321 Arheološki vestnik 71, 2020, 321–357; DOI: https://doi.org/10.3986/AV.71.11
Holocene macromammal remains 
from Grotta dell’Edera/Stenašca, Trieste Karst
(excavations in 1990–2001)
Ostanki velikih sesalcev holocenske starosti
iz jame Stenašca/Grotta dell’Edera na Tržaškem Krasu
(izkopavanja v letih 1990–2001) 
Francesco BOSCHIN
Izvleček
Predstavljeni so rezultati zooarheološke in tafonomske analize ostankov velikih sesalcev iz jame Stenašca/Grotta 
dell’Edera) na Tržaškem Krasu. Najstarejše človekove sledi so datirane v mezolitik (sovterjen). Ljudje so se namreč 
v jami občasno zadrževali že v predborealu, intenzivneje pa so jo začeli obiskovati v borealu in v začetku atlantika. 
Zadnja faza prisotnosti mezolitskih skupin je časovno umeščena v kastelnovjen. Označujejo jo ostanki domačih in 
lovnih živali, pri čemer večji del kosti izkazuje sledi neposredne izpostavljenosti vremenskim pojavom. To bi lahko 
pričalo o razmeroma počasnem odlaganju omenjene plasti, ki je povzročilo nastanek palimpsesta, vendar bi dokončna 
razjasnitev problematike terjala več geoarheoloških podatkov. Za skupek živalskih ostankov iz neolitskih in eneolitskih 
plasti je značilna očitna prevlada kosti in zob drobnice, pa tudi postopna rast števila najdb domačega prašiča in goveda. 
Na podlagi prisotnosti posameznih kosti ovčjih in/ali kozjih zarodkov je mogoče sklepati na naraven pogin teh živali. 
Slednje kliče k previdnosti pri oceni starostnih profilov domestikatov z najdišč, ki so bila uporabljana kot staje.
Ključne besede: Tržaški Kras; Grotta dell’Edera/Stenašca; sovterjen; mezolitik; neolitik; eneolitik; zooarheologija; 
tafonomija
Abstract
The zooarchaeology and taphonomy of large mammal remains from Grotta dell’Edera (Trieste Karst, northern Italy) 
are presented in this paper. A Mesolithic (Sauveterrian) Preboreal assemblage testifies to a first human occupation, 
followed by a second longer Sauveterrian frequentation from the Boreal until the beginning of the Atlantic. The last 
Mesolithic groups are represented by a Castelnovian assemblage, in which the bones of domesticates were detected with 
wild games remains. The presence of weathered bones in this layer indicates possible slow sedimentation, which may 
have led to the formation of a palimpsest, but more geoarchaeological data are needed to clarify this context. Neolithic 
and Eneolithic layers are characterized by an overwhelming presence of caprines and by an increase in the domestic 
pig and cattle through time. A number of caprine foetal remains points to the presence of individuals accumulated due 
to natural death and to the caution needed in evaluating caprine age profiles from sites used as stables.
Keywords: Trieste Karst; Grotta dell’Edera/Stenašca; Sauveterrian; Mesolithic; Neolithic; Eneolithic; zooarchaeology; 
taphonomy

322 Francesco BOSCHIN
INTRODUCTION
Grotta dell’Edera (Stenašca) (Figs. 1 and 2) is 
a key site for the study of the first colonization 
of the Karst plateau (northern Adriatic) and for 
better understanding its neolithization process. 
This area seems to have been uninhabited for 
several millennia (Boschian, Fusco 2007) at least 
from the onset of the Upper Palaeolithic until 
the Mesolithic; the first traces of Anatomically 
Modern Humans date to the early Holocene. In 
addition, in most of the archaeological cave sites 
widespread over the Karstic plateau, the Mesolithic-
Neolithic sequences are almost always incom-
plete (Biagi, 2003). From this perspective, Grotta 
dell’Edera is a unique site due to a stratigraphy 
that yielded Mesolithic Sauveterrian (Layers 3b–d) 
and Castelnovian (Layer 3a) remains, as well as 
Neolithic and Eneolithic ones (Layers 2–2a) (Fig. 
3). Being the only detailed Mesolithic-Neolithic 
carbon-dated sequence of the Trieste Karst (Fig. 
4), Grotta dell’Edera must be considered one of 
the most important cave-sites in the province of 
Trieste. Moreover, it was thoroughly excavated, 
with water sieving and flotation to recover small 
finds and plant remains. Excavation proceeded 
following stratigraphic criteria (arbitrary spits of 
5 or 10 cm were used within the different layers).
For this reason, the analysis of animal remains 
uncovered at the site is a unique opportunity to 
collect data useful for the reconstruction of subsist-
ence strategies of Mesolithic and Neolithic people 
that (re)colonized the plateau. Zooarchaeological 
remains were previously published in preliminary 
reports by Boschin and Riedel (2000; 2011) and, 
in a recent paper discussing caprines, mortality 
profiles (Boschin 2020) were published. The aim 
of this paper is to present a complete report on 
the zooarchaeology and taphonomy of the mac-
romammal assemblage.
Fig. 1: Position of Grotta dell’Edera and other sites cited 
in the text. Dary streams indicate the water sources closest 
to Grotta dell’Edera.
Sl. 1: Lega Stenašce in drugih v besedilu obravnavanih 
najdišč. Vrisani so tudi vodni viri v neposredni bližini jame.
1 Grotta dell’Edera/Stenašca; 2 Riparo di Biarzo/Pečina 
pri Bjarču; 3 Grotta del Mitreo/Mitrejeva jama; 4 Grotta 
Benussi; 5 Grotta Azzurra/Pečina na Leskovcu; 6 Vlaška 
jama; 7 Grotta degli Zingari/Ciganska jama; 8 Grotta dei 
Ciclami/Orehova pejca; 9 Mala Triglavca; 10 Viktorjev 
spodmol; 11, 12 Grotta degli Orsi/Medvedja jama, Grotta 
Piccola Pocala; 13, 14 Podmol pri Kastelcu, Acijev spodmol; 
15 San Rocco/Koromačnik; 16 Pupićina peć
Fig. 2: Grotta dell’Edera. Site planimetry. Excavated squares 
are marked in grey.
Sl. 2: Stenašca. Planimetrija najdišča. Posamezni izkopani 
kvadranti (označeni sivo).

323 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
The cave is located in the north-western sector 
of the Trieste Karst (northeastern Italy). It opens 
at the bottom of a small sinkhole, at an altitude of 
about 230 m a.s.l., some 3 km from the present-day 
coastline (Fig. 1); its 10 m wide entrance faces N. 
E. (Fig. 2). The cave was first excavated by Giorgio 
Marzolini (1970), and the fieldworks were resumed 
in 1990 by the Ca’Foscari University of V enice and 
the University of California at Berkeley (USA) and 
directed by Paolo Biagi and Barbara A. Voytek 
(Biagi, Starnini, Voytek 1993).
The stratigraphical sequence includes both 
Pleistocene and Holocene sediments (Boschian 
1997), but the archaeological layers cover only a 
Holocene time-span between the Preboreal and 
the Migration Period (Spataro 1999; Biagi, Spataro 
2001; Biagi, Starnini, Voytek 2008) (Fig. 3).
Available geoarchaeological data suggest that 
the deepest layers investigated during the first 
excavations are related to a quite wet and cold Late 
Pleistocene phase (layers up to Layer 6 in Boschian, 
1997, Fig. 3), which evolves towards slightly drier 
conditions (Layers 7–9). These phases were ten-
tatively related to the Dryas III (Boschian 1997). 
Stony layers corresponding to the Mesolithic (Layers 
10–12a) are characterized by intense human-related 
activities and thus are the uppermost Neolithic-
Eneolithic levels 13–15. In general, a decrease in 
the external input of sediments is visible, along 
the Holocene sequence, probably due to climate 
amelioration and afforestation (even if this trend 
is partially inverted during a possibly more arid 
climatic phase corresponding to the Eneolithic). 
This interpretation appears to be confirmed by 
available archaeobotanical data (Nisbet 2000). 
During the Neolithic and the Eneolithic, most of 
the sediments are related to the accumulation of 
domestic livestock droppings (Boschian 1997).
T o avoid misunderstanding, it has to be empha-
sized here that the naming of layers was changed 
during the most recent excavations. Layers com-
posing the about 4 m thick newly excavated stra-
tigraphy were labelled as follows, from bottom to 
top (Figs. 3 and 4):
– Layer 3d: it is related to the cave’ s oldest human 
frequentation (Mesolithic, Sauveterrian), which 
took place during the Preboreal period (9810±70 
uncal., GrN-23130; 9930±50 uncal., GrA-14108).
– Layer 3c: it yielded a living floor characterized 
by the presence of in situ faunal remains, lithics, and 
ash/charcoal lenses. It dates to the Boreal period 
(8350±120 uncal., GrN-25139; 8250±50 uncal., 
GrA-11818) and corresponds to a Sauveterrian 
occupation phase.
– Layer 3b: it yielded a Sauveterrian Mesolithic 
assemblage, and it is ascribable to the Boreal/be-
ginning of the Atlantic period. Dates span from 
8045±40 uncal. (GrA-14106) to 8110±90 uncal. 
(GrN-25138).
– Layer 3a: it is a hearth of about one metre in 
diameter which yielded a Late Mesolithic Castelno-
vian lithic assemblage together with a total amount 
of 17 potsherds; petrographic characterization of 
the latter revealed an allochthonous provenance 
Fig. 3: Grotta dell’Edera. Site’s stratigraphy; layers are 
labelled according to the 1990–2001 excavations. Layers: 
3d–b Mesolithic/Sauveterrian; 3a Mesolithic/Castelnovian; 
2a Neolithic (Vlaška group); 2 Late Neolithic/Eneolithic; 
1b Bronze Age; 1a Late Antiquity (Migration Period).
Sl. 3: Stenašca. Stratigrafija najdišča, izkopavanja 1990–2001. 
Plasti: 3d–b mezolitik/sovterjen; 3a mezolitik/kastelnovjen; 
2a neolitik (vlaška kulturna skupina); 2 pozni neolitik/
eneolitik; 1b bronasta doba; 1a pozna antika (obdobje 
preseljevanja ljudstev).

324 Francesco BOSCHIN
Fig. 4: Grotta dell’Edera. 
14
C Dates; calibration was performed with the software Oxcal 
v4.3.2 (Bronk Ramsey et al. 2013), using the IntCal13 atmospheric curve (Reimer et al. 
2013). Dates are from Spataro 1999; Biagi 2003; Biagi, Starnini, Voytek 2008.
Sl. 4: Stenašca. Datumi C
14
; kalibriranje je bilo izvedeno z uporabo programskega paketa 
OxCal 4.3 (Bronk Ramsey 2009) ob upoštevanju krivulje IntCal13 (Reimer et al. 2013). 
Podatki so povzeti po: Spataro 1999; Biagi 2003; Biagi, Starnini, Voytek 2008.

325 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
and a mineral composition, which differs from that 
of ceramics from the more recent Layer 2a (Spa-
taro 2001). A piece of charcoal dates to 6700±130 
uncal. (GX-19569), whilst a sheep bone dates to 
6620±60 uncal. (GrA-19912).
– Layer 3: it is a reworked level of about 20 
cm in thickness, where potsherds and few lithic 
flakes of local raw material were uncovered. Up 
to this layer, sediments are mainly characterized 
by a more reddish colour.
– Layer 2a: it is a thick palimpsest, composed of 
six superimposed lenses of ash and charcoal. Most 
of the sediments are composed of burned domestic 
ungulates droppings. Fragments of potsherds are 
ascribed to the Vlaška group, considered as a local 
variant of Danilo Neolithic culture. Dates range 
from 6305±285 uncal. (GX-19022) to 6615±390 
uncal. (GX-19568) (Biagi, Starnini, Voytek 2008).
– Layer 2: it is another palimpsest, whose up-
per part (at a depth of 170–180 cm) dates to an 
early phase of the Eneolithic (4680±325 uncal., 
GX-19019); another date, coming from a lower 
part of the layer (depth of 200–210 cm) is 5555±35 
uncal. (GrN-18968).
– Layer 1b: probably reworked, it yielded few 
early Bronze Age remains.
– Layer 1a: it yielded scarce archaeological re-
mains and a hearth dated to the Migration Period 
(1415±35 uncal., GrN-18968).
– Layer 1: it represents the top of the sequence, 
with scarce archaeological significance.
METHODS
Specimens discussed in this paper were identi-
fied at anatomic and taxonomic levels, using os-
teological comparative collections. The taxonomic 
nomenclature of domestic mammals was used 
following Gentry, Clutton-Brock, Groves (2004). 
In a previous report on Layers 2, 2a, 3, and 3a 
(Boschin, Riedel 2000), long bones were counted, 
and their different parts were considered as separate 
entities: for instance, if a femur was complete, it 
was counted three times (proximal, medial and 
distal), which means that the number of remains 
expressed in the tables in Boschin, Riedel (2000) 
does not correspond to the effective number of 
specimens found at the site. For this reason, the 
number of identified specimens (NISP) is used 
here to avoid mistakes in case other authors will 
use zooarchaeological data from Grotta dell’Edera. 
When possible, taxonomically unidentified speci-
mens were grouped according to anatomy (e.g., 
ribs, vertebrae, long bone diaphysis).
Minimum Number of Elements (MNE) and 
Minimum Number of Individuals (MNI) were 
counted considering side, size and ontogeny (see 
Lyman 2008 for a discussion on the meaning of 
these indexes). Proximal and distal portions of 
long bones are considered separately in counting 
MNE. MAU and MAU% (Minimum Animal Unit, 
Binford 1984) were used to evaluate skeletal fre-
quencies. Bone density values were used following 
Lyman (1984); utility indices were used following 
Metcalfe, Jones (1988); Rowley-Conwy, Halstead 
and Collins (2002); Morin (2007).
Measurements were taken according to von den 
Driesch (1976) and are listed in the Appendix. 
Wear stages were measured according to Grant 
(1982) for the genera Bos and Sus and Payne 
(1973) for caprines. Identification of caprine foetal 
elements was carried out following Habermehl 
(1975); Ahmed (2008); Richardson et al. (1976) 
and Mcdonald et al. (1977). Age-at-death of roe 
deer and red deer were evaluated following Tomé 
and Vigne (2003), and Mariezkurrena (1983) and 
Habermehl (1985) respectively. The status of shaft 
portions at the moment of breakage (fresh or dry) 
was analysed using the Fracture Freshness Index 
(FFI) (Outram 2002).
Bone surface modifications were analysed with 
a 10X lens and, when necessary, with a binocular 
microscope; natural modifications were studied 
using present comparative materials collected 
on the field. In order to give more reliable data, 
only unburned fragments larger than 3 cm were 
considered in the analysis of weathering (cracks 
and exfoliation on bone surfaces).
Radiocarbon dates (Fig. 4) were calibrated with 
the OxCal 4.3 software (Bronk Ramsey 2009), 
using the IntCal 13 curve (Reimer et al. 2013). 
Statistics and graphs were performed using PAST 
software (Hammer, Harper, Ryan 2001) and R (R 
Development Core Team 2008).
RESULTS
Exclusion of a part of the faunal
assemblage from the present study
Layer 2a was excavated in two distinct areas 
of the cave, the first represented by squares 3, 
4, 8 and 9, and the second by squares 27–30 
and 33–35. A different faunal composition was 

326 Francesco BOSCHIN
detected between the two groups of squares: in 
particular, a higher proportion of wild mammal 
remains was detected in the innermost portion of 
the excavated area (squares 27, 28, 29, 30, 33, 34, 
and 35; Fig. 2): even if the number of remains in 
this area is lower than in the squares closer to the 
cave entrance (number of specimens is between 91 
in square 27 and 556 in square 35, and 1326, 1429, 
989, 1197 in squares 3, 4, 8, and 9, respectively), 
the proportion of wild mammals ranges between 
7.6% (square 30) and 66.6% (square 33). This is 
particularly high, considering that in squares 3, 4, 
8, and 9 this proportion is less than 1% (Fig. 5). In 
addition to caprines (NISP = 101), red deer (NISP 
= 34), roe deer (NISP = 3), wild boar (NISP = 5), 
and badger (NISP = 3) were identified in Layer 2a 
in the innermost part of the cave. With regard to 
the preservation of bones, this differs between the 
two areas (see more details in the chapter dedicated 
to taphonomy), and sediment preserved in the 
marrow cavity of long bones varies from a red clay 
in squares 27–35 to a grey soil in squares 3, 4, 8, 
and 9. Noteworthy carnivore marks and digested 
remains are absent in Layer 2a in squares 27–35, 
thus excluding a bone accumulation by carnivores 
in this part of the cave. It is also important to 
mention that some possibly reworked areas were 
detected in squares 30 and 33 during the excava-
tion of Layer 2a (Voytek’s field notes; excavation 
year 1998). Reworking can be the reason for the 
presence of Neolithic caprines mixed together with 
Mesolithic wild games. A part of the bones (NISP 
= 19) from squares 30, 34, and 35 was found in a 
Neolithic hearth called “feature 8” (Voytek’s field 
notes; excavation year 1998). Among the identi-
fied specimens from this feature, 15 are ascribed 
to domestic caprines, three to “Sus sp. ” and one to 
red deer, but due to the presence of the possible 
stratigraphic disturbances as regards Layer 2a, in 
order to avoid misinterpretations, only remains 
coming from squares 3, 4, 8 and 9 will be con-
sidered in this paper.
Faunal composition
About 69,000 macromammal specimens were 
analysed, but due to strong fragmentation, only 
a small percentage was identifiable (Tab. 1). Sau-
veterrian samples are characterized by an over-
whelming presence of red deer, roe deer, and wild 
boar (together they count for about 90% of NISP). 
Fig. 5: Grotta dell’Edera. Layer 2a, abundance of wild taxa 
according to square.
Sl. 5: Stenašca. Plast 2a, količina ostankov divjadi po 
kvadrantu.
Fig. 6: Grotta dell’Edera. Layer 2, abundance of domestic 
taxa according to depth of artificial spits. Caprines (dark 
grey); cattle (white); pig (grey).
Sl. 6: Stenašca. Plast 2, količina ostankov domačih živali 
po globinah: drobnica (temno siva), govedo (bela) in 
domači prašič (siva).

327 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
Faunal composition of these levels does not show 
any clear diachronic trend.
The small assemblage from Layer 3a shows 
mixed characteristics: domestic animals are the 
most abundant, but there is also a high presence 
of wild ones (especially red deer). A very similar 
situation is also present in the small assemblage 
from the (possibly reworked) Layer 3.
In light of new revisions, the faunal composition 
of Layer 2a is characterized by a high proportion of 
domestic caprines (96.9% of NISP), few domestic 
cattle and pig remains, and rare wild mammal 
remains (0.4% of NISP).
Layer 2 reveals a slightly different situation: 
caprines are the most abundant mammals (79.7% of 
NISP), and both cattle and pig are more represented 
(4.3% and 8.7%, respectively). Furthermore, the 
proportion of wild taxa is slightly higher (3.7%). 
Even if geoarchaeological data indicate a possible 
reworking of domestic livestock droppings by 
prehistoric herders (Boschian 1997), it is interest-
ing to note an increase in domestic pig and cattle 
remains from the bottom to the top of Layer 2 (Fig. 
6). In particular, there is a significant correlation 
between a decrease in pig+cattle frequencies and 
the depth of spits (p = 0.007).
In Layers 1a and 1b, NISP is very low (67 and 32, 
respectively). Caprines are still the most abundant 
taxa, the domestic pig is more represented than 
cattle, and wild ungulates are absent.
Taxa NISP
Layer 1 1a 1b 2 2a 3 3a 3b 3c 3d
domesticates
Bos taurus 3 9 2 22 10 2 - - - -
Ovis/Capra 6 36 19 386 852 28 50 - - -
Ovis aries - - - 14 22 2 4 - - -
Capra hircus - - - 4 7 1 - - - -
Sus domesticus 1 20 8 44 4 - 1 - - -
Canis familiaris - - 1 2 1 - - 1 - -
d/w
Bos/Cervus - - - 12 9 1 - - - -
Sus sp. - - - 4 - 2 7 - - -
wild
Cervus elaphus - - - 9 3 24 49 355 527 76
Capreolus capreolus - - - 2 - 14 4 233 488 22
Sus scrofa - - - 2 - 9 2 175 428 23
Bos primigenius - - - - - - - - 2 -
Capra ibex - - - - - - - - 3 -
Canis lupus - - - - - - - - 3 -
Vulpes vulpes - - - 2 - 1 1 4 10 3
Meles meles - - 2 1 - 5 2 59 59 3
Martes sp. - - - - - - - 12 4 2
Lutra lutra - - - - - - - 4 5 -
Ursus arctos - - - - - - - 1 1 1
Felis sylvestris - - - - - - - 9 7 -
Lynx lynx - - - - - - - - 1 -
Lepus sp. - 2 - 2 1 2 - 12 28 6
Castor fiber - - - - - - - - 5 -
Marmota marmota - - - - - - - - 1 -
Erinaceus sp. - - - 1 - - - 1 - -
NISP 10 67 32 507 909 91 120 866 1572 136
Tab. 1: Grotta dell’Edera. Identified specimens (NISP) according to layers.
Tab. 1: Stenašca. Taksonomsko opredeljeni ostanki (NISP) po plasteh.

328 Francesco BOSCHIN
Fig. 7: Grotta dell’Edera. – A: Size of specimens. The total amount of specimens used in the analysis corresponds to NR 
in Table 4. – B: FFI scored on diaphyseal fragments.
Sl. 7: Stenašca. – A: velikost kostnih najdb. Skupno število primerkov, ki so bili upoštevani pri tej analizi, se ujema z 
vrednostjo NR v tabeli 4. – B: vrednosti indeksa FFI za odlomke diafiz.

329 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
Preservation state of faunal remains
Fragmentation patterns observed in the dif-
ferent layers are quite similar and show a strong 
abundance of very small fragments (Fig. 7a). The 
1 cm size class reaches a score between 45% (level 
2a) and 66% (level 3d). Size classes over 3 cm 
cover a range between 24% (level 2a) and 4% (level 
3d). None of the fragments smaller than 1 cm is 
considered here. The most fragmented samples 
are those from Layers 3d (the earliest one) and 
2, whilst the least fragmented one is the sample 
from Layer 2a (squares 3, 4, 8 and 9).
The FFI (Fracture Freshness Index, Outram 
2002) was counted using shaft fragments from all 
of the layers with the exception of levels 1, 1a, and 
1b. The results are shown in Fig. 7b. The mean FFI 
score is often under three, thus corroborating the 
idea that the fragmentation pattern was partially 
created whilst the bones were fresh or near-fresh, 
but the high proportion of specimens showing 
scores between 3 and 6 indicates that some dam-
age occurred after the bones were dried.
Bones from Layers 2 and 2a show fewer natural 
modifications. From Layers 3 to 3d, exfoliation, 
root marks, erosion cups, and cracks became more 
frequent on fragments.
W eathering processes (cracks and exfoliation on 
the surfaces) are visible on a number of remains 
from Layer 2 to 3c (Tab. 2). It is worth noting that 
there is a significant and positive correlation be-
tween the abundance of weathered specimens and 
the mean FFI values registered on diaphyses for 
each layer (P uncorr. = 0.001), thus corroborating 
the hypothesis that perturbative processes affected 
bone samples, producing both bone breakage and 
surface modifications. It is intriguing to note that 
in the Mesolithic samples, weathering decreases in 
Layer 3c, when deposition of sediments appears 
to be faster. Within the more recent part of the 
sequence, bones from Layer 2a are better preserved 
than those from Layer 2, which appears to be 
characterized by slower sedimentation (Fig. 4).
Description of faunal remains
Red deer
Red deer is the most common wild species from 
the Mesolithic to the Eneolithic layers (Tab. 1). 
This situation seems to be common in the north 
Adriatic region (Turk et al. 1993, Tab. 6; Toškan, 
Dirjec 2004; Miracle 1997). Red deer remains 
are mostly represented by teeth, mandibles, and 
maxillary elements. Mortality profiles indicate 
the presence of a wide spectrum of age classes, 
including both young and adult individuals (Tab. 
3). With regard to young individuals, it appears 
that most of them were killed from spring to late 
summer (Fig. 8).
Roe deer
At Grotta dell’Edera, as in other sites on the 
Karst (Cannarella, Cremonesi 1967; Cremonesi et 
al. 1984; Cremonesi 1984; Turk et al. 1992, Tab. 
Layer % of weathered remain Sample size
2 30,1 216
2a (squares 4–9)
18,8 335
2a (squares 27–35)
60,9 92
3 68,8 128
3a 50,4 141
3b 59,4 498
3c 42,6 857
3d 84,7 131
T ab. 2: Grotta dell’Edera. Percentage of weathered remains according to layer. Only unburned 
specimens larger than 3 cm are considered.
Tab. 2: Stenašca. Delež kosti s sledmi izpostavljenosti vremenskim razmeram po plasteh. 
Upoštevani so bili zgolj nesežgani drobci velikosti nad 3 cm.

330 Francesco BOSCHIN
2; Turk et al. 1993, Tab. 5; Toškan, Dirjec 2004), 
roe deer is less abundant than red deer. A differ-
ent mortality profile (Tab. 3) is observed: young 
individuals are more represented than older ones. 
Regarding the seasonality pattern, this is the same 
as that of the red deer (Fig. 8).
Wild Boar/Domestic Pig
Specimens of the genus Sus are found in all of 
the layers. With regards to biometry, domestic 
individuals seem to be introduced from the Late 
Mesolithic Layer 3a, but the distinction between 
Tab. 3: Grotta dell’Edera. Age-at-death of individuals (MNI). Only more representative layers 
and taxa are considered. Absolute ages have to be considered approximate.
T ab. 3: Stenašca. Starost ob poginu/zakolu za posamezno žival (MNI). Upoštevane so zgolj bolje 
zastopane vrste in bogatejše plasti. Ocene absolutne starosti je treba obravnavati s previdnostjo.
Age classes
Layers
2 2a 3 3a 3b 3c
Cervus elaphus
(Age classes in years)
0–1 3 4
1–3 2 2
3–5 1 2
5–7 2 2
>7 2 2
Σ 10 12
Capreolus capreolus
(Age classes in years)
foetus 0 1
0–1 4 4
1–2 1 2
>2 0 1
6 1 1
Σ 6 9
Sus scrofa
(Age classes in years)
foetus 1 1
0–1 3 6
1–2 3 3
3–5 0 3
>6 1 2
Σ 8 15
Ovis/Capra
Payne’s (1973) age classes
foetal 5 5 0 1
A (0–2 months) 1 2 0 0
B (2–6 months) 2 2 1 1
C (6–12 months) 2 2 0 1
D (1–2 years) 0 2 0 0
E (2–3 years) 0 0 0 0
F (3–4 years) 1 2 0 0
G (4–6 years) 1 0 0 0
H (6–8 years) 0 1 0 0
I (8–10 years) 0 0 0 0
deciduous teeth dropped 
from jaws during life
3 7 2 3
Σ 15 23 3 6

331 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
two groups is sometimes impossible at the site. Even 
if the sample comprises very few data, individuals 
from the Neolithic layers are of smaller body size 
(Fig. 9). Using the standard for postcranial elements 
proposed by Payne and Bull (1988), the body size 
of Mesolithic individuals is similar to that of wild 
boars from Burgäschisee Süd (Switzerland, Neo-
lithic, ca. 3000–2500 BC; Payne, Bull 1988) and 
also similar, but slightly smaller, to that of wild 
boars from Zalog pri Verdu (Slovenia; Toškan, 
Dirjec 2006). A putative domestic pig scapula 
from Layer 2 is much smaller, but its measurement 
(SLC), as suggested by Albarella, Payne (2005), 
is age-dependent and has to be considered with 
Fig. 8: Grotta dell’Edera. Season of death of selected species from Layers 3c and 3b. Each line represents a skeletal ele-
ment. Letters on the top indicate months of the first three years of animals’ life.
Sl. 8: Stenašca. Sezona pogina za izbrane vrste iz plasti 3c in 3b. Vsaka črta predstavlja posamezen skeletni element. Črke 
vzdolž zgornjega roba označujejo mesece v prvih treh letih življenja posamezne živali.
Fig. 9: Grotta dell’Edera. Body size of pigs: Neolithic-Eneolithic individuals (grey); Mesolithic individuals (white).
Sl. 9: Stenašca. Ocena velikosti prašičev: neolitsko-eneolitsko gradivo (siva); mezolitsko gradivo (bela).
care. Furthermore, the trend in tooth size is dif-
ferent between the Neolithic and Mesolithic layers 
(Fig. 9). Even if the sample is affected by a very 
reduced size, more unfused post-cranial elements 
were found in Neolithic layers than in Sauveter-
rian ones. In particular, considering Layers 2 and 
2a together, more than 90% of limb bones (whole 
sample, n=14) belong to individuals younger than 
31–35 months; in the Sauveterrian layers (whole 
sample, n=129), this percentage falls to 45%.
Sauveterrian wild boars show a different mor-
tality profile than that of red and roe deer (Tab. 
3). In particular, there are more young individuals 
(neonatal and foetuses included). Most of them 

332 Francesco BOSCHIN
were killed from the spring to the autumn. Even 
though these data confirm those of the red and 
roe deer (Fig. 8), they are of more difficult inter-
pretation, due to the fact that wild boar sows can 
have more than one litter a year. Indeed, due to 
trophic availability, two reproductive seasons can 
occur, respectively, around September and April-
May (Massei, Toso 1993).
Domestic caprines
Sheep and goat are the most important domestic 
animals at Grotta dell’Edera. The first domesticates 
are those from the Late Mesolithic Layer 3a, where 
a direct radiocarbon date of a sheep bone fragment 
yielded a result of 6620±60 uncal BP (GrA-19820; 
Biagi 2003). Following Boessneck et al. (1964) and 
Payne (1985), sheep is more abundant than goat, 
as is often the case. Sex has been identified on 
a few fragmentary horn cores and pelvis bones: 
one sheep female horn core is from Layer 2; two 
sheep female horn cores (one hornless and one 
atrophied) and one cf. goat male horn core are 
from Layer 2a. Among innominate fragments, one 
sheep male, one sheep/goat male, two sheep/goat 
females and one cf. goat female were identified. 
Age-at-death is estimated using both teeth and 
long bones (foetuses) (Tab. 3). Age of most of the 
young animals is obtained from Pd4 and Pd3. All 
M3-in-eruption-age observations are obtained from 
the abrasion of the Pd4, estimated “n” (following 
Grant 1982). Since all of these teeth seem to have 
dropped from jaws of still-living animals, the M3-
in-eruption-age was deleted from the mortality 
profile to avoid misinterpretations. Individuals 
corresponding to this age are reported as “decidu-
ous teeth dropped from jaws during life” in Tab. 
3. Indeed, even if the problem was highlighted 
in the previous report (Boschin, Riedel 2000), 
caprine individuals corresponding to that age were 
considered when mortality profiles from Grotta 
dell’Edera were compared with those from other 
sites (e.g. Mlekuž 2005). More than 35% of post-
cranial MNE (phalanxes excluded), both in Layers 
2 and 2a, is related to foetal individuals (Boschin 
2020). On the basis of bone length (Habermehl 
1975), the bulk of samples appears to belong to 
late or very late gestation foetuses (estimated age 
between 110–120 and 136–145 days) (Fig. 10).
Cattle/Aurochs
Cattle remains of both wild and domestic form 
are rare at the site. Wild animals are represented in 
the Mesolithic only by two remains from Layer 3c.
Individuals from other prehistoric levels have 
a very large size, comparable with that of female 
aurochs (about 130 cm high or more; Boschin, 
Riedel 2000) and of domesticates from other Neo-
lithic sites (such as Piancada in Friuli; Petrucci, 
Riedel, Pessina 2000).
Dog
The earliest dog remains are that from Layer 3b. 
It is a fused IV metatarsus that belongs to a small 
individual (withers height of about 48 cm, using 
the factor proposed by Clark 1995; measurements 
are: GL: 67.6; SD: 5.2; Bd: 8.5). Other Sauveterrian 
dogs similar in size were recovered in Slovenia 
Fig. 10: Grotta dell’Edera. Age of caprine foetuses (in 
weeks) from Layers 2 and 2a. Bones corresponding to the 
age “145+” can also belong to new-borns. Age is estimated 
here following Habermehl (1975) and considering the 
bones as belonging to sheep. For a detailed discussion of 
foetal mortality at site see Boschin (2019).
Sl. 10: Stenašca. Starost ob poginu (v tednih) za zarodke 
drobnice iz plasti 2 in 2a. Kosti, ki so bile uvrščene v razred 
“145+” , bi načeloma lahko pripadale komaj skotenim živalim. 
Starost ob poginu je bila ocenjena na podlagi smernic, ki 
jih je objavil Habermehl (1975), s predpostavko, da najdbe 
pripadajo ovcam. Za podrobnejšo obravnavo problematike 
smrtnosti zarodkov v Stenašci glej Boschin (2019).

333 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
(Pohar 1984), whilst a Castelnovian remain was 
identified at Viktorjev spodmol in the Slovenian 
Karst (Toškan, Dirjec 2004). Other bones from 
Layer 2 of Grotta dell’Edera also belong to small 
individuals. A talus from Layer 1b (Bronze Age) 
is 25.6 mm long, belonging to an individual that 
is still showing a small size. Dogs similar in size 
are present elsewhere during the same period (for 
instance, see De Grossi Mazzorin, T agliacozzo 1997).
Other taxa
An upper premolar, a proximal metacarpus and 
a distal femur from Layer 3c are attributed to ibex. 
Few specimens of this taxon were recognized in 
the Grotta Azzurra/Pečina na Leskovcu on the 
Trieste Karst (Cannarella, Cremonesi 1967). Due to 
the high fragmentation, the femur was previously 
attributed with a high level of uncertainty to a 
chamois (Boschin, Riedel 2011), but the comparison 
with other modern ibex bones seems to exclude 
this possibility. With regards to neighbouring 
regions, Rupicapra specimens were identified in 
Late Pleistocene/early Holocene sites in Slovenia 
at Betalov spodmol (Toškan, Dirjec, Bavdek 
2014), Istria (Miracle 1997; Miracle, Galanidou, 
Forenbaher 2000) and in Riparo di Biarzo in the 
Natisone/Nadiža valley (Rowley-Conwy 1996).
Badger is present in most of the layers (Tab. 1). 
It is the most common carnivore, as observed in 
other cave sites of the area (Cannarella, Cremonesi 
1967; Cremonesi 1984; Cremonesi et al. 1984; Bon 
1996; Miracle 1997; Toškan, Dirjec 2004). Age 
profiles show the presence of adult and young 
individuals. One from Layer 3b is younger than 
3–4 months, and one in Layer 3d is about 4 months 
old (following Habermehl 1985).
Marten is represented by few specimens from 
several layers. With regards to mandible, skull, and 
teeth, Mesolithic remains appear to belong to the 
pine marten Martes martes (following Hans, Steiner 
1986). Specimens from Layers 3 and 2a were given 
only generic determination. Beech marten Martes 
foina is present in the Neolithic site of the Grotta 
degli Zingari/Ciganska jama (Bon 1996) and its 
occurrence at Grotta dell’Edera cannot be ruled out.
The otter is represented by some long bones, 
one talus and one mandible from Layers 3b and 
3c. All of the long bones are completely ossified 
and belong to adult individuals.
Fox bones were identified in both the Neolithic 
and Mesolithic levels. All of the individuals are adult.
Wild cat remains were found both in Mesolithic 
and Neolithic layers, whilst a lynx mandible comes 
from level 3c. This animal is very rare in the ter-
ritory and was recognized only in few cave sites 
(Cremonesi 1984; Toškan, Dirjec 2004; possibly 
also Turk et al. 1993, Tab. 5). Measurements (M/1, 
length = 15.4) fit in the variability of extant Lynx 
lynx (Hemmer 1993).
The brown bear is present in the three earliest 
levels with one specimen in each. This taxon is 
not frequent in the other sites of the area (Toškan, 
Dirjec 2004; Turk, Toškan, Dirjec 2004).
The wolf is represented by three specimens from 
Layer 3c. Hare is represented by few remains. In 
other sites, the species was identified as Lepus eu-
ropaeus (Cannarella, Cremonesi 1967; Cremonesi 
1984; Cremonesi et al. 1984; Bon 1996; Toškan, 
Dirjec 2004). Since diagnostic elements at Grotta 
dell’Edera are missing (skull fragments), distinguish-
ing between the various taxa (i.e. L. europaeus, L. 
timidus and L. corsicanus) is not possible.
Marmot is present with a distal femur in Layer 
3c. It would be the only identified Holocene speci-
men of this taxon from the Karst, even if putative 
Holocene marmot remains were found at Betalov 
spodmol (Postojna, SW Slovenia; Toškan, Dirjec, 
Bavdek 2014).
Beaver is represented by few bones from Layer 3c. 
This animal was also identified in the Grotta Azzurra/
Pečina na Leskovcu (Cannarella, Cremonesi 1967), and 
an unpublished beaver femur, found by Karl Moser 
in the Vlaška jama, is part of the zooarchaeologi-
cal collections of the Naturhistorisches Museum 
Wien. In addition, this taxon was also identified 
at Mala Triglavca (Turk, Toškan, Dirjec 2004).
Hedgehog is present only in Layers 2 and 3b. 
The taxonomic identification is uncertain between 
the species europaeus and roumanicus, whose 
geographical distributions show an overlap in the 
Karst territory today (Lapini et al. 1996).
Human modifications on bones
Burned bones
Burned fragments were retrieved in the whole 
sequence and are very abundant in Layers 2 and 
3d. In this level, they are concentrated in squares 
3, 8, 9, and 23. In Layers 3a and 3b, they are less 
numerous (they are concentrated in squares 3 and 
8 in 3b), whilst in the other Layers (2a, 3, 3c) they 
never rich 20% (Tab. 4).

334 Francesco BOSCHIN
Other human modifications
Butchering marks were analysed throughout the 
sequence. Samples from Layers 2 and 2a are quite 
poor in this kind of traces, and only some cuts 
on shaft portions and some butchering marks on 
few caprine and pig vertebrae were observed. In 
addition, rare impact scars were identified among 
diaphyseal fragments (one in Layer 2), and cone 
flakes due to bone breakage were detected both in 
Layer 2 (n = 11) and 2a (n = 4). Only two impact 
scars and two cone flakes were identified among 
remains from Layer 3a.
Butchering marks are more frequent in Layers 
3b and 3c. Indeed, there are anthropic modifica-
tions related to all phases of carcass treatments 
on ungulate remains (skinning, disarticulation, 
filleting and bone breakage). There are 13 impact 
scars on diaphysis in Layer 3b and 44 in Layer 3c; 
18 cone flakes are  in Layer 3b and 44 in Layer 3c. 
Marrow extraction or grease-rendering activities 
are also indicated by phalanges that show a stand-
ardized fragmentation. In addition, a number of 
human modifications were detected on lagomorph, 
marmot, and carnivore remains. These latter are 
summarized in Tab. 5. W orked bones are very rare 
(Biagi, Starnini, Voytek 2008).
Evidence of commensalisms
Evidence of bone modifications by carnivores and 
small rodents was recognized in the cave. Most of 
the gnawing marks were observed in Layers 3b (on 
38 specimens) and 3c (on 53 specimens) and appear 
to have been largely caused by small carnivores, 
such as dog, fox, or badger. They consist of small 
punctures, pitting furrows, or intensively chewed 
or scooped-out epiphysis. Some small specimens 
from Layer 2a (n = 6) and 3c (n = 6) appear to be 
affected by digestive processes.
A proximal roe deer humerus from Layer 3b shows 
both gnawing marks and burned surfaces, whilst 
two roe deer tarsals from Layer 3c, which belong 
to the same individual, show gnawing marks (on 
the talus) and cut marks (on the cubonavicular).
Body part profiles
Body parts are shown in Tabs. 6–8, but they were 
analysed in-depth only for Layers 2 and 2a (domestic 
caprines), and 3b and 3c (red deer, roe deer and 
wild boar) (Tab. 9); data from other layers were not 
taken into account due to the sample size. With 
regards to the main taxa, with the exception of Layer 
3c, there is no significant correlation between body 
part representation (MAU%) and structural bone 
density. An insignificant correlation was detected in 
most of the samples also between body part repre-
sentation (MAU%) and Standardized Food Utility 
Index (SFUI) (Tab. 10). It is worth noting that the 
relationship is always negative. Since percussion 
marks testified to the exploitation of phalanges for 
marrow extraction in Sauveterrian Layers 3b and 
3c, a relation between the Unsaturated Marrow 
Index (UMI) and MAU% was tested. P-values are 
always not significant (for red deer, roe deer, and 
Neolithic caprines), ranging from 0.16 (caprines, 
Layer 2) and 0.93 (roe deer, Layer 3b).
Among the identified specimens, head ele-
ments are the best represented in all cases. This is 
probably due to the good identifiability of teeth. 
Among the Sauveterrian taxonomically unidenti-
fied specimens, the most abundant are shaft frag-
ments, ribs, vertebrae, and teeth fragments (Tab. 
11). Among caprines in Layers 2 and 2a, the lack 
of phalanges has to be underlined.
DISCUSSION
Sauveterrian
Hunter-gatherers from Layer 3d are among the 
first humans to frequent the Trieste Karst after 
the Late Glacial. Several data seem to suggest that 
the area was uninhabited (or rarely frequented 
% of burned
specimens
% of calcined
among burned
NR
2 36,5% 8,5% 8061
2a 17% 2% 4941
3 11% 0,3% 1655
3a 22,6% 0,5% 2179
3b 23% 3,5% 15107
3c 15% 3,1% 31212
3d 35% 0% 4106
Tab. 4: Grotta dell’Edera. Abundance of burned specimens 
per layer. (NR = total number of remains: burned and 
unburned; in each layer).
Tab. 4: Stenašca. Številčnost sežganih ostankov po plasteh. 
(NR = skupno število ostankov: sežganih in nesežganih; 
po plasteh).

335 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
Taxon Element Anthropic modification Layer NISP Supposed Action
Canis lupus pubis cut marks 3c 1 disarticulation/filleting
Canis lupus scapula cut marks 3c 1 disarticulation
Martes sp. cuboid calcined 3c 1 intentionally burned
Meles meles femur carbonized 3b 1 burned
Meles meles humerus carbonized 3b 1 burned
Meles meles mandible carbonized 3b 1 burned
Meles meles metapodial carbonized 3c 1 burned
Meles meles metapodial carbonized 3c 6 burned
Meles meles metapodial carbonized 3c 2 burned
Meles meles phalanx carbonized 3b 1 burned
Meles meles phalanx carbonized 3b 1 burned
Meles meles phalanx carbonized 3c 1 burned
Meles meles phalanx carbonized 3c 1 burned
Meles meles phalanx carbonized 3c 1 burned
Meles meles phalanx carbonized 3c 1 burned
Meles meles radius carbonized 3c 1 burned
Felis silvestris radius cut marks 3c 1 filleting
Lynx lynx mandible cut marks 3c 1 skinning
small carnivore calcaneum cut marks 3c 1 disarticulation
small carnivore caudal vertebra calcined 3c 1 intentionally burned
small carnivore innominate carbonized 3b 1 burned
small carnivore metapodial calcined 3c 1 intentionally burned
small carnivore metapodial carbonized 3c 1 burned
small carnivore metapodial carbonized 3c 1 burned
small carnivore metapodial cut marks+calcined 3c 1 burned + disarticulation
small carnivore metapodial fresh bone fracture 3c 1 disarticulation
small carnivore metapodial fresh bone fracture 3c 1 disarticulation
small carnivore phalanx calcined 3c 1 intentionally burned
small carnivore phalanx calcined 3c 1 intentionally burned
small carnivore phalanx carbonized 3c 1 burned
small carnivore proximal ulna cut marks+carbonized 3c 1 burned + disarticulation
small carnivore radius fresh bone fracture 3c 1 disarticulation
small carnivore ulna carbonized 3c 1 burned
Lepus sp. calcaneum carbonized 3c 1 burned
Lepus sp. cervical vertebra carbonized 3b 1 burned
Lepus sp. metapodial calcined 3c 1 intentionally burned
Lepus sp. metapodial calcined 3b 1 intentionally burned
Lepus sp. phalanx calcined 3c 1 intentionally burned
Lepus sp. phalanx calcined 3c 1 intentionally burned
Lepus sp. phalanx calcined 3c 1 intentionally burned
Lepus sp. ulna carbonized 3b 1 burned
Marmota marmota femur carbonized 3c 1 burned
Tab. 5: Grotta dell’Edera. Anthropic modifications on lagomorph, rodent and carnivore remains.
Tab. 5: Stenašca. Popis zajčjih, glodavskih in zverskih kosti s sledmi človekovih dejavnosti.

336 Francesco BOSCHIN
by humans) during the Upper Palaeolithic due to 
environmental constraints (harsh conditions, arid-
ity, rocky landscape poorly covered by vegetation; 
Boschian, Fusco 2007). At the beginning of the 
Holocene, with the onset of more humid conditions, 
followed by the spread of wooded environments, 
the Karst became more suitable for humans. At 
the same time, human groups were also pushed 
northwards by the marine transgression, which 
led to the reduction of the Adriatic Plain. A dif-
ferent situation, more vegetated and suitable for 
hunter-gatherers, characterized the nearby regions, 
such as Istria, where both Upper Palaeolithic and 
Mesolithic sites have been found (Boschian, Fusco 
2007; Jamnik et al. 2015; Pilaar-Birch, Miracle 
2017). Human presence in Layer 3d occurred just 
after the abrupt warming oscillation at the end of 
the Greenland Stadial 1 (Rasmussen et al. 2007). 
It appears as an isolated episode of frequentation, 
followed by a long period of absence, until the 
9.3kyrs event, a period of widespread climatic 
deterioration (Fleitmann et al. 2008), when hu-
Bt C/O CH Oa Sd
Layer 2 2a 3 2 2a 3 3a 2 2a 3 2 2a 3 3a 2 2a 3a
Antler
- - - - - - - - - - - - - - - - -
Horn core
- - - - - - 2 1 1 - 1 1 - - - - -
Skull
1 3 - 32 52 3 1 - - - 1 1 - - 4 - -
Maxilla
1 1 - 3 7 - 2 - - - - - - - - - -
Mandible
- 1 - 5 26 - 1 - - - - - - - 1 - -
Isolated teeth
10 1 - 139 217 20 15 - - - - 2 - - 18 - 1
Hyoid
- - - 2 2 - - - - - - - - - - - -
Vertebrae
1 - - 26 32 1 10 - - - - 1 1 - 4 - -
Ribs
- 1 - 71 332 2 5 - - - - - - - 2 - -
Sternum
- - - - - - - - - - - - - - - - -
Scapula
3 - - 6 9 - 1 - 1 - - - - 2 2 - -
Humerus
- - - 8 17 - - - - - - - - - - - -
Radius
- - - 4 16 1 2 1 - - - - - - - - -
Radius-ulna
- - - - 1 - - - - - - - - - - - -
Ulna
- - - 5 11 - - - - - - 1 - - - - -
Carpals
- - - 14 3 - 4 1 - - - - 1 - - - -
Metacarpal
- - 1 11 19 - 3 - - - 1 4 - - - - -
Plesiometacarpal
- - - - - - - - - - - - - - - - -
Innominate
- 1 - 3 6 - - - 1 - - 2 - - - - -
Femur
1 - - 4 13 - 2 - - - - 1 - - 1 1 -
Patella
- - - 1 2 - - - - - - - - - - - -
Tibia
- - - 6 21 - - - - - - - - - - - -
Fibula
- - - - - - - - - - - - - - 1 2 -
Malleolar
- - - 1 - - - - - - 1 - - - - - -
Tarsals
- 1 - 7 7 1 - - 1 1 - 1 - - - - -
Metatarsal
1 1 - 18 33 - - 1 1 - 4 2 - - 2 - -
Metapodial
2 - - 14 18 - 2 - - - 3 4 - - 2 1 -
Phalanxes
2 - - 6 7 - - - 2 - 3 2 - 2 5 - -
Sesamoidal bones
- - 1 - 1 - - - - - - - - - 2 - -
Σ
22 10 2 386 852 28 50 4 7 1 14 22 2 4 44 4 1
Tab. 6: Grotta dell’Edera. Body parts (NISP) of domestic ungulates.
Tab. 6: Stenašca. Zastopanost anatomskih regij (NISP) pri domačih kopitarjih. 
Bt: Bos Taurus; C/O: Capra/Ovis; Ch: Capra hircus; Oa: Ovis aries; Sd: Sus domesticus

337 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
Tab. 7: Grotta dell’Edera. Body parts (NISP) of wild ungulates. 
Tab. 7: Stenašca. Zastopanost anatomskih regij (NISP) pri divjih kopitarjih.
Cc: Capreolus capreolus; Ce: Cervus elaphus; Ss: Sus scrofa; Ci: Capra ibex; Bp: Bos primigeniusv
man frequentation is testified to by evidence from 
Layer 3c. The presence of hunter-gatherer groups 
in the Edera Cave became more stable, as testified 
by 
14
C dates of the overlying Layer 3b (Fig. 4). 
Evidence of the latter layer is the last occurrence of 
Sauveterrian techno-complex at Grotta dell’Edera; 
indeed, it seems that a lack of frequentation which 
lasted for about one thousand years, may also 
be partially related to the 8.2kyrs event, another 
period of climatic deterioration (Rasmussen et al. 
2007). This climatic event can be detected more 
clearly (was it also more intense?) than the one 
at 9.3kyrs in the isotopic record registered in a 
stalagmite from the Grotta Savi on the Trieste 
Karst (Borsato et al. 2003).
Sauveterrian faunal composition of Grotta 
dell’Edera is not different from those from other 
cave sites in the Karst territory and Istria (Can-
narella, Cremonesi 1967; Cremonesi 1984; Cremo-
nesi et al. 1984; Bon 1996; Miracle 1997; Miracle, 
Galanidou, Forenbaher 2000; Toškan, Dirjec 
2004). Identified thanatocoenoses appear to be 
Cc Ce Ss Ci Bp
Layer 2 3 3a 3b 3c 3d 2 2a 3 3a 3b 3c 3d 2 3 3a 3b 3c 3d 3c 3c
Antler
- 1 - 1 - - - - - - 3 20 1 - - - - - - - -
Horn core
- - - - - - - - - - - - - - - - - - - - -
Skull
- - 1 10 22 2 - - 1 5 4 16 1 - - - 2 14 1 - -
Maxilla
- - - 2 2 1 - - - - 1 4 1 - - - 1 2 - - -
Mandible
- 1 - 1 10 - 3 - 2 - 7 11 2 - 1 - 5 3 - - -
Isolated teeth
1 2 1 28 43 3 1 1 3 15 92 103 33 1 2 - 19 50 5 1 -
Hyoid
- - - - - - - - - - 1 - - - - - - - - - -
Vertebrae
- 3 - 32 42 - 1 - 4 2 23 31 1 - - - 6 38 1 - -
Ribs
- 1 - 18 32 - - - 1 4 34 23 2 - - - 4 6 - - -
Sternum
- - - - - - - - - - 1 - - - - - - - - - -
Scapula
- - - 3 4 - - - 1 1 5 - 1 - - - 3 5 - - -
Humerus
- - - 3 12 2 - - 1 - 5 5 1 - - - 3 11 1 - -
Radius
- - - 2 10 - - - - 1 1 6 1 - - - 3 18 - - -
Radius-ulna
- - - - - - - - 1 - - - - - 1 - - - - - -
Ulna
- 1 - 6 6 1 1 1 - 3 6 6 - - - - 8 16 1 - -
Carpals
- - - 6 10 - - - - 1 6 11 3 - - - 3 10 - - -
Metacarpal
- - - 9 34 1 - - 1 3 25 30 1 - 1 - 8 20 - 1 -
Innominate
- 1 - 4 3 - - - - - 6 9 - - - - 2 6 1 - -
Femur
- 1 - 9 5 - - - 1 - 2 9 - - - - 2 2 1 1 -
Patella
- - - - - - - - - - - 1 - - - - 1 2 - - -
Tibia
- 1 1 7 10 1 - - - - 3 5 1 - - - 1 12 - - -
Fibula
- - - - - - - - - - - - - - - - 3 9 - - -
Malleolar
- - - - 1 - - - - - - 3 - - - - - - - - -
Tarsals
- - - 15 28 - - - 1 3 8 17 4 - - - 4 6 - - 2
Metatarsal
1 1 - 12 31 1 1 - 1 27 38 3 - - - 4 21 4 - -
Metapodial
- - - 7 16 - 1 - 2 1 5 15 4 - 1 2 20 38 - - -
Phalanxes
- 1 1 49 134 9 1 1 4 9 82 142 12 1 3 - 68 116 6 - -
Sesamoidal bones
- - - 9 33 1 - - - 1 8 22 4 - - - 5 23 2 - -
Σ
2 14 4 233 488 22 9 3 24 49 355 527 76 2 9 2 175 428 23 3 2

338 Francesco BOSCHIN
composed by taxa that may have been hunted in 
the wooded environments surrounding the cave. 
Exceptions consist of taxa that could have been 
hunted in other areas more or less far from Grotta 
dell’Edera: in this regard, otter and beaver probably 
lived in freshwater habitats around the Karst, such 
as along the rivers Timavo (Reka), Isonzo (Soča), 
Vipacco (Vipava), Ospo (Osp) or Rižana. As for 
the otter, the small polje of Doberdò (Doberdob) 
and that of Pietrarossa (Prelosno jezero), located 
on the western side of Karst plateau, may have 
also represented suitable environments.
Elk was not identified at Grotta dell’Edera, but its 
remains, even if rare, were yielded by early Holocene 
deposits of the area: Viktorjev spodmol (Toškan, 
Dirjec 2004), Vlaška jama (unpublished material 
collected by Karl Moser; Naturhistorisches Museum 
Wien, Archäologisch-Zoologische Sammlung) and 
Grotta Benussi (Riedel 1976).
A suitable environment for some of the above-
mentioned taxa was probably also represented by 
the Gulf of Trieste: indeed, sedimentological and 
geological evidence (Marocco 1991; Boschian 1993) 
indicates a considerably reduced sea covered area 
(especially in the north-western side) during the 
early Holocene. A relatively small plain crossed 
by small freshwater streams (there are many un-
derwater springs along the present-day coastline) 
must have been a very resourceful environment 
for Mesolithic groups. Pond-turtle remains found 
at Grotta dell’Edera (Delfino, Bressi 2000) also 
suggest the exploitation of this area.
Marmot and ibex definitely inhabited the 
Karst plateau during the various arid phases of 
Cf C1 Vv Mm M sp.
Layer
2 2a 3b 3c 2 3 3a 3b 3c 3d 2 3 3a 3b 3c 3d 3b 3c 3d
Skull - - - - - - - - - - - - - 5 3 - - - -
Maxilla - - - - - - - - 1 - - - - - 1 - 1 - -
Mandible - - - - - - - - - - - - - 4 - - - 1 -
Isolated teeth - 1 - - - - - - 3 1 1 - - 16 8 1 5 - -
Vertebrae - - - - - - - 1 1 - - - - 1 2 - - - -
Ribs - - - - - - - - - - - - - 3 2 - - - -
Sternum - - - - - - - - - - - - - 1 - - - - -
os penis - - - - - - - - - - - - - 1 - - - - -
Scapula - - - 1 - - - - - - - - - - - - - 1 -
Humerus - - - - - - - 1 - - - - - - 1 - 1 - -
Radius - - - - - - - - - - - - - 2 1 - 1 1 -
Radius-ulna - - - - - - - - - - - - - - - - - - -
Ulna - - - - - - - - 1 - - - - - - - 1 1 -
Carpals - - - - - - - - - - - - - 3 2 - - - -
Metacarpal - - - - - - - 1 - - - 1 1 6 8 - 2 - 1
Innominate - - - 1 - - - - - - - - - - - - - - 1
Femur - - - - - - - - - - - - - 3 1 - - - -
Patella - - - - - - - - - - - - - - - - - - -
Tibia - - - - - - - - - - - - - 1 - - - - -
Tarsals - - - - - 1 - - - - - - - 1 1 - - - -
Metatarsal 1 - 1 - 2 - - - 1 - - - - 4 7 - - - -
Metapodial - - - - - - - - 1 - - 1 - - 12 - - - -
Phalanxes 1 - - 1 - - 1 1 2 2 - 3 1 8 10 2 1 - -
Σ 2 1 1 3 2 1 1 4 10 3 1 5 2 59 59 3 12 4 2
Tab. 8: Grotta dell’Edera. Body parts (NISP) of carnivores, lagomorphs, rodents and insectivores.
Cf: Canis familiaris; Cl: Canis lupus; Vv: Vulpes vulpes; Mm: Meles meles; M sp.: Martes sp; Lut: Lutra lutra; Ua: Ursus 
arctos; Fs: Felis silvestris; Ll: Lynx lynx; L. sp.: Lepus sp; Cas: Castor fiber; Mar: Marmota marmota; E. sp.: Erinaceus sp.

339 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
the Upper Pleistocene: ibex was recovered in a 
number of cave sites ascribable to this period 
(Caverna degli Orsi; Boschian 2001; Caverna 
Piccola Pocala, unpublished material identified 
by the author, collections of the Natural History 
Museum of Trieste) and in some quarries along 
rocky slopes behind the city of Trieste (unpub-
lished material identified by the author, collec-
tions of the Natural History Museum of Trieste). 
Furthermore, marmot was identified in some 
Upper Pleistocene sites (for instance Bartolomei 
2004), but the specimen from Grotta dell’Edera 
is the only one from a Holocene deposit on the 
Trieste Karst. Noteworthily, the presence of both 
ibex and marmot at Grotta dell’Edera corresponds 
to the cooler oscillation at 9.3kyrs (Rasmussen 
et al. 2007).
Faunal composition from the Sauveterrian layers 
indicates the presence of wooded vegetation (red 
deer, roe deer, wild boar) and open environments 
(ibex, marmot, hare), whose presence decreased 
moving from the Boreal to the Atlantic period. 
This reconstruction is in agreement with archaeo-
botanical data (Galizia Vuerich, Princivalle 1994; 
Nisbet 2000) that indicate the presence of a boreal 
forest of Pinus and Betula, probably mixed with 
grasslands, during the early Holocene. Moreover, 
sedimentological evidence indicates a scarcely af-
forested territory with easily erodible soils during 
this phase (Boschian 1997). Therefore, especially 
during Preboreal and early Boreal, residual open 
landscapes may have favoured ibex and marmot. 
The presence of taxa related to arid open environ-
ments was also detected among small mammals in 
Lut Ua Fs L1 L sp. Cas Mar E sp.
Layer
3b 3c 3b 3c 3d 3b 3c 3c 2 2a 3 3b 3c 3d 3c 3c 2 3b
- - - - - 1 - - - - - - - - - - - - Skull
- - - - - - - - - 1 - - 2 - - - - - Maxilla
1 - - 1 - - 1 1 - - - - - - - - - - Mandible
- - 1 - 1 1 - - - - 1 - 11 3 1 - - - Isolated teeth
- 2 - - - - - - - - - 1 1 - 1 - - - Vertebrae
- - - - - - - - - - - - - - - - - - Ribs
- - - - - - - - - - - - - - - - - - Sternum
- - - - - - - - - - - - - - - - - - os penis
- - - - - - - - - - - - - - - - - - Scapula
1 1 - - - - - - - - 1 1 1 - - - - - Humerus
- - - - - - 1 - - - - 1 - - - - - - Radius
- - - - - - - - - - - - - - - - - - Radius-ulna
- 2 - - - - - 1 - - 2 - - - - - - Ulna
- - - - - - - - - - - - - - - - - - Carpals
- - - - - 2 1 - - - - - - - - - - - Metacarpal
- - - - - 1 1 - - - - 1 - - 1 - 1 1 Innominate
1 - - - - - - - - - - - - - - 1 - - Femur
- - - - - - - - - - - 1 - - - - - - Patella
- - - - - - - - - - - 2 1 1 - - - - Tibia
1 - - - - - - - - - - 2 1 - - - - - Tarsals
- - - - - 2 1 1 - - - 5 - - - - - Metatarsal
- - - - - 1 - - - - - - - - - - - - Metapodial
- - - - - 1 2 - - - - 1 6 2 2 - - - Phalanxes
4 5 1 1 1 9 7 1 2 1 2 12 28 6 5 1 1 1 Σ
Tab. 8: Stenašca. Zastopanost anatomskih regij (NISP) pri zvereh, zajcih in glodavcih.
Cf: Canis familiaris; Cl: Canis lupus; Vv: Vulpes vulpes; Mm: Meles meles; M sp.: Martes sp; Lut: Lutra lutra; Ua: Ursus 
arctos; Fs: Felis silvestris; Ll: Lynx lynx; L. sp.: Lepus sp; Cas: Castor fiber; Mar: Marmota marmota; E. sp.: Erinaceus sp.

340 Francesco BOSCHIN
MNE Capra/Ovis Capreolus capreolus Cervus elaphus Sus scrofa
Layer 2 2a 3b 3c 3b 3c 3b 3c
Maxilla 5 12 5 9 7 8 13 33
Mandible 10 7 6 7 10 11 12 22
Hyoid 2 2 0 0 1 0 0 0
Atlas 0 1 2 0 0 0 0 0
Axis 2 0 0 1 0 0 0 0
Cerv. Vert. 4 4 2 4 5 3 1 19
Thor.Vert. 2 10 6 4 3 5 1 10
Lumb. Vert. 4 5 7 7 3 5 4 4
Sacrum 0 0 0 1 2 2 0 3
Caud. Vert. 3 0 0 2 1 1 0 0
Ribs 0 20 0 0 0 0 4 6
Sternum 0 0 0 0 0 0 0 0
Scapula 4 6 2 2 2 0 3 5
Humerus p. 5 6 0 3 2 1 0 1
Humerus d. 6 9 2 5 2 2 3 4
Radius p. 3 6 2 2 1 4 1 7
Radius d. 4 7 0 4 0 1 0 5
Ulna p. 4 12 5 4 5 5 6 11
Ulna d. 0 0 0 1 1 0 2 3
Carpals 15 2 5 10 6 11 3 10
Pelvis 2 5 3 1 4 3 2 6
Femur p. 1 3 1 2 2 0 1 1
Femur d. 2 3 2 3 0 4 1 0
Patella 1 1 0 0 0 0 1 2
Tibia p. 2 7 3 5 1 2 0 3
Tibia d. 3 6 2 3 1 2 1 6
Fibula/Malleollar 0 0 1 1 0 3 3 5
Tarsals 8 5 15 26 7 17 4 6
Metapodial p. 11 13 1 11 8 12 10 17
Metapodial d. 13 15 9 10 10 12 6 18
Metapodial lat. 0 0 0 9 0 0 6 18
Phalanx 1 5 5 17 38 16 23 15 27
Phalanx 2 3 1 10 22 12 23 14 22
Phalanx 3 0 1 10 15 7 11 16 18
Phalanx lat. 1 0 0 0 8 2 8 6 12
Phalanx lat. 2 0 0 0 5 4 21 7 15
Phalanx lat. 3 0 0 1 0 0 0 10 22
Sesamoides 0 1 9 33 8 22 5 23
Σ 124 175 128 258 133 222 161 364
Tab. 9: Grotta dell’Edera. Minimum number of elements of main taxa. 
Tab. 9: Stenašca. Vrednost indeksa najmanjše število ostankov (MNE) za posamezne skeletne elemente najpomembnejših 
taksonov.
Vert. : V ertebra. – Cerv.: cervical. – thor.: thoracic. – Lumb.: lumbar. – Caud.: caudal. – p.: proximal. – d.: distal. – lat.: lateral.

341 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
the Boreal sample from Mala Triglavca (Toškan, 
Kryštufek 2006; Toškan 2009) and Grotta degli 
Zingari (Bartolomei 2001). A broadleaf forest 
covered the territory during the Atlantic period 
(Nisbet 2000), leading to the extinction of open-
environments-related taxa.
Slicing cut marks, percussion marks, cones, 
and burned specimens point to Mesolithic people 
as the main accumulators of Sauveterrian faunal 
assemblages. In addition, among the few gnawed 
bones, some of them were previously modified by 
humans. Taphonomy also ruled out an intrusive 
origin of rodents, lagomorphs and carnivores, in-
dicating the exploitation of a wide variety of taxa, 
as observed in other Mesolithic sites of northern 
Italy (Crezzini et al. 2014).
A reduction of skeletal portions by means of 
percussion is attested on limb bones. Lower fre-
quency of axial elements can be due to their lower 
identifiability, and this is testified by the high 
quantity of ribs and vertebrae among unidentified 
fragments. Thus, ungulate carcasses were likely 
brought complete to the site. Available data do not 
support the idea that selected parts were brought 
to site depending on their utility as food resources 
or, more specifically, as sources of fat. Regardless, 
once carcasses were processed on-site, every part 
was exploited, including the small quantity of mar-
row within a single phalanx.
Differences in age-class distribution observed 
among main wild ungulate taxa (red deer, roe deer 
and wild boar) are probably due to the different 
composition of herds and to their behaviour. Data 
on tooth eruption confirm that hunting activities 
were mainly carried out in a period between spring 
and autumn (Fig. 8).
Among mammal taxa, hunting activities are 
mainly focused on three species, no one of which 
is clearly dominant considering NISP . In addition, 
the spectrum of exploited animal resources is com-
pleted by other mammals, pond turtle (Delfino, 
Bressi 2000) and possibly birds (the analysis is still 
in progress), whilst consumption of land snails 
is difficult to evaluate (Girod 2003; Bonizzoni et 
al. 2009). As seen in other sites, the diet was also 
integrated with water molluscs and fish. In par-
ticular, a shift from the exploitation of fresh-water 
resources, to marine ones was observed through 
the Mesolithic (Cannarella, Cremonesi 1967) and 
can be linked to marine transgression. The fact 
that hunting activities are not focused on a single 
taxon, together with the evidence of transport to 
the site of complete carcasses and of their intensive 
exploitation, as well as the killing season mainly 
covering spring to early autumn months, make 
the Sauveterrian bone assemblage from Grotta 
dell’Edera very similar to those from Croatian 
sites (Pilaar-birch, Miracle 2017). At this stage of 
research, more data, especially those coming from 
the study of lithic implements (raw materials, lithic 
technology, use-wear) are needed to thoroughly 
understand the role played by Grotta dell’Edera in 
Tab. 10: Grotta dell’Edera. p-values of non-parametric 
Spearman’s correlation between MAU% and Bone density 
and MAU% and SFUI according to main taxa and layers.
T ab. 10: Stenašca. p-vrednosti za neparametrični Spearma-
nov test soodvisnosti med MAU % in strukturno gostoto 
kostnine ter med MAU % in SFUI za najpomembnejše 
taksone in plasti.
Tab. 11: Grotta dell’Edera. Sauveterrian layers: elements 
identified only at general anatomic level.
Tab. 11: Stenašca. Sovterjenske plasti, skeletni elementi so 
opredeljeni zgolj na ravni splošne anatomske pripadnosti.
Layer
MAU% vs
Bone density
MAU%
vs SFUI 
Ovis/Capra 2 0,09 0,72
Ovis/Capra 2a 0,57 0,8
Cervus elaphus 3b 0,14 0,14
Cervus elaphus 3c 0,0003 0,05
Capreolus 
capreolus
3b 0,06 0,38
Capreolus 
capreolus
3c 0,01 0,02
Sus scrofa 3b - 0,06
Sus scrofa 3c - 0,12
Layer
Skeletal elements 3b 3c 3d
Head elements 48 27 1
Isolated teeth 65 116 12
Vertebrae 53 45 1
Ribs 94 165 4
Sternum - 1 -
Innominate 6 7 -
Diaphysis 161 233 67
Basipodial 7 11 -
Metapodial 11 22 1
Phalanges 4 26 7
Sesamoidal bones 1 14 1

342 Francesco BOSCHIN
the subsistence strategies of Sauveterrian groups 
(hunting camp, shelter for the whole group, etc.).
Castelnovian
In Layer 3a, faunal composition radically 
changes from that of the underlying phase, due 
to the appearance of domestic ungulates (mainly 
caprines). In light of revisions, no cattle are found 
in this late Castelnovian level (it has been dated 
to 6390±60 BP, GrA-19820; Biagi 2003) and only 
one specimen can be ascribed quite confidently 
to a domestic pig. Domesticates account for 44% 
of NISP. These values are totally different from 
the slightly older Castelnovian spit of Viktorjev 
spodmol near Škocjan (Slovenian Karst), where 
domesticates, with the exception of a dog, are 
absent (Toškan, Dirjec 2004, Sipts 10–19 in Table 
16.7; NISP = 131). Radiometric date of Layer 3a 
of Grotta dell’Edera is similar to the one of the 
middle Neolithic level I at Pupićina peć in Istria 
(Forenbaher, Kaiser, Miracle 2004). In that case, 
the faunal composition is very different, showing 
a higher proportion of domesticates (considering 
cattle, pig and caprines, it ranges between 94.5% 
and 96.5%, depending on the quantity of domestic 
pig remains among the category “Sus”). Dates of 
Layer 3a are also similar to those of overlaying Layer 
2a, where caprines are the dominant species. The 
high number of fragments affected by weathering 
indicates a possible slow burial of remains and 
point to the formation of a short-term palimpsest, 
whose components (Neolithic and Castelnovian) 
would be difficult to tell apart. Archaeological 
and chronological data from the region seem to 
indicate possible interrelationships between the 
last hunter-gatherers and the first farmers (Biagi 
2003), and re-analysis of faunal data from Layer 
3a cannot confirm or reject the hypothesis that 
the Castelnovian hearth would attest a contact 
between the two groups. Regardless of whether 
the presence of domesticates in the Castelnovian 
phase due to a palimpsest or to the effective contact 
between Mesolithic hunter-gatherers and Neolithic 
groups, the direct date of an Ovis bone (7566–7470 
cal BP, GrA-19912; Biagi 2003) points to a quite 
old presence of domesticates in the area of the 
Trieste bay. Indeed, it is contemporaneous with 
the oldest Neolithic dates obtained at Pupićina 
Cave (7463–7326 cal BP, OxA-8471; 7621–7465 
cal BP, Beta-131625; Miracle, Forenbaher 2006), 
thus indicating that the spread of Neolithic (Da-
nilo) influences to the Karst plateau did not take 
place after the arrival of this culture in Istria but 
simultaneously.
Neolithic-Eneolithic
Layer 2a (Vlaška group) is characterized by an 
overwhelming presence of domesticates (Tab. 1), 
among which caprines are dominant. The same 
situation was recognized in Layer 6 at Grotta 
del Mitreo/Mitrejeva jama (Petrucci 1997), and 
in Levels G, H, and I at Pupićina peć (Miracle 
2006); wild game is more represented in Layer 5 
of Grotta degli Zingari/Ciganska jama (Bon 1996) 
and in Layer 8 of Grotta dei Ciclami/Orehova pejca 
(Riedel 1969) but some doubts have been cast on 
the stratigraphic reliability of this site (Radmilli 
1981). At Grotta dell’Edera, sheep is more abundant 
than goat (Tab. 1) like at Pupićina peć and Grotta 
del Mitreo/Mitrejeva jama. The high proportion 
of goat horn cores at Grotta dei Ciclami/Orehova 
pejca (Riedel 1969) can be due to the very selective 
recovering of bones at the site.
The percentage of wild animals NISP varies 
between 0 and 43% (Me = 13.5%) throughout 
the Neolithic-Eneolithic sequence unearthed at 
Podmol pri Kastelcu in the Slovenian Karst (Turk 
et al. 1993, Tab. 6). At this site, caprines are the 
most abundant mammals among domesticates, 
while the most important wild game is red deer. 
During phase 1 (Neolithic-Eneolithic) at the nearby 
Acijev spodmol (Slovenian Karst, Turk et al. 1992, 
Tab. 2), domesticates represented between 68% up 
to 82% of the total NISP . Again, caprines account 
for the great majority of this finds.
In any case, when faunal composition is discussed, 
some important factors have to be considered: 
the small size of some samples, the uncertain 
stratigraphic interpretations of some excavations 
(e.g., Grotta dei Ciclami/Orehova pejca), and the 
possible incomplete recovering of faunal remains 
in others (at Grotta degli Zingari/Ciganska jama 
there are 239 identified remains out of 261, whilst 
at Grotta del Mitreo/Mitrejeva jama 224 out of 
237). In some caves, the high proportion of badger 
remains could suggest a possible disturbance 
caused by the digging activity of this animal. At 
Grotta degli Zingari/Ciganska jama, for instance, 
this species is the most abundant (NISP = 66), and 
the presence of a badgers’ sett was hypothesized 
(Bon 1996).

343 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
Regarding anthropogenic stratigraphic dis-
turbances, the possibly reworked area detected 
in Layer 2a of Grotta dell’Edera is paradigmatic: 
the apparent contribute of hunting to Neolithic 
subsistence economy, testified by the presence of 
wild game remains (published in Boschin, Riedel 
2000), can be mainly due to the reworking of cave 
sediments by Neolithic shepherds. The latter burned 
coprogenic materials, mainly composed of caprine 
dung, organizing it in small heaps (Boschian 1997). 
An ab antiquo reworking due to Neolithic people 
is testified to at Grotta Azzurra/Pečina na Lesko-
vcu also by the occurrence of Mesolithic artefacts 
together with the dung of domesticated animals 
(Boschian, Montagnari-Kokelj 2000). Reworking of 
the deposit was also detected at Mala Triglavca in 
the Slovenian Karst (Mlekuž et al. 2008). At Grotta 
dell’Edera, 
14
C dates testify for other reworkings 
of cave soil by shepherds (Spataro 1999) in the 
later Neolithic/Eneolithic Layer 2.
The overwhelming presence of caprines in 
Layer 2a was probably favoured by the rocky 
environment and by the vegetation of the Karst: 
Selserio-Quercetum petraeae is considered as the 
climax vegetation of the area (Poldini 1980), and 
its undergrowth is mainly made of Poaceae. Even 
if secondary successions could have characterized 
vegetation dynamics of the area in Neolithic times 
(Turk et al. 1993; Kaligarič, Culiberg, Kramberger 
2006), anthropogenic modification of vegetation 
was not necessary for Neolithic shepherds, who 
probably took advantage of these particular woods 
that offer optimal food for grazers.
In Layer 2, caprines are still the most abundant 
domesticates at Grotta dell’Edera, but cattle and pig 
remains also increase, especially in the uppermost 
spits of the layer. A decrease in caprines can also 
be observed in the small samples from the younger 
Layers 1b and 1a. The same trend is observed at 
Pupićina peć from Layers I and H (Vlaška group) 
to Layer G (Eneolithic). At Grotta del Mitreo/
Mitrejeva jama, in the small sample from Layer 
6, there are no pig and cattle specimens (Vlaška 
group, NISP = 34), while these taxa became more 
abundant in the overlaying Layers 5 to 3 (Late 
Neolithic-Eneolithic) (Petrucci 1997). Layer 2 of 
Grotta dell’Edera is also characterized by a slight 
increase in wild game (from 0.4% of NISP in Layer 
2a to 3.7% in Layer 2). This could indicate a change 
towards more diversified subsistence strategies.
Given that Payne’s age classes A, B and C (0–2 
months, 2–6 months and 6–12 months respec-
tively) are all represented at the site, slaughtering 
(or natural death) of caprines is compatible both 
with a continuous presence at the site across all 
seasons (if one litter a year is hypothesized) or 
with a seasonal frequentation, if two litters are 
considered.
The high percentage of possible caprine foetal 
individuals (5 out of 23, 21.7% of MNI in Layer 
2a; 5 out of 15, 33.3% of MNI in Layer 2) could 
be indicative of a high mortality, thus testifying 
to livestock whose survival was still strongly de-
pendent on natural constrains (pathogenic agents, 
stress events during gestation). A more conserva-
tive interpretation is that this abundance could be 
related to the nature of the site (i.e., a sheepfold) 
and its use. Indeed, a site frequentation during 
ewes’ pregnancy and birth season can favour a 
selective accumulation of bones belonging to foetal 
individuals due to abortion (Boschin 2020). One 
foetal individual was also detected both in Layers 
2 and 2a among cattle remains. Even though age 
estimation of foetuses is based on modern refer-
ence data and some of these individuals could 
be new-borns, caprine foetuses appear to cover 
a variety of prenatal ages, not only placed at the 
end of gestation; they can also testify the death 
of pregnant individuals (Fig. 10). This fact also 
points to a significant problem in evaluating the 
presence of young individuals belonging to Payne’ s 
A and B age classes (0–2 months and 2–6 months 
respectively) (Payne 1973). Were they killed by 
humans following peculiar production strategies, 
or did they die due to adverse conditions? Indeed, 
it was observed that the causes of miscarriage 
(e.g., infections, nutritional stress) can also lead 
to the natural death of lambs (Boschin 2020 and 
references therein).
Both in Layer 2a and 2, only a few caprine indi-
viduals are represented. Even if data are sparse, it 
was demonstrated that caprine mortality profiles 
from Grotta dell’Edera resemble the theoretical 
model (Payne 1973) considered as indicative of the 
exploitation of milk and dairy products (Boschin 
2020). Nonetheless, the same study pointed to the 
fact that if the presence of foetuses and very young 
lambs can be related to natural mortality, kill-off 
data should be considered with care in the analysis 
of a site with a specific function (i.e., a sheepfold). 
The exploitation of milk can be better detected 
by the analysis of chemical residues on vessels. 
In this perspective, interesting results, indicating 
the presence of dairy products at Mala Triglavca 
on the Karst plateau already at 7565–7474 cal BP 
(Poz-48540), have been achieved by Budja et al. 

344 Francesco BOSCHIN
(2013). Even if dropped deciduous teeth confirm the 
use of the cave as a stable for animals, butchering 
marks testify to an occasional in situ consumption 
of domestic mammals, thus indicating that the 
cave was not only used as a sheepfold. Percussion 
marks and cones detected both in Layers 2 and 2a 
point to the need for shepherds to extract marrow, 
perhaps for feeding purposes. The presence of 
dry-bone fractures on limb bone elements is not 
in contrast with the abovementioned picture and 
can be explained by a further post-depositional 
bone breakage due to ungulate trampling. The 
integration of diet with animal fat (and also other 
animal resources, see Budja et al. 2013) would not 
be surprising, if we consider that few (or perhaps 
no) evidence of agricultural activities is available 
for the Karst in this period, even if it has been 
demonstrated that plants were surely part of the 
diet of Neolithic groups of the area (Budja et al. 
2013). While Neolithic settlements might have 
been submerged by the rise of sea level during 
the Holocene (Benjamin, Bonsall 2009), recent 
revision of materials from old excavations and 
the analysis of new finds (Bernardini, Betic 2008; 
Bernardini et al. 2017) point to the presence of 
possible open-air sites (San Rocco and Zaule) in 
the relatively fertile territory corresponding to 
the present-day coastal turbiditic hilly belt and 
plains in the south-western part of the province 
of Trieste. Unfortunately, no evidence is available 
so far on the agricultural vocation of these sites 
and on their possible relationships with the cave 
sites on the Karst plateau. Similarly, even if the 
presence of Danilo potsherds in open-air sites 
of the Friuli Plain, such as Sammardenchia and 
Fagnigola, points to the spread of the Danilo cul-
ture westwards, there are still no clear data on the 
relationships occurred between the settlements of 
the plain and the cave sites on the Karst plateau.
CONCLUSION
Faunal remains from Grotta dell’Edera represent 
the most important early Holocene bone assemblage 
from the Trieste Karst. The results show a first at-
tempt of exploitation of this territory by Sauveter-
rian hunter-gatherers about 11,000 years ago (Layer 
3d) in a period in which the plateau was probably 
still degraded, arid, and poorly vegetated. A more 
stable frequentation took place during the Boreal 
period/Early Atlantic at about 9300–8800 years 
ago. Hunting activities seem to follow a seasonal 
pattern and affect a broad spectrum of resources, 
whilst carcasses were intensively exploited. These 
data make it possible to compare Grotta dell’Edera 
with Mesolithic sites of Croatia (Pilaar-Birch and 
Miracle 2017).
The neolithization process and the arrival of 
domesticates were placed by Biagi, Starnini, V oytek 
(2008) in the first half of the 8
th
 millennium BP 
and the first livestock was mainly composed by 
caprines, probably more adapted to the pastures 
offered by the Karst plateau. The territory was 
highly suitable for Neolithic shepherds, due to 
the presence of caves that were used to stable 
flocks. Cattle and pig became more important in 
later times, but they were always subordinated to 
caprines.
Even if the Trieste Karst is characterized by a 
high concentration of cave sites that yielded Meso-
lithic and Neolithic remains, there are no extensive 
excavations, and some faunal assemblages were 
not studied from a taphonomic point of view and 
sometimes not even from a zooarchaeological one. 
As a result, available raw data about the exploita-
tion of faunal resources by Mesolithic, Neolithic 
and Eneolithic human groups are very scarce, and 
research is far from the possibility of drawing a 
synthesis about hunting/herding strategies, and 
human mobility across the territory. As an example, 
the faunal assemblage from Pupićina peć should 
be considered the only “representative enough” 
Neolithic sample for the whole Karst and Istria. 
Data from Grotta dell’Edera can be improved in 
future research by means of isotopic studies and 
more in-depth age estimations of individuals and 
seasonality data (e.g., by means of thin slices), but 
more excavations would be necessary to better 
contextualize any new result.
Acknowledgements
The author is grateful to Prof. Paolo Biagi for the pos-
sibility to study mammal remains from Grotta dell’Edera. 
This work is dedicated to the memory of Alfredo Riedel.
Illustrations: Fig. 3 (drawing: Paolo Biagi) / Slikovno gradivo: Sl. 3 (risba: Paolo Biagi)

345 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
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najstarejše sledi prisotnosti anatomsko moder-
nega človeka datirajo v zgodnji holocen. Ob tem 
je treba navesti, da je v večini jamskih najdišč 
s Krasa nepopolno zastopano tudi mezolitsko-
neolitsko sosledje plasti (Biagi 2003). Prav v tem 
je Stenašca v lokalnem kontekstu edinstvena, saj 
so tu mezolitski (sovterjenski in kastelnovjenski), 
neolitski in eneolitski sedimenti (sl. 3). Ker gre za 
edino najdišče s Krasa z dobro datiranim preho-
dom med mezolitikom in neolitikom (sl. 4), ga je 
treba razumeti kot eno najpomembnejših jamskih 
najdišč v tržaški pokrajini.
Stenašca leži na severozahodnem delu Tržaškega 
Krasa, slabe 3 km od današnje morske obale. Njen 
10 m širok vhod se odpira proti severovzhodu na 
dnu manjšega brezna na nadmorski višini pribl. 
230 m (sl. 1 in 2). Prva izkopavanja je v jami orga-
niziral G. Marzolini (1970), leta 1990 sta terensko 
raziskovanje nadaljevali univerza Ca’ Foscari iz 
Benetk in Kalifornijska univerza iz Berkeleyja 
(ZDA) pod vodstvom arheologov P . Biagi in B. A. 
Voytek (Biagi, Starnini, Voytek 1993). V tem času 
je bilo izkopanih pribl. 4 m sedimenta, pri čemer 
so bile prepoznane naslednje plasti (sosledje je 
podano od spodaj navzgor; sl. 3 in 4).
− Plast 3d: gre za plast s sledmi iz časa prvih 
človekovih obiskovanj jame, tj. iz predboreala 
(9810±70 uncal., GrN-23130; 9930±50 uncal., 
GrA-14108). Zbrano arheološko gradivo sodi v 
mezolitk (sovterjen).
− Plast 3c: vključuje ostanke hodne površine z in 
situ odkritimi živalskimi in kamnitimi najdbami ter 
lečami pepela/oglja. Datirana je v boreal (8350±120 
uncal., GrN-25139; 8250±50 uncal., GrA-11818), 
kar se časovno ujema z drugo sovterjensko fazo 
človekovih obiskovanj.
− Plast 3b: nastala je v obdobju prehoda med 
borealom in atlantikom (razpon datacij: od 8045±40 
uncal. [GrA-14106] do 8110±90 uncal. [GrN-
25138]). Vključuje mezolitsko (sovterjensko) 
arheološko gradivo.
−Plast 3a: gre za ognjišče premera pribl. 1 m, 
kjer je bilo odkrito kamnito gradivo iz mlajšega 
mezolitika (kastelnovjen) in 17 odlomkov lončenine. 
Kot je pokazala petrografska analiza, naj bi se 
minerološka sestava slednjih razlikovala od sestave 
lončenine iz mlajše plasti 2a, kar priča o njihovem 
alohtonem izvoru (Spataro 2001). Starost drobca 
oglja iz te plasti je bila z metodo radioaktivnega 
Ostanki velikih sesalcev holocenske starosti
iz jame Stenašca/Grotta dell’Edera na Tržaškem Krasu
(izkopavanja v letih 1990–2001) 
Povzetek

349 Ostanki velikih sesalcev holocenske starosti iz jame Stenašca/Grotta dell'Edera na Tržaškem Krasu ...
ogljika ocenjena na 6700±130 uncal. (GX-19569), 
starost odlomka ovčje kosti pa na 6620±60 uncal. 
(GrA-19912).
−Plast 3: gre za premešano plast debeline pribl. 
20 cm s posameznimi odlomki lončenine in nekaj 
kamnitimi odbitki lokalnega izvora. Do te plasti 
sedimente označuje rdečkasta obarvanost.
−Plast 2a: debela plast, palimpsest, ki ga sesta-
vlja navpično sosledje šestih leč pepela in oglja. 
Sediment večinoma vključuje sežgane iztrebke 
kopitarjev. Odlomki lončenine so bili pripisani vlaški 
kulturni skupini, razumljeni kot lokalna različica 
neolitske danilske kulture. Datumi, pridobljeni z 
metodo radioaktivnega ogljika, se razvrščajo med 
6305±285 uncal. (GX-19022) in 6615±390 uncal. 
(GX-19568) (Biagi, Starnini, Voytek 2008).
−Plast 2: palimpsest, katerega zgornji del (globina 
od 170 do 180 cm) datira v zgodnjo fazo eneolitika 
(4680±325 uncal., GX-19019). Drug razpoložljiv 
datum 
14
C, ki se nanaša na vzorec iz spodnjega dela 
plasti (globina od 200 do 210 cm), je bil ocenjen 
na 5555±35 uncal. (GrN-18968).
−Plast 1b: najverjetneje premešana; vključevala 
je nekaj zgodnjebronastodobnih ostankov.
−Plast 1a: vključevala je pičlo število arheolo-
ških najdb in ognjišče, datirano v pozno antiko, 
(1415±35 uncal., GrN-18968).
−Plast 1: vrhnja plast s pičlim številom arheo-
loških ostankov.
Taksonomsko in anatomsko opredeljevanje 
živalskih ostankov je potekalo ob sprotni kon-
zultaciji primerjalnih osteoloških zbirk. Nomen-
klatura domačih živali sledi smernicam, ki so jih 
objavili Gentry, Clutton-Brock, Groves (2004). V 
preliminarni arheozoološki objavi subfosilne favne 
iz plasti 2, 2a, 3 in 3a (Boschin, Riedel 2000) je 
količina najdb podana kot število ohranjenih delov 
posameznih kosti in ne kot dejansko število doku-
mentiranih kostnih drobcev. Tako je bil, denimo, 
nepoškodovan primerek stegnenice štet kar trikrat, 
tj. po enkrat za ohranjen proksimalni, medialni in 
distalni del kosti. Kvantitativni podatki iz omen-
jene objave zato ne zrcalijo dejanskega števila 
odkritih in analiziranih najdb, temveč posredno 
tudi stopnjo njihove fragmentiranosti. Zaradi 
enostavnejše primerjave rezultatov iz Stenašce 
z izsledki podobnih analiz z drugih najdišč je v 
tem članku količina živalskih ostankov podana 
kot število opredeljenih primerkov (Number of 
Identified Specimens; NISP). Taksonomsko neo-
predeljene najdbe so bile, kjer je bilo to mogoče, 
grupirane na podlagi anatomske pripadnosti (npr. 
rebra, vretenca, diafize dolgih kosti). Pri računanju 
najmanjšega števila elementov (Minimum Number 
of Elements; MNE) in najmanjšega števila oseb-
kov (Minimum Number of Individuals; MNI) so 
bili upoštevani anatomska stran (tj. levo-desno) 
in velikost posamezne najdbe, kakor tudi starost 
pripadajoče živali ob poginu (za razpravo o pomenu 
obeh navedenih indeksov glej Lyman 2008). Podatki 
o gostoti kostnine posameznih skeletnih elemen-
tov izbranih živalskih vrst so povzeti po Lymanu 
(Lyman 1984). Pri indeksih uporabnosti so bile 
upoštevane smernice, ki so jih objavili Metcalfe, 
Jones (1988), Rowley-Conwy, Halstead, Collins 
(2002) in Morin (2007). Metrični podatki so bili 
zbrani v skladu z navodili von den Driescheve (von 
den Driesch 1976) in so podani v prilogi. Beleženje 
stopnje obrabe žvekalne površine kočnikov pri 
živalih iz rodov Bos in Sus sledi smernicam Gran-
tove (Grant 1982), pri drobnici pa objavi Payna 
(Payne 1973). T aksonomska opredelitev kosti in zob 
ovčjih oziroma kozjih zarodkov je bila izvedena na 
podlagi kriterijev, ki so jih predstavili Habermehl 
(1975), Ahmed (2008), Richardson et al. (1976) 
in Mcdonald et al. (1977). Starost ob poginu pri 
jelenjadi in srnjadi je bila ocenjena po metodi, kot 
jo podajajo Tomé, Vigne (2003), Mariezkurrena 
(1983) in Habermehl (1985). Svežost posameznih 
kosti ob njihovem razbitju je bila ugotavljana z 
uporabo indeksa svežosti loma (Fracture Freshness 
Index oziroma FFI; Outram 2002). Spremembe 
na površini kosti so bile analizirane pod najmanj 
10-kratno povečavo in ob uporabi primerjalne 
zbirke modernega kostnega gradiva, ki je bilo 
nabrano na planem. Proučevanje sledi izpostavl-
jenosti vremenskim razmeram (tj. razpok, luščenja 
površine kosti) je zajelo zgolj nesežgane kostne 
odlomke velikosti nad 3 cm. Datumi, pridobljeni 
z metodo radioaktivnega ogljika, so bili kalibrirani 
z uporabo programskega paketa OxCal 4.3 (Bronk 
Ramsey 2009) in ob upoštevanju krivulje IntCal 13 
(Reimer et al. 2013). Statistična analiza in priprava 
grafikonov sta bili izvedeni s pomočjo programskih 
paketov PAST (Hammer, Harper, Ryan 2001) in R 
(R Development Core Team 2008).
Rezultati
Analiziranih je bilo približno 69.000 ostankov 
velikih sesalcev, vendar je bila stopnja določljivosti 
gradiva zaradi izrazite razdrobljenosti najdb 
skromna (tab. 1). Med ostanki iz sovterjenskih 
plasti močno prevladujejo kosti in zobje jelena, 

350 Francesco BOSCHIN
srne in divjega prašiča (njihov skupni delež pres-
ega 90 odstotkov vseh taksonomsko opredeljenih 
najdb). Vrstna sestava tega gradiva ne izkazuje 
diahronih sprememb.
Za skromen skupek kosti iz plasti 3a je značilna 
večinska zastopanost domačih živali, vendar je 
soliden tudi delež divjadi (predvsem jelen). Po-
dobno sliko kaže skromen nabor najdb iz (morda 
premešane) plasti 3.
Za plast 2a je treba na podlagi rezultatov revi-
zije (kvadranti 27–35 pri tem niso bili upoštevani 
zaradi morebitne prisotnosti divjadi iz mezolitskih 
kontekstov; sl. 5) ugotoviti očitno prevlado drobnice 
(96,9 % NISP), prisotnost nekaj posameznih ostan-
kov goveda in domačega prašiča ter maloštevilne 
najdbe divjadi (0,4 % NISP). Zelo podobno sliko 
kaže plast 2. Najbolje zastopan takson je tudi 
v tem primeru drobnica (79,9 % NISP), deleži 
goveda (4,3 % NISP), domačega prašiča (8,7 % 
NISP) in divjadi (3,7 % NISP) so komaj kaj večji. 
Zanimivo je, da je od spodnjega proti zgornjemu 
delu omenjene plasti zaznati postopno rast števila 
govejih in prašičjih najdb (sl. 6). Vendar pa je ob 
tem treba opozoriti, da je na podlagi razpoložljivih 
geoarheoloških podatkov obravnavana plast dom-
nevno premešana, saj naj bi prazgodovinski pastirji 
prerazporejali v jami akumulirane živalske iztrebke 
(Boschian 1997). Število taksonomsko opredeljenih 
živalskih ostankov v plasteh 1a in 1b je pičlo (tj. 67 
oziroma 32 primerkov). Najbolje zastopan takson 
ostaja drobnica, delež ostankov prašiča presega 
delež ostankov goveda, najdb divjadi pa ni.
Vzorec fragmentiranosti gradiva se med posa-
meznimi plastmi bistveno ne razlikuje, saj so 
vzdolž celotnega stratigrafskega stolpca znatno 
zastopani zelo majhni kostni drobci (sl. 7A). Indeks 
svežosti loma (FFI; Outram 2002) je bil izračunan 
za odlomke iz vseh plasti z izjemo plasti 1, 1a in 
1b. Rezultati so prikazani na sliki 7B.
Sledi izpostavljenosti vremenskim razmeram 
(razpokanost, luščenje površine) so bile opažene na 
številnih najdbah iz plasti 2 do 3c (tab. 2). Zanimivo 
je, da v okviru sedimentov mezolitske starosti od 
plasti 3c navzdol številčnost takšnih najdb upade, 
domnevno zaradi hitrejše sedimentacije. Podobno 
so kosti iz plasti 2a bolje ohranjene od tistih iz 
plasti 2, ki naj bi se odlagala počasneje (sl. 4).
Sežgani ostanki so bili odkriti vzdolž celotnega 
stratigrafskega stolpca, vendar so najbolje zasto-
pani v plasteh 2 in 3d (tab. 4). Zasekanine in vrezi 
so bili odkriti predvsem na kosteh kopitarjev, v 
posameznih primerih pa tudi na ostankih zajcev, 
svizcev in zveri (tab. 5).
V eč deset analiziranih živalskih najdb je obgrize-
nih, tako od zveri kot glodavcev. Pretežni del 
takšnih kosti izvira iz plasti 3b (38 primerkov) in 
3c (53 primerkov). Posebno omembo zasluži prst-
nica srne iz plasti 3b, ki je obgrizena in sežgana, 
ter dve nartni kosti iste vrste iz plasti 3c, izmed 
katerih je ena (skočnica) obgrizena, na drugi 
(kvadratnočolničasta kost) pa so bili opaženi vrezi.
Podatki o zastopanosti posameznih skeletnih 
elementov so podani v tabelah 6 do 8, vendar so 
podrobneje obravnavani le v primeru plasti 2 in 
2a (drobnica) oziroma 3b in 3c (jelen, srna, divji 
prašič) (tab. 9). Zgolj v navedenih primerih je 
bilo namreč mogoče oblikovati zadovoljivo velike 
in torej kolikor toliko reprezentativne vzorce. Z 
izjemo plasti 3c soodvisnost med številčnostjo 
kosti iz posameznih anatomskih regij (MAU 
%) in strukturno gostoto teh kosti ni statistično 
značilna. Enako praviloma velja za korelacijo med 
zastopanostjo posameznih anatomskih regij (MAU 
%) in standardiziranim indeksom prehranske upo-
rabnosti (SFUI) pripadajočih kosti (tab. 10). Med 
taksonomsko opredeljenimi ostanki so najbolje 
zastopani skeletni elementi glave, in to v vsaki 
izmed analiziranih plasti. Ugotovitev kaže pripisati 
enostavnejšemu opredeljevanju zob v primerjavi 
s kostnimi drobci. Med taksonomsko neopredel-
jenimi ostanki sovterjenske starosti prevladujejo 
odlomki diafiz dolgih kosti, reber, vretenc in zob.
Sklep
Živalski ostanki iz Stenašce so najpomembnejši 
arheozoološki zbir zgodnjeholocenske starosti z 
območja Tržaškega Krasa. Prve sledi prisotnosti 
sovterjenskih lovcev in nabiralcev v tej jami so 
datirane v čas pred približno 11.000 leti (plast 3d), 
ko je bila planota najbrž še degradirana, sušna in 
slabo poraščena. Intenzivnejše obiskovanje jame 
se je domnevno začelo v borealu in na začetku 
atlantika, v obdobju pred približno 9.300 do 8.800 
leti. Lov naj bi se tedaj večinoma izvajal sezonsko 
(sl. 8), nabor plenjenih živali pa je bil razmeroma 
širok (veliko je predvsem ostankov jelena, srne in 
divjega prašiča). V tem smislu kaže Stenašca številne 
podobnosti s sočasnimi najdišči na Hrvaškem 
(Pilaar-Birch, Miracle 2017).
V plasti 3a se vrstna sestava zbira živalskih 
ostankov v primerjavi z ugotovitvami iz spodaj 
ležečih sedimentov bistveno spremeni, saj se 
pojavijo najdbe domačih kopitarjev (44 % NISP; 
prevladujejo ostanki drobnice). Pri tem je revizija 

351 Ostanki velikih sesalcev holocenske starosti iz jame Stenašca/Grotta dell'Edera na Tržaškem Krasu ...
gradiva iz te plasti, datirane v obdobje mlajšega 
kastelnovjena (6390±60 BP, GrA-19820, Biagi 
2003), pokazala na odsotnost kosti ali zob goveda. 
Datacija plasti 3a je podobna dataciji plasti 2a, ki 
leži neposredno nad njo in kjer prav tako prevla-
dujejo ostanki drobnice. Pogostnost najdb s sledmi 
izpostavljenosti vremenskim razmeram bi lahko 
kazala na počasno sedimentacijo in oblikovanje 
kratkotrajnega palimpsesta s kastelnovjenskimi 
in neolitskimi elementi. Arheološki in kronološki 
podatki iz regije pričajo o verjetni interakciji med 
zadnjimi lovci-nabiralci in prvimi kmetovalci (Biagi 
et al. 2003). Tukaj predstavljeni rezultati analize 
živalskih ostankov iz plasti 3a teze o tem, da bi 
tam odkrito kastelnovjensko ognjišče utegnilo 
dokazovati stik med navedenima skupinama, niti 
ne potrjujejo niti ne zavračajo.
Biagi, Starnini in V oytek (2008) postavljajo začetek 
procesa neolitizacije Krasa in pojav domestikatov v 
prvo polovico 8. tisočletja pred sedanjostjo. V eliko 
naj bi bilo predvsem drobnice, ki je bila domnevno 
bolj prilagojena na pašo v tedanji lokalni krajini. 
Kras je bil neolitskim živinorejcem naklonjen tudi 
zaradi možnosti uporabe številnih jam za staje. 
Pomen goveda in domačega prašiča se je sčasoma 
nekoliko okrepil, vendar ni nikoli presegel vloge 
ovce in koze.
Podatki o starosti drobnice ob poginu/zakolu 
dovoljujejo razmišljanja o tem, da so bile črede v 
jami vse leto. To naj bi veljalo, če je bila kotitev 
na leto le ena. V gradivu so bili namreč zastopani 
vsi trije najzgodnejši starostni razredi po Paynu: A 
(0–2 meseca), B (2–6 mesecev) in C (6–12 mese-
cev). Če sta bili kotitvi dve, bi te iste podatke o 
starosti ovci in koz ob smrti lahko razumeli kot 
indic za predvsem sezonsko prisotnost čred v 
obravnavani jami.
Pomenljiv je podatek o bržčas znatni zastopanosti 
ostankov zarodkov ovc in/ali koz v plasteh 2 in 2a 
(tab. 3), saj bi to lahko podkrepilo tezo o uporabi 
jame za stajo. Če je bila Stenašca obiskovana (tudi) 
v času neposredno pred kotitvijo in med njo, je 
prisotnost kosti zarodkov kot posledica splavov 
pravzaprav pričakovana (Boschin 2019). Žal je 
bila metodologija za oceno starosti zarodkov ob 
poginu oblikovana na podlagi študije sodobnih 
ovc in koz, zaradi česar bi utegnile nekatere od 
omenjenih kosti pripadati že skotenim živalim. 
V endar pa naj bi vsaj nekateri primerki iz Stenašce 
poginili precej pred kotitvijo, kar prepričljivo 
dokazuje pogin brejih samic (sl. 10). Odprto ostaja 
še vprašanje o razlogih za smrt jagnjet in kozličev 
obeh najzgodnejših starostnih razredov po Paynu, 
tj. razredov A (0–2 meseca) in B (2–6 mesecev) 
(Payne 1973). So bili ti načrtno zaklani v skladu 
z izbrano politiko reje ali so poginili zaradi neu-
godnih razmer? Dejavniki, ki lahko sprožijo splav 
(npr. okužbe, prehranski stres), utegnejo namreč 
povzročiti tudi naravni pogin jagnjet/kozličev 
(Boschin 2019 in tam navedena literatura).
Smrtnostni profil za drobnico, ki sicer temelji na 
razmeroma pičlem zbiru podatkov, je v splošnem 
zelo podoben teoretičnim modelom vzreje teh 
živali s ciljem pridobivanja mleka in mlečnih 
izdelkov (Payne 1973; Boschin 2019). V endar pa je 
treba zaradi odkritja ostankov zarodkov in komaj 
skotenih mladičev, izmed katerih so vsaj nekateri 
utegnili poginiti naravne smrti, ocene starosti 
drobnice ob poginu/zakolu jemati zelo previdno. 
Toliko bolj zato, ker je bila Stenašca zelo verjetno 
periodično uporabljana kot staja za živino. Prido-
bivanje mleka je mogoče verodostojneje oceniti z 
analizo organskih ostankov na lončenini. V tem 
smislu kaže omeniti rezultate takšnih raziskav na 
gradivu iz še ene kraške jame s tega območja, tj. 
Male Triglavce pri Divači. Zanimivo je namreč, 
da rezultati dokazujejo človekovo uporabo mleka 
in mlečnih izdelkov že pred okvirno 7500 leti 
(7565–7474 cal BP; Poz-48540) (Budja et al. 2013).
Nedavna revizija že pred časom izkopanega 
arheološkega gradiva in analiza novih najdb (Ber-
nardini, Betic 2008; Bernardini et al. 2017) sta 
nakazali verjeten obstoj najdišč na prostem (San 
Rocco, Žavlje/Zaule) na razmeroma rodovitnem 
območju turbiditnega hribovja in nižin vzdolž 
današnjega obalnega pasu na jugozahodu tržaške 
pokrajine. Žal podatki o kmetijskih aktivnostih 
tedanjih prebivalcev teh naselbin niso znani, tako 
kot tudi ne podatki o njihovem odnosu do jamskih 
najdišč s Krasa. To velja kljub odkritju lončenine 
danilske kulture na nekaterih tedanjih najdiščih 
na prostem iz Furlanije (npr. Sammardecchia, 
Fagnigola), ki sicer kažejo na širitev te kulture tudi 
na območja zahodno od kraške planote.
Kljub velikemu številu jamskih najdišč z me-
zolitskim in neolitskim gradivom na območju 
T ržaškega Krasa ta nikoli niso bila obsežneje terensko 
raziskovana. Tudi živalski ostanki so bili pogosto 
analizirani zgolj parcialno ali pa še to ne. Zato na 
podlagi razpoložljivih arheozooloških podatkov 
za obdobje mezolitika, neolitika in eneolitika ni 
mogoče podati celovite slike o strategijah živinoreje 
in lova ter o vzorcih človekove mobilnosti na tem 
območju. Pravzaprav kaže kot edino zares reprezen-
tativno neolitsko najdišče za celoten prostor Krasa 
in Istre omeniti hrvaško Pupićino peć. Podatke iz 

352 Francesco BOSCHIN
Stenašce bi bilo mogoče v prihodnje obogatiti z 
analizami stabilnih izotopov, detajlnejšo študijo 
starosti živali ob poginu/zakolu, preučevanjem 
indicev o sezoni človekovih obiskov jame (npr. s 
histološkimi analizami kostnine) ter predvsem z 
nadaljnjim terenskim raziskovanjem.
Prevod: Borut Toškan
Francesco Boschin
Universita degli Studi di Siena
Dipartimento di Scienze Fisiche, della Terra e 
dell'Ambiente
Via Laterina 8
I-53100 Siena
fboschin@hotmail.com
https://orcid.org/0000-0001-5795-9050
App. 1: Grotta dell’Edera. Measurements taken on animal bones. 
Meas: Measurement; M\: Upper molar tooth; M/: lower molar tooth; D\: Upper deciduous tooth; D/: lower deciduous 
tooth; Mc: Metacarpal; Mt: Metatarsal. (Measurement’s codes are from: Von den Driesch 1974).
Pril. 1: Stenašca. Metrični podatki za živalske ostanke. 
Meas: dimenzija; M\: zgornji kočnik; M/: spodnji kočnik; D\: zgornji mlečni zob; D/: spodnji mlečni zob; Mc: dlančnica; 
Mt: stopalnica. (Označbe posameznih dimenzij so povzete po: von den Driesch 1974).
Taxon Layer Element Meas. 1 Meas. 2 Meas. 3 Meas. 4
Bos sp. 2a Talus GLl 69 GLm 64,1 Dl 38,4 Bd 42,5
Bos taurus 1a M/3 L 36,5 B 14
Ovis aries 2a Scapula GLP 32,2
Ovis aries 2a Radius Bp 27,1 BFp 24,7
Ovis aries 2a Metacarpus Bd 24,8
Ovis aries 2a Metacarpus Bd 24,2
Ovis aries 2a Talus GLl 31,5 GLm 29,5 Dl 17,6 Bd 19,6
Ovis aries 2a Talus GLl 29 GLm 27,7 Dl 16,3 Bd 18,8
Ovis aries 2a Metatarsus Bd 22,8
Ovis aries 2a Metatarsus Bd 27
Ovis aries 2a Phalanx 1 GLpe 31,5 Bp 10,8 SD 8,6 Bd 9,8
Ovis aries 2a Phalanx 1 GLpe 38,8 Bp 13,7 SD 11,5 Bd 13,2
Ovis aries 2a Phalanx 1 GLpe 36,2 Bp 12,6
Ovis aries 2a Phalanx 1 GLpe 36,5 Bp 11,3
Ovis aries 3a Phalanx 1 GLpe 31,1 SD 9,2
Ovis aries 3a Phalanx 2 GL 18,5 Bp 11,5 SD 8,4 Bd 8,5
Capra hircus 2a Scapula SLC 19
Sus domesticus 1b M\3 L 29 B 18
Sus domesticus 2 M\3 L 34,5
Sus domesticus 2 M/2 L 21
Sus domesticus 2 M/3 L 35
Sus domesticus 2 Scapula SLC 18,2
Sus scrofa 3 M\3 L 38,8
Sus scrofa 3b M/1 L 19,4
Sus scrofa 3c D\4 L 18,5

353 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
Taxon Layer Element Meas. 1 Meas. 2 Meas. 3 Meas. 4
Sus scrofa 3c M\1 L 18,3
Sus scrofa 3c M\1 L 19,3
Sus scrofa 3b Radius Bp 32
Sus scrofa 3b Ulna DPA 56 SDO 40,6 LO 72,6
Sus scrofa 3b MC IV Bp 18,8
Sus scrofa 3b MC IV Bp 21,8
Sus scrofa 3b Talus GLl 47,1 GLm 42,5
Sus scrofa 3b Phalanx 1 BP 15,6
Sus scrofa 3b Phalanx 1 BP 23,6
Sus scrofa 3b Phalanx 1 BP 21
Sus scrofa 3b Phalanx 1 BP 22,1
Sus scrofa 3c Scapula GLP 44 LG 36,4 BG 28,6
Sus scrofa 3c Humerus Bd 54,4 BT 41,3
Sus scrofa 3c Humerus Bd 55,1 BT 40,4
Sus scrofa 3c Humerus Bd 44,3 BT 36,3
Sus scrofa 3c Radius Bp 32,2
Sus scrofa 3c Radius Bp 31,4
Sus scrofa 3c Radius Bd 41,5
Sus scrofa 3c Ulna DPA 37,5 SDO 27,5 BPC 23,4
Sus scrofa 3c Ulna BPC 22,5
Sus scrofa 3c Ulna BPC 20,7
Sus scrofa 3c MC IV Bp 23,4
Sus scrofa 3c Tibia Bd 36,3
Sus scrofa 3c Talus Gll 48,9
Sus scrofa 3c Talus Gll 49,6 GLm 45,5
Sus scrofa 3c Calcaneum GL 95,5
Sus scrofa 3c Mt III Bp 18,3
Sus scrofa 3c Mt III Bp 20
Sus scrofa 3c Phalanx 1 BP 18,6
Sus scrofa 3c Phalanx 1 BP 18,1
Sus scrofa 3c Phalanx 1 BP 18,1
Sus scrofa 3c Phalanx 1 BP 19,5
Sus scrofa 3c Phalanx 2 GL 26,1
Sus scrofa 3c Phalanx 2 GL 29 Bp 20,1 SD 15,8 Bd 18,1
Sus scrofa 3c Phalanx 2 GL 30 Bp 21,5 SD 17,1 Bd 21,5
Sus scrofa 3c Phalanx 2 GL 30,2 Bp 19,6 SD 16,4 Bd 15,8
Sus scrofa 3c Phalanx 2 GL 30,3 Bp 19,3 Bd 16,8
Sus scrofa 3d Humerus BT 43
Cervus elaphus 3a Radius Bd 59
Cervus elaphus 3a Ulna BPC 28,5
Cervus elaphus 3b M/3 L 36,6
Cervus elaphus 3b M/3 L 29,1
Cervus elaphus 3b Scapula GLP 67,1 LG 55,9 BG 49,5

354 Francesco BOSCHIN
Taxon Layer Element Meas. 1 Meas. 2 Meas. 3 Meas. 4
Cervus elaphus 3b Humerus Bd 58,4 BT 50,3
Cervus elaphus 3b Radius Bp 59,2 BFp 54,5
Cervus elaphus 3b Ulna DPA 49,2 SDO 48,8 LO 70,8 BPC 26,6
Cervus elaphus 3b Ulna DPA 48,9
Cervus elaphus 3b Metacarpus Bd 37,9
Cervus elaphus 3b Talus GLl 54,3 GLm 49,4 Dl 29 Bd 32,5
Cervus elaphus 3b Metatarsus Bd 42
Cervus elaphus 3b Metatarsus Bd 43,6
Cervus elaphus 3b Metatarsus Bd 42,2
Cervus elaphus 3b Phalanx 1 Bp 21,2
Cervus elaphus 3b Phalanx 1 Bp 19,8
Cervus elaphus 3b Phalanx 1 Bp 23,6
Cervus elaphus 3b Phalanx 1 Bp 21,8
Cervus elaphus 3b Phalanx 1 Bp 25
Cervus elaphus 3b Phalanx 1 Bp 21,8
Cervus elaphus 3b Phalanx 1 Bp 19,5
Cervus elaphus 3b Phalanx 2 Bp 18,8
Cervus elaphus 3b Phalanx 2 Bp 20,3
Cervus elaphus 3b Phalanx 2 Bp 19
Cervus elaphus 3c Maxilla 20 114,4 21 69,2 22 45,3
Cervus elaphus 3c Maxilla 22 46,4
Cervus elaphus 3c Mandible 7 125,1 8 79,4 9 45,8
Cervus elaphus 3c Mandible 7 49,3
Cervus elaphus 3c M/3 L 33,4
Cervus elaphus 3c M/3 L 33,9
Cervus elaphus 3c Radius Bp 49,4 BFp 46,7
Cervus elaphus 3c Ulna DPA 26,3 SDO 22,5 LO 30
Cervus elaphus 3c Ulna BPC 34,2
Cervus elaphus 3c Metacarpus Bp 44,6
Cervus elaphus 3c Metacarpus Bp 41,9
Cervus elaphus 3c Metacarpus Bd 40,7
Cervus elaphus 3c Metacarpus Bd 41,8
Cervus elaphus 3c Talus GLl 57,6 GLm 53 Dl 30,4 Bd 35,4
Cervus elaphus 3c Talus GLl 55,4 GLm 52,1 Dl 29,2 Bd 34,1
Cervus elaphus 3c Calcaneum GL 116,5
Cervus elaphus 3c Metatarsus Bd 42,1
Cervus elaphus 3c Phalanx 1 Bp 21,2
Cervus elaphus 3c Phalanx 1 Bp 23,1
Cervus elaphus 3c Phalanx 1 Bp 22,4
Cervus elaphus 3c Phalanx 1 Bp 20,2
Cervus elaphus 3c Phalanx 1 Bp 21,3
Cervus elaphus 3c Phalanx 1 Bp 21,6
Cervus elaphus 3c Phalanx 1 Bp 24

355 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
Taxon Layer Element Meas. 1 Meas. 2 Meas. 3 Meas. 4
Cervus elaphus 3c Phalanx 1 Bp 19,4
Cervus elaphus 3c Phalanx 1 Bp 20,5
Cervus elaphus 3c Phalanx 2 Bp 19,2
Cervus elaphus 3c Phalanx 2 Bp 20,4
Cervus elaphus 3c Phalanx 2 Bp 20,7
Cervus elaphus 3c Phalanx 2 Bp 20,6
Cervus elaphus 3c Phalanx 2 Bp 22,3
Cervus elaphus 3c Phalanx 2 Bp 23
Cervus elaphus 3c Phalanx 2 Bp 18,1
Cervus elaphus 3c Phalanx 2 Bp 22,2
Cervus elaphus 3c Phalanx 2 Bp 19,9
Cervus elaphus 3c Phalanx 2 Bp 17,8
Cervus elaphus 3d Mandible 8 75,6
Cervus elaphus 3d M/3 L 31,4
Cervus elaphus 3d Metacarpus Bd 39,4
Cervus elaphus 3d Talus GLl 56 GLm 52,4 Dl 29,3 Bd 35,1
Cervus elaphus 3d Phalanx 2 GL 38,7 Bp 19,6 Bd 17,3
Cervus elaphus 3d Phalanx 2 Bp 21,8
Capreolus capreolus 3b Maxilla 22 30,4
Capreolus capreolus 3b Maxilla 21 28,3
Capreolus capreolus 3b Scapula SLC 8,9
Capreolus capreolus 3b Humerus Bd 27,5 BT 25,3
Capreolus capreolus 3b Humerus Bd 27,2 BT 23,3
Capreolus capreolus 3b Ulna DPA 25,6 SDO 21,6 LO 31 BPC 15,9
Capreolus capreolus 3b Ulna DPA 24,1 SDO 21,7 LO 29,6 BPC 15,3
Capreolus capreolus 3b Tibia Bd 27,6
Capreolus capreolus 3b Talus GLl 29,7 GLm 29,4 Dl 17,1 Bd 20,5
Capreolus capreolus 3b Talus GLl 31,3 GLm 30,2 Dl 16,9 Bd 20
Capreolus capreolus 3b Talus GLl 31,2 Bd 20,4
Capreolus capreolus 3b Talus GLl 28,7 GLm 27,6 Dl 15,4 Bd 18,7
Capreolus capreolus 3b Talus GLl 31,7 Dl 16,9 Bd 19,9
Capreolus capreolus 3b Metatarsal Bd 25,1
Capreolus capreolus 3b Phalanx 1 Bp 12,3
Capreolus capreolus 3b Phalanx 1 Bp 10,6
Capreolus capreolus 3b Phalanx 2 Bp 12,3
Capreolus capreolus 3b Phalanx 2 Bp 11,8
Capreolus capreolus 3b Phalanx 2 Bp 10,6
Capreolus capreolus 3c Maxilla 22 28
Capreolus capreolus 3c Maxilla 21 35,4
Capreolus capreolus 3c M/3 L 15,6
Capreolus capreolus 3c M/3 L 15,8
Capreolus capreolus 3c Scapula SLC 7,9 BG 18,8 LG 20,7 GLP 27,6
Capreolus capreolus 3c Scapula BG 20,4 LG 22,8

356 Francesco BOSCHIN
Taxon Layer Element Meas. 1 Meas. 2 Meas. 3 Meas. 4
Capreolus capreolus 3c Humerus Bp 31,3
Capreolus capreolus 3c Humerus Bd 27
Capreolus capreolus 3c Radius Bp 54,1 BFp 50,8
Capreolus capreolus 3c Radius Bd 25
Capreolus capreolus 3c Radius Bd 26,8
Capreolus capreolus 3c Ulna DPA 26,6 SDO 22,6 LO 32
Capreolus capreolus 3c Ulna BPC 13,8
Capreolus capreolus 3c Metacarpus Bp 19,8
Capreolus capreolus 3c Metacarpus Bd 22,9
Capreolus capreolus 3c Metacarpus Bd 21,8
Capreolus capreolus 3c Tibia Bd 27,1
Capreolus capreolus 3c Talus GLl 29,5 GLm 28,8 Dl 15,9 Bd 19,3
Capreolus capreolus 3c Talus GLl 30,3 GLm 28,7 Dl 16,9 Bd 19,5
Capreolus capreolus 3c Talus GLl 32 GLm 29,1 Dl 18,2 Bd 19,1
Capreolus capreolus 3c Talus GLl 29,2 GLm 28,5 Dl 15,8 Bd 19,2
Capreolus capreolus 3c Talus GLl 29,9 GLm 28,7 Dl 16,5 Bd 17,6
Capreolus capreolus 3c Talus GLl 31 GLm 28,6 Dl 17,3 Bd 20,9
Capreolus capreolus 3c Talus GLl 31,4 GLm 30,2 Dl 16,7 Bd 19,7
Capreolus capreolus 3c Talus GLl 30,1 GLm 28,8 Dl 16,9 Bd 20,5
Capreolus capreolus 3c Talus GLl 30,4 GLm 28,7 Dl 17,5 Bd 19,3
Capreolus capreolus 3c Talus GLl 28 Dl 15,4
Capreolus capreolus 3c Calcaneum GL 64,3
Capreolus capreolus 3c Calcaneum GL 65,1
Capreolus capreolus 3c Calcaneum GL 64,5
Capreolus capreolus 3c Phalanx 1 Bp 12
Capreolus capreolus 3c Phalanx 1 Bp 11,8
Capreolus capreolus 3c Phalanx 1 Bp 11,6
Capreolus capreolus 3c Phalanx 1 Bp 12,9
Capreolus capreolus 3c Phalanx 1 Bp 11,6
Capreolus capreolus 3c Phalanx 1 Bp 10,4
Capreolus capreolus 3c Phalanx 1 Bp 11,5
Capreolus capreolus 3c Phalanx 1 Bp 12,3
Capreolus capreolus 3c Phalanx 1 Bp 12,2
Capreolus capreolus 3c Phalanx 1 Bp 10,2
Capreolus capreolus 3c Phalanx 1 Bp 13,4
Capreolus capreolus 3c Phalanx 1 Bp 11,8
Capreolus capreolus 3c Phalanx 1 Bp 13,2
Capreolus capreolus 3c Phalanx 1 Bp 11,7
Capreolus capreolus 3c Phalanx 2 Bp 9,6
Capreolus capreolus 3c Phalanx 2 Bp 12,2
Capreolus capreolus 3c Phalanx 2 Bp 10,6
Capreolus capreolus 3c Phalanx 2 Bp 10,7
Capreolus capreolus 3c Phalanx 2 Bp 11,8
Capreolus capreolus 3c Phalanx 2 Bp 10,8

357 Holocene macromammal remains from Grotta dell’Edera/Stenašca, Trieste Karst ...
Taxon Layer Element Meas. 1 Meas. 2 Meas. 3 Meas. 4
Capreolus capreolus 3c Phalanx 2 Bp 13,8
Capreolus capreolus 3c Phalanx 2 Bp 12,3
Capreolus capreolus 3d Phalanx 2 Bp 10,9
Capreolus capreolus 3d Phalanx 2 Bp 10,2
Capreolus capreolus 3d Phalanx 2 GL 26,6
Canis familiaris 2 Phalanx 2 GL 10 Bp 6,5 Bd 6
Canis familiaris 1b Talus GL 25,6
Canis familiaris 3b Mt IV GL 67,6 SD 5,2 Bd 8,5
Canis lupus 3c Scapula GLP 38,5 SLC 30,8 LG 29,8 BG 22
Canis lupus 3c Phalanx 1 GL 28,1 Bp 10,2 SD 6,6 Bd 8,3
Vulpes vulpes 3b Mc IV GL 44,2 Bd 5,2
Meles meles 3b Mc I GL 28,3
Meles meles 3b Mc 3 GL 29,9 Bd 6,6
Meles meles 3b Mt IV GL 32,1 Bd 6,2
Meles meles 3c Mc III GL 34,4 Bd 6,8
Meles meles 3c Mc III GL 31,7 Bd 6,2
Meles meles 3c Mc IV GL 29,8 Bd 7
Meles meles 3c Mt III GL 32,8 Bd 6,7
Meles meles 3c Mt III GL 31,8 Bd 6
Meles meles 3c Mt III GL 33,2 Bd 6,9
Meles meles 3c Mt IV Bd 6,8
Martes sp. 3b Humerus Bd 15
Martes sp. 3b Radius GL 52,2 Bp 6,2 SD 3,4 Bd 8,5
Martes sp. 3c Radius Bd 6,1
Lutra lutra 3b Humerus Bd 27,4
Lutra lutra 3b Talus GL 18,4
Lutra lutra 3c Ulna DPA 12,7 SDO 11,6 BPC 7,5
Lutra lutra 3c Ulna DPA 11,8 SDO 10,5
Lynx lynx 3c Mandibula 7 15,5