples that show a high destruction
level and/or effects of extreme digestion must be included
in the highest categories.

Carnivore mammals like Conepatus chinga and the marsupial

Didelphis albiventris, must be submitted to the extreme
category of modification, like the carnivore species studied
by Andrews (1990). A great destruction of cranial and
postcranial elements is due to chewing and the great and
extreme digestion degrees, rounded edges and salient of
some postcranial fragments, especially in breakage areas,
characterize these species. In sum, the different Pampean
region species can be classified according to the following
graphic, using the criteria that have been used for their
categorization (Table 10 and Figure 9).

Poliborus planctus and Milvago chimango are species with

scavenger habits; therefore they produce a great destruction
of their prey. Puma concolor has very destructive digestion
effects over the material. Small fragments of bone, not
belonging to micromammals, which have important digestion
effects, were recovered from their scats (Figure 8). It should
be considered that these species belong to the maximum
modification category.

Predator categorization is important because a wide

spectrum of micromammal species appears in many
archaeological sites in the Pampean region; their bones have
scarcely been studied from a taphonomic perspective.
Nowadays, the taphonomic analysis of these species offers
a range of information concerning the different processes
that could have
happened
(including
p r e d a t o r
activities) which
would not have
been recorded in
the site using
other kinds of
evidence.

Regarding Athene cunicularia, typical modifications from

category 5 can be observed, although some of them are
characteristic of category 1. It differs from, Athene noctua,
a European species because they present features of major
destruction and modification (Andrews 1990). This can be
due to size differences between the two species, being the
European one smaller than the American, thus producing a
major breakage of skeletal elements.
There is a great coincidence between the features observed
in Circus buffoni and the European species Circus cyaneus,
whose data are presented by Andrews (1990). Both of them
fall into the category of great modification. They are, both,

Figure 8. SEM image shows bone fragment (no micromammal) with a high digestion degree
produced by Puma concolors gastric juices.

!

Figure 9. Summary of the categories of the

South American predators.

R elative
A bundance

Pc/c

T+r/f+h

Postcranial
Breakage

M axilla with
zygom atic

M axilla
Loose teeth

M andible
Breakage

M andible
Loose teeth

C ategory 1
Tyto alba

C ategory 2
Bubo
Virginianus
D idelphis
abiventris

Bubo virginianus
Asio flam m eus
Tyto alba

Athene cunicularia
Asio flam m eus
Conepatus
Chinga
Bubo virginianus
Tyto alba
Athene cunicularia
Asio flam m eus
Tyto alba

Asio flam m eus

Athene cunicularia
Bubo virginianus
Tyto alba
Circus buffoni
D idelphis
albiventris
Conepatus chinga
H erpailurus
Jaguaroundi
Canis Pseudalopex
gym nocercus
Asio flam m eus

Asio flam m eus

Tyto alba
Athene cunicularia

C ategory 3
Athene cunicularia

Circus buffoni
M ilvago chim ango
H erpailurus
jaguaroundi
O ncifelis geoffroyi

D idelphis
albiventris

Athene
cunicularia
Bubo
virginianus

Bubo virginianus
Asio flam m eus
D idelphis
albiventris
Conepatus chinga

C ategory 4
Circus buffoni
Asio flam m eus
Conepatus
chinga
D idelphis
albiventris
Conepatus
chinga
Athene
cunicularia
Puma concolor

C ategory 5
Canis Pseudalopex
gym nocercus
M ilvago chim ango
and felids

Circus buffoni
O ncifelis
geoffroyi

Puma concolor

D idelphis
albiventris

O ncifelis geoffroyi
Pum a concolor
M ilvago chim ango

Tyto alba
Asio flam m eus

Bubo
virginianus
D idelphis
albiventris
Conepatus
chinga
Asio flam m eus

D idelphis
albiventris

Athene
cunicularia
Bubo virginianus
Circus buffoni
D idelphis
albiventris
Conepatus
chinga
H erpailurus
jaguaroundi
Circus buffoni
H erpailurus
jaguaroundi
M ilvago
chimango

Tyto alba
D idelphis
albiventris
Circus buffoni

Athene
cunicularia
Bubo virginianus
Circus buffoni
D idelphis
albiventris
Conepatus
chinga
H erpailurus
jaguaroundi
M ilvago
chimango

Table 10. Summary of the categories of the South American predators.

"

Isolated teeth

C ategory 1
Tyto alba
Asio flam m eus

B reakage
T eeth

Tyto alba
Asio flam m eus

M olars w ith
digestion

Incisors w ith
digestion

C ategory 2
M ilvago
chim ango
Bubo
virginianus
C ircus buffoni
Bubo
virginianus
Asio flam m eus
Tyto alba

Tyto alba

Postcranial
w ith
digestion

Asio flam m eus

Tyto alba
Bubo
virginianus
D idelphis
albiventris

C ategory 3
Athene cunicularia
H erpailurus
jaguaroundi

Bubo virginianus

C ategory 4
C onepatus
chinga
D idelphis
albiventris
C onepatus
chinga

C ategory 5
C ircus buffoni
M ilvago chim ango

C onepatus
chinga
D idelphis
albiventris

Athene cunicularia
C ircus buffoni
C onepatus
C hinga
O ncifelis G eoffroyi
H erpailurus
jaguaroundi
C ircus buffoni
O ncifelis geoffroyi
H erpailurus
jaguaroundi
M ilvago chim ango

Asio flam m eus

Athene cunicularia
C onepatus chinga
Bubo virginianus
D idelphis
albiventris
C ircus buffoni
Athene cunicularia
C onepatus chinga

D idelphis
albiventris

Poliborus planctus
M ilvago chim ango
O ncifelis geoffroyi
H erpailurus
jaguaroundi

Table 10. Second part.

On the other hand, it is important to distinguish bones that

were deposited by predators and from those of the species
that lived in the site. These analyses are important not only
to distinguish bones deposited by predators, but also to
identify potential disturbances such as galleries and dens
that predators can produce in the archaeological record.

Andrews, P. and Y. Fernndez Jalvo

1992 Small mammal taphonomy of Gran Dolina, Atapuerca
(Burgos), Spain. Journal of Archaeological Science
19: 407-428.

Bochenski, Z., V. Korovin, A. Nekrasov and T. Tomek

1997 Fragmentation of bird in food remais of imperial eagles
(Aquila heliaca). International Journal of
Oseoarchaeology 7:165-171.

Acknowledgements

Crivelli Montero, E., M. Silveira, E. Eugenio, P. Escola, M.

Fernndez and N. Franco

This investigation was carried out under the research

program INCUAPA (Investigaciones Arqueolgicas y
Paleontolgicas del Cuaternario Pampeano), directed by Dr.
Gustavo G. Politis and Lic. Jos L. Prado (FACSO, UNICEN).
The material was analysed using a binocular lens Wild M8
de 50X, from Museo Nacional de Ciencias Naturales, Madrid,
Spain. The most significant specimens were also analysed
in a SEM from the Natural History Museum of London,
England. I would like to thank La Mxima County Zoos
director and staff, as well as Dr. Mara Teresa Alberdi, Dr.
Peter Andrews, Dr. Yolanda Fernndez Jalvo, Dr. Mara A.
Gutierrez, and to all the INCUAPA researcher staff.

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