Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective

Abstract

The studies of the small mammal fossil and recent assemblages from the Limay river basin (Northwestern Patagonia) have strongly increased during the last decade. Taxonomic, taphonomic, and paleoecological information about small rodents and marsupials recovered from fossil sites offers the opportunity to discuss the periods of change and stability in the conformation of their communities through the Anthropocene. Here, we used two large data matrix of fossil and recent small mammal samples. As starting point, we considered the small mammal record of Epullán Grande cave (LL thereafter), which covers the Early Holocene/Post-hispanic Period, in order to assess the impact of anthropic activities on the small mammal communities during the Anthropocene. The taphonomic analysis performed on the newest samples from LL confirms the predatory activity of Tyto furcata on sigmodontines and human consumption on caviomorphs (mostly for the last ca. 1000 years). The analysis of manganese oxide staining suggested higher levels of moisture during the earliest formation of the LL sequence. The taxonomic results indicate a major diversity in the small mammal fossil assemblages to the later periods of LL and other fossil sequences of the Limay basin of the Anthropocene. Conversely, opportunistic sigmodontines (Abrothrix olivacea, Calomys musculinus, Eligmodontia spp. and Oligoryzomys longicaudatus) experienced a growth in the recent samples. Some stenoic and specialist species (Euneomys spp., Lestodelphys halli, Loxodontomys micropus and Reithrodon auritus) were abundant in the temporal units associated with the Anthropocene, but now are in retraction. Additionally, the drop in the diversity of recent assemblages supports a restructuration of small mammal communities from Limay river basin occurred in the 20th century.

Keywords

diversity Epullán Grande cave marsupials recent impoverishment rodents

Introduction

For the last four decades, especially during the last one, the studies of the small mammal Late Quaternary fossil and recent assemblages from the Limay river basin (Northwestern Patagonia, Argentina) have strongly increased (e.g. Andrade, 2015; Cordero, 2011; Crivelli Montero et al., 1996, 2017; Fernández et al., 2016, 2018; Guillermo et al., 2020a, 2020b, 2021; Pardiñas, 1999; Pardiñas and Teta, 2013; Pearson, 1987; Pearson and Pearson, 1993; Silveira and Cordero, 2023; Tammone et al., 2014, 2020; Teta et al., 2005). The pool of taxonomic, taphonomic and paleoecological information about small rodents and marsupials recovered mostly from archeological contexts of the Limay river basin offers the opportunity to discuss the occurrence of periods of change and stability in the conformation of their communities through the Holocene and post-hispanic times, taken into account the main climatic events and the anthropic impact.

Recent global climate and environmental changes promote, first informal and then formal, the definition of a new human-dominated geological epoch so-called the Anthropocene, which started for some authors at the beginning of the 17th century, according to the levels of the atmospheric CO₂ measured in high-resolution Antarctic ice core records (Lewis and Maslin, 2015, and references therein). The broad range and magnitude of human-induced changes during the Anthropocene Epoch embraces the land surface transformation and change in the composition of the atmosphere and oceans (Lewis and Maslin, 2015). The effect of modern humans (Homo sapiens) on the Earth has been sharply discussed because the paleoclimatic and paleoenvironmental changes happened regardless of human existence on Earth, independently of atmospheric CO₂ concentrations and the use of hydrocarbons (e.g. Foscolos, 2017; Singer and Idso, 2009).

On the other hand, high resolution stratigraphic event markers characterized a distinctive Anthropocene chronostratigraphic base, highlighting a great acceleration event array around mid-20th century (e.g. Waters et al., 2022; Waters and Turner, 2022). In fact, since mid-20th century the influence of the expansion in human population, together with their livestock, their cultivation and their industrial activities on the environments has increased markedly, affecting millions of species on Earth and probably triggered the sixth mass extinction in planet’s history (e.g. Barnosky et al., 2011; Ceballos et al., 2015, 2020; Waters et al., 2022; Waters and Turner, 2022). Conversely, Patagonian-Fueguian mammals showed few extinctions during the Anthropocene Epoch, including two species of canids: the foxes Dusicyon avus and Dusicyon australis (e.g. Prevosti et al., 2015, and references therein). Nonetheless, according to IUCN Red List, Lista Roja SAREM and the fossil record, several species of mammals –mostly small rodents and marsupials- have experienced drastic reductions in their range or abundance in the course of the Anthropocene Epoch (e.g. Andrade and Monjeau, 2016; Fernández, 2014; Fernández et al., 2021a; Pardiñas et al., 2000, 2012; Prevosti et al., 2015; Rebane, 2002; Tammone et al., 2020; Teta et al., 2014).

In this contribution, we have summarized the available taxonomic and taphonomic knowledge about the small mammals from the Anthropocene Epoch and recent pellet samples from Limay river basin (Northwestern Patagonia, Argentina) to better understand how climate and/or anthropic activities could have structured the abundance and distribution of the small mammal species and communities. As starting point, we considered the small mammal record of Epullán Grande cave (LL thereafter), an emblematic archeological site placed at the transition between the Patagonian steppe and the Monte desert (Figure 1). LL has a rich zooarchaeological sequence, spanning the Early Holocene (Period A), mid-Holocene (periods B and C), Late-Holocene (periods D and E) and /Post-hispanic (Period F). Here we studied the newest small mammal samples from all periods considering taxonomical, taphonomical, and paleoecological aspects. In order to assess the impact of human intensive activities on the small mammal communities, we will take special emphasis on the Post-hispanic Period / Anthropocene Epoch.

Figure 1.

Map of the study area, Limay river basin, northwest Patagonia, Argentina, superimposed on the main phytogeographic districts (cf. Oyarzabal et al., 2018), including the location of the fossil sites (black triangles) and recent pellet localities (red dots) used in this work (a). Cañadón del Tordillo area showing Epullán Grande (white triangle) and Epullán Chica (black triangle) sites (b). picture of Epullán Grande cave entrance taken by Luis Cesar Teira Mayolini (c). The abbreviations of the fossil sites and recent pellet localities are indicated in Tables S1 and S3.

Regional settings and archeological contexts

Environments at Limay river basin and Epullán Grande

The Limay river rises at the eastern end of the glacial lake Nahuel Huapi and flows about 500 km in northeastern direction, adding important tributaries such as the Traful, Pichileufú, Collón Curá and Picún Leufú rivers, until it meets the Neuquén River, forming at its confluence the Negro River. The climate and environment of the area shows significant west-east gradient from 3000 mm in the deciduous forest to 200–150 mm to the east in the Patagonian shrub-grass steppe and the Monte shrub-steppe (Figure 1a) due to the Pacific anticyclone and the topographic barrier of the Andean Cordillera, which generates drier conditions to the east (Paruelo et al., 1998; Prieto et al., 2021).

On the one hand, different paleoclimatic proxies from Limay River basin display similar climatic conditions than today and a general stability through the Late-Holocene, with some short episodes of colder/wetter or warmer/drier conditions possibly associated to the strengthening of interannual/interdecadal climate variability of the El Niño-Southern Oscillation –ENSO- (e.g. Bianchi, 2007; Heusser, 1993; Iglesias et al., 2014, 2017; Markgraf, 1983; Mercer, 1976; Prieto et al., 2021; Prieto and Stutz, 1996; Rabassa, 2008; Whitlock et al., 2006).

During the last four centuries the northwestern portion of Patagonia has been strongly affected by human activities. However, the most negative impact over the environment took place during the 20th century, when the human impact became more intensive trough urban growth, introduction of exotic plants (mostly Rosa rubiginosa and Pseudotsuga menziesii) and animals (e.g. Sus scrofa scrofa, Cervus elaphus, Neovison vison, Mus musculus, Rattus spp.), cattle raising (mostly Ovis orientalis aries and to a lesser extent Capra aegagrus hircus, Equus ferus caballus, Bos primigenius taurus and Sus scrofa domestica), intentional fires, wood industry, extraction of hydrocarbons and minerals, and hydric control by installing dams across the Limay river basin (e.g. Aagesen, 2000; Bertiller and Bisigato, 1998; Fernández and de Mendoza, in press; Tammone et al., 2020; Teta et al., 2014; Valenzuela et al., 2023; Victoria Lantschner et al., 2011).

LL (40°23′’21″S, 70° 11′ 40″W, 680 m asl) is an archeological cave site eroded in a rocky outcrop of volcanic tuffs of the Collón Curá Formation and is emplaced in the middle Limay river basin, specifically in the Cañadón del Tordillo from ~5 km north of the Limay River (Neuquén Province, Argentina; Figure 1b). LL is located at the Monte-Patagonian Ecotone and near the Occidental District of the Patagonian Phytogeographic Province at the middle part of the Limay River basin (Oyarzabal et al., 2018). Its plant composition embraces Patagonian shrubs, herbs, and grasses such as Chuquiraga, Cortaderia, Distichlis, Mulinum, Poa, Senecio and Stipa, and Monte shrubs such as Larrea, Prosopis and Schinus (Oyarzabal et al., 2018).

Chronostratigraphic context of Epullán Grande cave

LL cave (Figure 1c) is about 7 × 5 m. However, it cannot be discarded that the cave was probably larger due to the collapse of the boulder rock fall toward the opening during the 19th–20th centuries (Crivelli Montero et al., 1996; Prieto et al., 2021). The excavations were carried out between 1988 and 1992, under the direction of Dr. Eduardo Crivelli Montero. The entire surface of the cave was excavated (~50 m²) following the methodology proposed by Harris (1989), where each stratum was described, given a unique number in a continuous sequence beginning with 1, and its stratigraphic position recorded by noting its physical relationship with the adjacent strata. Only in a few occasions the surface of the cave was also excavated in natural layers and artificial levels of 5–10 cm; the stratigraphic thickness reached ~100–120 cm depth, depending on the place of the cave (Crivelli Montero et al., 1996).

The cave was periodically used as a shelter by humans and predators from ca. 11,500 cal yr BP to the 20th century (Crivelli Montero et al., 1996). Based on archeological and chronostratigraphical criteria six main periods have been suggested: Period A (11,444-8670 cal yr BP), Period B (7847 cal yr BP), Period C (5829 cal yr BP), Period D (2991-2319 cal yr BP), Period E (942-364 cal yr BP) and Period F (Post-hispanic). Radiocarbon calibrations (each one expressed in probabilistic median) were made using OxCal 4.4 program in conjunction with SHCal20 Southern Hemisphere curve (Hogg et al., 2020). Period A, ranges between bedrock (~100–120 cm) to ~80 cm from datum, embraces the earliest human burials of the cave and the small mammal samples here studied came from the artificial levels of 80 to 85 and 95–100 cm depth. In Period B, which comprises from ~85 to ~75 cm from datum, combustion structures, lithic tools and faunal remains (including the small mammal samples here analyzed which came from artificial levels of 75–80 cm and 75–85 cm depth) were found. A thin layer of grasses and shrubs was also found associated with some cacti Austrocactus bertinii and numerous micro flakes of silex that indicate a change in the use of the cave. In Period C, ranging between ~75 and ~55 cm from datum, small mammal samples from the artificial levels of 55 to 65 cm and -65–75 cm depth were studied. Period D, comprises from ~55 to ~35 cm from datum, embraces the strata #4 and #5, layers 4, 5 and 6, and artificial levels of -35–40 cm, -45–50 cm and -45–55 cm depth here studied. Period E, ranges from ~35 to ~15 cm from datum, includes the strata #3, #18, #25, #70, #75, #76, #78, #87, #96, #142, Layer 3, and the artificial level -15–20 cm depth analyzed here. During periods C-E, the site was preferentially used for storage of cacti A. bertinii, and the non-local rock was used specially for bifacial reduction, which proves that specialized lithic artifacts were employed for long-range hunting and processing. Two hiatuses in the sedimentary sequence associated with the anthropic activities are recorded: 1) one after ca. 5800 cal yr BP (hiatus 1) and 2) other in recent times (hiatus 2) in correspondence with gaps in the chronology. Petroglyphs (footprint style) were engraved in the walls and ceiling of the cave before ca. 3000 years BP. Period F, comprises from ~15 to datum, includes the strata #77, #170, #171, #172, #173, #175, #176, and layers 1, 2 and 2-consolidated sheep-goat manure studied here. Toward the end of Period E and during Period F A. bertinii storage continued at the time, but the site was also utilized for others domestic activities: processing meat, hides and the products of gathering and pottery manufactures. LL continued in use during the Post-hispanic period, as indicated by the find of glass and metal beads, a layer of consolidated manure, horsehair, burlap, rags, and scraps of newspaper from 1889, 1902, and 1904 (Crivelli Montero et al., 1996). The only Post-hispanic burial of the site was also found in Period F, which includes characteristic elements of the Patagonian Andean region. In the 20th century, fox hunters occupied LL and left behind various modern artifacts (Crivelli Montero et al., 1996).

The zooarchaeological and taphonomic studies of the mollusks, fishes, birds, and medium and large mammals (more than 15,000 specimens) have revealed that the subsistence of the human groups that inhabited LL was based on the exploitation of Lama guanicoe and was complemented with Rhea pennata and small size mammals such as Conepatus chinga, Chaetophractus villosus, Lycalopex gymnocercus, Lycalopex culpaeus, and Leopardus sp. (Guillermo et al., 2022). One exotic species was also identified O. o. aries, although its incorporation into human subsistence for historical context was secondary (Guillermo et al., 2022).

Material and methods

Archeological small mammal samples

In this work, 1075 cranial and postcranial specimens of small rodents and marsupials were recovered through 3 mm-sized mesh from six temporal periods of LL (Table S1), as it was indicated in the previous section.

In order to compare the taxonomical, taphonomical and paleoecological information from newest small mammal remains from LL, we used the small mammal samples dated on Post-hispanic times/Anthropocene Epoch obtained from 11 archeological and one paleontological cave sites emplaced through the Limay river basin, within a 87 km radius from LL. Studied sites were placed in the Patagonian steppe and in the ecotones between Monte-Patagonian steppe and Patagonian steppe-Forest (Figure 1a, Table S1).

Small mammal remains recovered from LL were observed under a Leica A60 binocular magnifying glass with a zoom of up to 40×. Anatomical identification was based on osteological atlas (e.g. Pearson, 1995; Udrizar Sauthier et al., 2020) and reference materials housed at collections of Grupo de Estudios en Arqueometría of Facultad de Ingeniería (GEArq-FIUBA, Buenos Aires) and Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” (MACN, Buenos Aires). The average body mass of the small mammal species represented (⩽1 kg) was obtained from the Macroecological Database of Mammalian Body Mass (MOM), version 4.1 (Smith et al., 2003). The minimum number of specimens (NISP), minimum number of skeletal elements (MNE), and minimum number of individuals (MNI) were calculated following Lyman (1994).

Taphonomic analysis

The taphonomic analysis followed the methodology developed by Andrews (1990), Fernandez-Jalvo and Andrews (1992), Fernández et al. (2017a) and Montalvo and Fernández (2019). Breakage of bones was analyzed according to the methodology proposed by Andrews (1990). The categories of digestive corrosion on incisors and molars were evaluated considering Fernández et al. (2017a) and Montalvo and Fernández (2019), who distinguished corrosion patterns on different tooth morphologies of South American rodents (caviomorphs and cricetids) and marsupials (didelphimorphs). Digestive corrosion on the postcranial elements was assessed using proximal portions of femora and distal portions of humeri (Andrews, 1990). The relative abundance of elements and other taphonomic indices (see Andrews, 1990; Montalvo and Fernández, 2019, and references therein) could not be calculated because during the excavation of LL the recovering of cranial elements was prioritized over postcranial ones (Table S2).

Anthropic exploitation of small rodents was studied based on the taphonomic attributes of cutting marks and thermoalteration, distinguishing between partial (burnt on the distal ends of premaxillaries, dentaries, incisors, and zeugopodial bones) and complete (burnt affected all bone). Both kind of thermoalteration features help to differentiate between human cooking from exposing small mammals directly to charcoal fire and natural/human-indirect fires (generally burnt or calcined bones on the whole surface, respectively), as demonstrated by neotaphonomical and zooarchaeological studies (Fernández et al., 2017b; López and Chiavazza, 2020; Medina et al., 2012; Pardiñas, 1999). In addition, taxonomical concepts that include abundance of large species and/or species with diurnal and social habits, were evaluated (e.g. Fernández et al., 2017b; Pardiñas, 1999; and references therein).

Paleoenvironmental analysis

The use of small mammals as indicators of environmental conditions are usually based on frequencies of MNI and presence/absence of some stenoic species (e.g. Andrews, 1990, 1995, 2006; Andrews et al., 2016; Fernández, 2012; García-Morato et al., 2021; López et al., 2021). Raptor pellet samples are considered good estimators of prey abundance and as tools for yielding paleoenvironmental models (e.g. Andrade et al., 2016; López et al., 2021). In this sense, 35 samples of small mammals from pellets of avian raptors Tyto furcata, Bubo magellanicus (Strigiformes), and Geranoaetus melanoleucus (Accipitriformes) taken within a 110 km radius from LL archeological site, placed in Patagonian steppe and in the ecotones Monte-Patagonian steppe and Patagonian steppe-Forest, were used as actualistic parameters (Andrade, 2015; Formoso, 2013; Massoia, 1988a, 1988b; Massoia et al., 1991, 1999; Massoia and Lartigau, 1995; Massoia and Pardiñas, 1986, 1988a, 1988b; Pardiñas et al., 2003; Pardiñas and Massoia, 1989; Pardiñas and Teta, 2013; Tammone et al., 2014, 2020; Teta et al., 2005; Trejo and Lambertucci, 2007; Figure 1a; Table S3).

In addition, habitats were inferred applying the Habitat Weighting Method following the procedures explained by Evans et al. (1981), Andrews (2006) and Andrews et al. (2016). The habitat classification for each of the small mammal species found in archeological sites and in recent pellet samples is shown in Table S4. A total of seven habitats were distinguished based on trapping data and further analyses published by Guillermo et al. (2021):

1) Forests – vegetation formations of one–three species of Nothofagus plus Austrocedrus. These formations can be open or dense, with or without bamboo at the understory.

2) Shrub – bushes of more than1 m tall including small trees (e.g. Berberis, Chacaya, Colletia, Discaria, Fabiana or bushy Nothofagus antarctica and bamboo).

3) Steppe – formations of low bushes with less than1 m tall, mixed with bunchgrasses and open areas of bare grounds. Bushes that can be found belong to the genus Acaena, Baccharis, Berberis, Mulinum and Senecio.

4) Bunchgrass – relatively pure stands of one or more species of bunchgrasses.

5) Weeds and turfs – mixed formations of dense weeds (dandelions, thistles, and grasses), non-bunch grasses and herbs (white clovers and filarees) that are usually present in moist places or bordering moist areas.

6) Rocks – cliffs or tumbled boulders and rocks that are large enough to provide refugia for mice. Usually linked to mountainous or hilly areas.

7) Bare grounds – habitats with a lack of vegetation cover, characterized by the presence of a substrate of fine scree or rocks in which rocky refugia are scarce or absent. This habitat is usually found on windswept hilltops.

Considering the variation in sample size of the different archeological and recent pellet samples, rarefaction curves were calculated for each sample in order to assess reliability of richness (S) and Shannon diversity index (H) with the relative abundance of species in terms of MNI. In addition, Shannon diversity index (H) was also used to evaluate temporal changes in diversity. Possible size biases in the present and archeological samples produced by prey-predator preferences or preservation contexts were analyzed applying a Detrended Correspondence Analysis (DCA) to avoid the arch artifact usually present in Correspondence Analyses (CA), when non-linear correlations are present between the axes. Small mammal species were classified into four different body size categories (0–40 gr, 40–80 gr, 100–170 gr and 250–500 gr) following García-Morato et al. (2021). A Principal Component Analysis (PCA) was also applied for evaluating the differences in the abundance of the species represented in recent (raptor pellets) and archeological samples. This exploration was made on a data matrix of relative abundance (MNI), taken from Tables S1 and S2, log-transformed (ln). Statistical analyses were performed using the software PAST (Paleontological STatistics), version 4.12 (Hammer, 1999).

Results

Taxonomic structure of small mammals from Epullán Grande and other sites of Limay River basin

Of the 15 recorded taxa of LL, mainly at species level, two correspond to marsupials, three to caviomorph rodents, and 10 to cricetid sigmodontine rodents. The caviomorph Ctenomys sp. (Figure 2a) and the sigmodontine Phyllotis cf. P. pehuenche (Figure 2b) were abundant and registered in all periods of LL. Other taxa found in various periods were the caviid Microcavia australis (periods A, C-F; Figure 2c), the sigmodontines Eligmodontia sp. (Figure 2d) and Reithrodon auritus (periods A-B, D-E; Figure 2e), the sigmodontine Euneomys chinchilloides (periods A-C, E; Figure 2f) and the marsupial Thylamys pallidior (periods C-F; Figure 2g). Nevertheless, most species were mainly found in the later periods: the sigmodontines Abrothrix olivacea (Figure 2h), Abrothrix hirta (Figure 2i), Akodon iniscatus (Figure 2j), Loxodontomys micropus (Figure 2k) and Oligoryzomys longicaudatus (Figure 2l), the caviomorph Galea leucoblephara (Figure 2ll), and the marsupial Lestodelphys halli (Figure 2m). The sigmodontine Calomys musculinus (Figure 2n) was only recorded in the earliest period (A).

Figure 2.

Dentaries of the small mammals (one example for each taxon) recovered at Epullán Grande site: Ctenomys sp. from Period E (a). Phyllotis cf. P. pehuenche from Period E (b). Microcavia australis from Period F (c). Eligmodontia sp. from Period B (d). Reithrodon auritus from Period A (e). Euneomys chinchilloides from Period A (f). Thylamys pallidior from Period E (g). Abrothrix olivacea from Period E (h). Abrothrix hirta from Period B (i). Akodon iniscatus from Period (e) (j). Loxodontomys micropus from Period F (k). Oligoryzomys longicaudatus from Period E (l). Galea leucoblephara from Period E (ll). Lestodelphys halli from Period F (m). Calomys musculinus from Period A (n). Scales = 10 mm.

Rarefaction analyses revealed that the species diversity curves of fossil samples have more asymptotic tendency than species richness curves (Figure S1). Therefore, for fossil samples, species diversity comparisons are more reliable than richness. Both rarefaction curves for recent raptor samples (Figure S2) are consistent in excluding from the analysis some samples (#8, #12, #13, #18, #26, see Table S3) that have been biased mostly by their small sample sizes (MNI < 63).

Figure 3 shows that periods E and F of LL and Post-hispanic unit of ECh also present higher values of species diversity index than the recent pellet samples recovered from the LL cave and the nearest localities (Figure 1). Other fossil site so-call Álvarez 4 (AZ4), emplaced in the middle Limay river basin in a typical Patagonian environment, exhibits the highest species diversity (Figure 3). However, some fossil exceptions with lower diversity values were found in the same section of the river basin, such as Alero Nestares, Casa de Piedra de Ortega (CPO), La Marcelina 1 (Mar 1) and Sarita II, possibly because they were mostly accumulated by humans with their dietary preferences for caviomorphs (for more details see Discussion). Toward the upper Limay river basin, in the ecotone Patagonia-Forest environment, the Post-hispanic/Anthropocene units of Cueva Traful I (CTI), Cueva del Caballo and Puma II yield higher taxonomic diversity than recent pellet samples recovered in the same caves (Figure 3).

Figure 3.

Estimates, based on MNI, of Shannon species diversity (H) in fossil (selected temporal units, see Table S1) and recent raptor samples (#8, #12, #13, #18, #26 were excluded, see Table S3) from Limay River basin (Northwestern Patagonia). The abbreviations of the fossil sites and recent pellet localities are indicated in Tables S1 and S3.

PCA results (Figure 4) ordered the fossil samples from the later periods of LL (E and F) close to the fossil sample of ECh and near to some recent raptor samples to the negative values on component 1 (30.5% of the total variance) and positive values on component 2 (16.7% of the variance). All these samples were characterized by a mixture of Patagonian (L. halli) and Monte taxa (A. dolores, A. iniscatus, C. musculinus and T. pallidior). The samples from earlier temporal periods of LL, along with both fossil (AZ4, Alero Nestares, Cañadón Las Coloradas 1 [CLC1], CPO, Mar 1, Sarita II), and other recent samples recovered from Patagonian environments (Figure 1a) are grouped toward the negative values of both axes, according to the abundance of the caviids G. leucoblephara and M. australis. On the contrary, the fossil (Cueva del Caballo, CTI, Puma II) and recent samples emplaced toward the Forest-Patagonian ecotone and their typical species (A. hirta, Geoxus valdivianus, Irenomys tarsalis, L. micropus, Paynomys macronyx) are ordered to positive values of component 1 and negative values of component 2 (except Cueva del Caballo).

Figure 4.

Principal Component Analysis (PCA) based on the log-transformed MNI of the different species of fossil (black dots) and recent (red dots) samples from Limay river basin (Northwestern Patagonia). The abbreviations of the fossil sites and recent pellet localities are indicated in Tables S1 and S3.

PCA results (Figure 4) also show that most of the generalist or opportunistic species (A. olivacea, Eligmodontia sp. and O. longicaudatus) were situated toward to the positive values of both axes along with some Patagonia and Patagonian-Forest ecotone recent and fossil samples. Generalist species allied to Monte desert (C. musculinus) are also separated at the positive values of axis 2, but grouped together with most of fossil (including LL and ECh) and recent samples associated to Monte-Patagonian ecotones to negative values of axis 1.

Results obtained for the Habitat Weightings (Figure 5a) calculated for LL show that steppes, shrubs and rocky areas were the most common environments, even in Post-hispanic times/Anthropocene Epoch. Only the sites located in the Subandean-Andean Patagonian forests region still showing a similar percentage of forest cover to the one observed in the lower levels of LL. Nonetheless, it is remarkable the high abundance of bare ground areas in CTI amongst all the archeological sites analyzed. In relation to the habitats inferred from recent pellets (Figure 5b), steppes increase in comparison with the archeological units, especially in the Monte-Patagonia ecotone and Central and Occidental Districts. Finally, shrubs, weed and turfs habitats increase with respect to steppes in the Subandean District and Andean Patagonian forests ecotone.

Figure 5.

Results for the Habitat Weighting Method based on the small mammal assemblages analyzed from 12 archeological sites (a) and 35 recent pellet samples (b) located in the Limay river basin. The abbreviations of the fossil sites and recent pellet localities are indicated in Tables S1 and S3.

Taphonomy analysis of Epullán Grande small mammals

The main taphonomic attributes recorded in the studied samples of LL are shown in Table 1. The fragment of a pellet was recovered from Period B (Figure 6a), and other three fragments were found in Period E (Figure 6b). Of the analyzed remains (incisors in situ (retained in the alveoli) and isolated, molars in situ and isolated, distal end of humeri, and proximal end of femora = 1318), only 5.2% exhibited signs of digestive corrosion, all in light category (Table 1), revealing sparsely pitting on bone surfaces and little reduction of the enamel or loss of shine (Figure 6c). None of them presented digestive corrosion in the categories of greatest damage (i.e. moderate, heavy and extreme). The proportion of digested teeth and bones remained at low grade and at quite constant values throughout the entire sequence (Period A = 3.3%; Period B = 5.3%; Period C = 2.8%; Period D = 5.3%; Period E = 7.6%; Period F = 2.9%).

Table 1.

Taphonomic attributes of small mammal samples from the different temporal periods of Epullán Grande cave site (Limay river basin, Northwestern Patagonia, Argentina).

Epullán	Period A		Period B		Period C		Period D		Period E		Period F
Grande	N	%	N	%	N	%	N	%	N	%	N	%
Complete pellets preserved	0		0		0		0		0		0
Fragment pellets preserved	0		1		0		0		3		0
Digestive corrosion
Incisor in situ (light)	5	12.5	2	6.7	2	4.4	0	0	12	12	2	5
Incisor isolated (light)	2	8.7	0	0	0	0	2	15.4	6	11.8	0	0
Molar in situ (light)	1	0.8	7	8.2	1	2.2	1	2.5	7	2.8	3	2.3
Molar isolated (light)	0	0	0	0	0	0	0	0	0	0	0	0
Humerus (light)	0	0	0	0	0	0	0	0	0	0	1	100
Femur (light)	1	100	0	0	0	0	2	100	11	50	0	0
Termoalteration in caviomorphs
Partially carbonized specimens	13	7.8	2	2	5	5.9	4	6.1	15	10.3	4	11.1
Termoalteration in cricetids and marsupials
Total carbonized specimens	1	1.7	1	3.1	0	0	0	0	5	1.9	1	0.9
Breakage
Complete skull	0	0	0	0	0	0	0	0	0	0	0	0
Maxillary with zygomatic	4	25	7	43.8	1	11.1	1	8.3	6	8.7	3	15
Maxillary without zygomatic	12	75	9	56.2	8	88.9	11	83.3	63	91.3	17	85
Complete mandible	16	17.6	7	12.5	9	30.0	2	9.5	27	19.2	17	20
Mandible with ascendant ramus broken	29	31.9	24	42.9	9	30.0	11	52.4	74	52.4	31	36.5
Without ascendant ramus	41	45.0	15	26.8	5	16.7	7	33.3	27	19.2	25	29.4
Mandible with inferior edge broken	5	5.5	10	17.8	7	23.3	1	4.8	13	9.2	12	14.1
Broken postcranial bones	0	0	4	80.0	1	16.7	11	91.7	56	57.7	2	66.7
Postdepositional processes
Weathering	0	0	0	0	0	0	0	0	0	0	0	0
Root traces	0	0	0	0	0	0	0	0	0	0	0	0
Transport	0	0	0	0	0	0	0	0	0	0	0	0
Manganese oxide staining	24	10.8	10	7.8	2	2.2	0	0	3	0.7	0	0

Figure 6.

Examples of taphonomic attributes found at different periods of Epullán Grande site: fragments of pellet preserved from Period B (a) and Period E (b). Light digestive corrosion in sigmodontine lower incisor from Period F (c). Premaxillary of Ctenomys sp. partially carbonized from Period E (d). Dentary of Phyllotis cf. P. pehuenche with manganese oxide staining from Period A (e). Arrows indicates the areas affected. Scales = 1 min.

On the one hand, 43 skull and dentary elements (7.3%) of Ctenomys sp. (MNE = 42 in all periods) and M. australis (MNE = 1 in Period E) with the anterior parts (premaxillary and dentary diastema) partially carbonized (Figure 6d) were observed. This taphonomic attribute is considered one of the most reliable signatures to assign human consumption (see Material and Methods, Taphonomic analysis section) and it was detected in higher percentages at the later periods, especially on caviomorph rodents (these animals are easily hunted by humans due to their social and predictable behavior; Period A = 7.8%; Period B = 2%; Period C = 5.9%; Period D = 6.1%; Period E = 10.3%; Period F = 11.1%). On the other hand, lower proportions of burned bones, without this specific pattern (1.6%), were found in smaller and solitary cricetids and marsupials (Period A = 1.7%; Period B = 3.1%; Period C = 0%; Period D = 0%; Period E = 1.9%; Period F = 0.9%).

Most of the studied bones were fractured (89.3%), showing sharp edges and rough surfaces, reaching almost the totality in some periods (Period A = 99.1%; Period B = 73.1%; Period C = 88.9%; Period D = 97.8%; Period E = 86.6%; Period F = 99.1%). All crania were heavily broken, thus only the maxillae with or without zygomatic arches were preserved (Table 1). Conversely, about 10–30% of dentaries were completed, although most of these elements were found with the ascendant rami broken or without it (Table 1). Despite the sullegic factors (i.e. recent fossil modifications occurred during the excavation, such as do not recover all elements) afore-mentioned (see Material and Methods, Taphonomic analysis section), lower proportion of postcranial bones were broken (59.7%) in comparison with the cranial ones (Table 1).

Finally, regarding the postdepositational processes (Table 1), evidences of small and sparse blackish dots-shaped in bones related to manganese oxide staining (Figure 6e) were registered with higher incidence in the earlier periods (Period A = 10.8%; Period B = 7.8%; Period C = 2.2%; Period E = 0.7%). Neither of the analyzed bones showed cracking/exfoliations associated with the exposure to the meteoric agents nor polish/round/shine edges produced by water/wind transport.

Discussion

Different taphonomic trajectories and their small mammal taxonomic biases

Previous taphonomic studies performed with small mammal samples from LL indicate the actions of both barn owl (T. furcata) predation and human exploitation (Crivelli Montero et al., 1996; Pardiñas, 1999; Pardiñas and Teta, 2013). The finding of pellets along with the low proportion of light digestive corrosion found in the analysis of newest samples are in agreement with an accumulation produced by this strigiform (Andrews, 1990; Fernández et al., 2017a; Montalvo and Fernández, 2019; Pardiñas, 1999). T. furcata usually nests in caves and rockshelters, inhabits open landscapes and displays an opportunistic trophic behavior, feeding mostly on nocturnal sigmodontine rodents (e.g. Bellocq, 2000; Montalvo and Fernández, 2019, and references cited therein). In fact, part of the studied assemblages from LL are composed by the sigmodontine rodents (Table S1). The other part of the LL samples are formed by caviomorph rodents; even, some of its premaxillae and dentaries presented the typical burning pattern of human cooking direct on charcoal (Fernández et al., 2017b; López and Chiavazza, 2020; Medina et al., 2012; Pardiñas, 1999). Although this taphonomic signature was recorded throughout the entire LL sequence, it increased toward the last ca. 1000 years BP (periods E and F), in line with the exploitation of more faunal resources at this site (Guillermo et al., 2022), and at regional level (e.g. Cordero, 2011; Fernández et al., 2016, 2017b; Guillermo et al., 2020a, 2020b, 2021).

Body size analyses indicate a certain separation between recent pellets and fossil samples of Limay river basin (Figure 7). Although some exceptions can be observed due to sample size or to the intervention of different taphonomic processes, most of fossil sites are usually characterized by a major abundance of species of larger body sizes than in current pellet samples. Most of the archeological sites show the intervention of different accumulation agents, usually raptors and humans. In the case of humans, rodent species of more than 100 gr and gregarious-grouped habits are usually predated (i.e. Ctenomys sp., G. leucoblephara and M. australis), favoring an increase of the representation of these large sized species. In addition, the presence of Ctenomys sp. in archeological sites is remarkable in comparison with the pellet samples. Species belonging to this rodent genus are fossorial and could be incorporated to the sites by natural death (e.g. Fernández, 2012; García-Morato et al., 2021). However, the abundance of caviomorph rodents decrease toward the more recent levels of LL, which could be the consequence of the introduction of ovicaprids in the area, as they compact the soils trougth trampling (Pardiñas and Teta, 2013; Teta et al., 2014). Both facts (feeding habits of birds of prey and soil degradation) could also explain why Ctenomys sp. is absent in most of the recent samples.

Figure 7.

Detrended Correspondence Analysis (DCA) for the different recent raptor pellet and archeological samples from Limay river basin, based on the body size classification applied. The abbreviations of the fossil sites are indicated in Tables S1.

Finally, a good preservation and a rapid deposition of the small mammal assemblages in the stratigraphic context is demonstrated by the scarce postdepositional taphonomic processes (e.g. absence of weathering, rodent marks, hydraulic transport, root action) observed in LL. Otherwise, some of the skeletal remains analyzed, especially in periods A and B, display manganese coatings suggesting the presence of wet or damp conditions (Fernández-Jalvo and Andrews, 2016) during the earliest formation of the LL sequence, between ca. 11.5 and 7.8 cal ky BP. The high breakage proportions of elements contrasts with the values and types of digestion associated with the activity of the predator involved (T. furcata) in the formation of the small mammal assemblage (see Andrews, 1990; Montalvo and Fernández, 2019). The evidence of sharp edges and rough fracture surfaces indicates trampling action, a typical taphonomic process occurred at caves (Andrews, 1990; Fernández, 2012; Pardiñas, 1999), as the main taphonomic process causing the high breakage values observed.

Small mammals at Limay River basin from latest Pleistocene to Anthropocene

In general terms, the small mammal communities of Limay river basin have changed abruptly after the end of the Last Glacial Maximum (Pearson, 1987; Pearson and Pearson, 1993; Tammone et al., 2014). Postglacial small mammal assemblages, as occurred in other areas of Patagonia and Central Andes, have shown a taxonomic stability or minor variations in species representation (e.g. Crivelli Montero et al., 1996; Fernández, 2012; Fernández et al., 2012, 2016, 2018, 2021a, 2021b; López et al., 2021; Pardiñas and Teta, 2013; Pearson, 1987; Pearson and Pearson, 1993; Tammone et al., 2020; Teta et al., 2005). However, glacial, charcoal, and pollen records indicate several paleoenvironmental changes during the Early and mid-Holocene, whilst the current environmental conditions in the area were established during the last 3000 years, although with some intervals of colder or warmer events (e.g. Bianchi, 2007; Heusser, 1993; Iglesias et al., 2014, 2017; Markgraf, 1983; Mercer, 1976; Prieto et al., 2021; Prieto and Stutz, 1996; Rabassa, 2008; Whitlock et al., 2006). For example, for the upper Limay river basin, pollen record from Lake El Trébol (41º04’S, 71º29’W; Río Negro province) yielded the development of the modern vegetation at the site, embracing the High Andean and mixed forest, although with some temporal element coming from the Patagonian steppe communities (Iglesias et al., 2014, 2017; Whitlock et al., 2006). For the middle Limay river basin, more accentuated climate variability was detected at polinic record of LL and ECh between ca. 2.5 and 1.6 cal ky BP, promoting an increase in shrubs over grasses and the establishment of a mosaic of Monte-Patagonian steppe, similar to the recent conditions (Prieto et al., 2021).

Such stability in the small mammal communities through the Holocene had been understood as resilience feature of most of the species to environmental changes aforementioned (e.g. Fernández et al., 2016, 2018; Pardiñas and Teta, 2013). Among the scarce variations recorded, we can find the expansion of Holochilus brasiliensis, an amphibious cricetid rodent that inhabit temperate to subtropical habitats, during the warm pulses the Medieval Climate Anomaly, and Lestodelphys halli, an endemic marsupial of Patagonian affinities, toward the cold and dry pulse of the Little Ice Age (Teta et al., 2005). However, is clear that the major change in the taxonomic composition of small mammal communities had been occurred at recent times. Then one might ask: are these expansions/retractions and general variations related to climatic changes or anthropic disturbances? When did the deep change occur in the small mammal communities? At the beginning of the Anthropocene Epoch or in its most pronounced along the 20th century?

Removing those fossil samples strongly bias for the abundance of caviomorph rodents (Ctenomys sp., G. leucoblephara and M. australis) by the human hunting (including the periods A-C of LL), the fossil samples from both the upper and middle Limay River basin associated to Anthropocene Epoch had greater diversity than recent pellet samples recovered in the same environments. This scenario has also been documented in several other areas of Patagonia (e.g. Andrade and Monjeau, 2016; Fernández et al., 2012, 2021a; Pardiñas et al., 2000, 2012; Tammone et al., 2020; Teta et al., 2014) and some adjacent regions of central Argentina (e.g. Fernández, 2012, 2014; García-Morato et al., 2021; López et al., 2021; Teta et al., 2014). Since the Post-hispanic period/Anthropocene Epoch, a gradual entry of domestic exotic fauna (horse, cow, and sheep) to northwestern Patagonia took place (Fernández and de Mendoza, in press; Valenzuela et al., 2023), along with intentional fires, the extraction of hydrocarbons and minerals gave rise to a series of environmental modifications, mainly in the soil and vegetation cover, by the action of trampling and overgrazing (e.g. Bertiller and Bisigato, 1998). This also led to a restructuring of the small mammal communities in the area, with the prevalence of generalist or opportunistic species, such as A. olivacea, C. musculinus, Eligmodontia sp., and O. longicaudatus, to the detriment of other more stenoic species, such as Euneomys spp., L. halli, L. micropus and R. auritus, that were more abundant and, some of them dominant, during the Holocene even up to the Anthropocene Epoch (Crivelli Montero et al., 1996; Fernández et al., 2016; Guillermo et al., 2021; Pardiñas and Teta, 2013; Pearson, 1987; Pearson and Pearson, 1993; Tammone et al., 2020; Teta et al., 2005).

On the one hand, some changes in habitats were recorded in some fossil sites of the upper Limay river respect to the recent ones. For example, the archeological small mammal samples from CTI, Cueva del Caballo, and Puma II reflect a large dominance of bare ground areas, mainly inferred through the high abundance of Euneomys spp., contrasting with its almost absence in recent samples from the upper Limay river basin (Pardiñas and Teta, 2013; Pearson, 1987; Pearson and Pearson, 1993; Rebane, 2002; Tammone et al., 2020). In turn, the high abundance of O. longicaudatus was linked to the expansion of the exotic plant Rosa rubiginosa, which was used by this rodent as a refuge and food source (Pardiñas and Teta, 2013; Tammone et al., 2020; Andrade et al., in this volume; and references cited therein). On the other hand, the restructuration of small mammal communities from middle Limay river basin embraces some species (e.g. A. olivacea, C. musculinus, Eligmodontia spp.) allied to shrubby and overgrazed open environments (Teta et al., 2014).

Finally, most of the authors cited in the previous paragraphs indicated that the modern configuration of small mammal communities took place about 400-150 years ago (e.g. Fernández et al., 2016; Guillermo et al., 2021; Pardiñas and Teta, 2013; Tammone et al., 2020; Teta et al., 2014). However, the pool data analyzed here for Limay river basin, comparing fossil samples from the last 400 years with recent pellet samples generated by birds of prey, revealed that the restructuring of small mammal communities occurred more recently, toward the final stretch of the current Anthropocene Epoch, along the last century (e.g. Barnosky et al., 2011; Ceballos et al., 2015, 2020; Waters et al., 2022; Waters and Turner, 2022), when human populations and their activities grew remarkably seriously affecting the environment. Congruently, in the adjacent northern region of the Central Andes, the amphibian and endemic sigmodontine rodent Holochilus lagigliai was lastly documented at the Atuel river (Mendoza province) toward the middle half of the last century, after some fossil findings for the Late-Holocene (Fernández et al., 2017c; Fernández and Pardiñas, 2018; López et al., 2021). The biological extinction of this rodent was also accompanied by declines in the diversity of small mammal communities and an increase of opportunistic species (Fernández, 2012, 2014; López et al., 2021).

Conclusions

The taphonomic analysis carried out on the newest samples of LL confirms the conclusions made in previous works: 1) the predatory activity of the common owl T. furcata on sigmodontine rodents as main agent in the formation of archeological small mammal assemblages; 2) human consumption on social and diurnal caviomorph rodents; in addition, in this work we observed that the anthropic incidence on caviomorphs increased toward the last 1 cal ky BP, in line with what occurred on a general scale in Limay river basin. Otherwise, through the novel analysis of manganese oxide staining, we could associate higher levels of moisture during the earliest formation of the LL sequence, between ca. 11.5–7.8 cal ky BP.

The taxonomic results indicate a major diversity in the small mammal fossil assemblages to the upper periods of LL and other fossil sequences of the Limay basin of the Anthropocene Epoch. Conversely, opportunistic sigmodontines such as A. olivacea, C. musculinus, Eligmodontia spp. and O. longicaudatus greatly growth their proportions in recent owl pellet samples. Apparently, in the Limay river basin, as it was observed in other areas of Patagonia, the environmental degradation took place very recently (i.e. the last century).

Finally, the evidence presented here indicated that the stenoic and specialist species, such as Euneomys spp., L. halli, L. micropus and R. auritus were present and, some of them, even were abundant in the temporal units associated with the Anthropocene Epoch, but now they are in retraction. This fact, in addition with the drop in the diversity of recent pellet assemblages, shows that the restructuring of small mammal communities possibly occurred in the final stretch of the current Anthropocene Epoch, mostly during the last century, when human populations and their activities grew remarkably.

Supplemental Material

sj-jpg-1-hol-10.1177_09596836241231450 – Supplemental material for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective

Supplemental material, sj-jpg-1-hol-10.1177_09596836241231450 for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective by Fernando J Fernández, Ailín A Guillermo, José Agustín Cordero, Pablo Teta and Sara García-Morato in The Holocene

Supplemental Material

sj-jpg-2-hol-10.1177_09596836241231450 – Supplemental material for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective

Supplemental material, sj-jpg-2-hol-10.1177_09596836241231450 for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective by Fernando J Fernández, Ailín A Guillermo, José Agustín Cordero, Pablo Teta and Sara García-Morato in The Holocene

Supplemental Material

sj-xlsx-3-hol-10.1177_09596836241231450 – Supplemental material for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective

Supplemental material, sj-xlsx-3-hol-10.1177_09596836241231450 for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective by Fernando J Fernández, Ailín A Guillermo, José Agustín Cordero, Pablo Teta and Sara García-Morato in The Holocene

Supplemental Material

sj-xlsx-4-hol-10.1177_09596836241231450 – Supplemental material for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective

Supplemental material, sj-xlsx-4-hol-10.1177_09596836241231450 for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective by Fernando J Fernández, Ailín A Guillermo, José Agustín Cordero, Pablo Teta and Sara García-Morato in The Holocene

Supplemental Material

sj-xlsx-5-hol-10.1177_09596836241231450 – Supplemental material for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective

Supplemental material, sj-xlsx-5-hol-10.1177_09596836241231450 for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective by Fernando J Fernández, Ailín A Guillermo, José Agustín Cordero, Pablo Teta and Sara García-Morato in The Holocene

Supplemental Material

sj-xlsx-6-hol-10.1177_09596836241231450 – Supplemental material for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective

Supplemental material, sj-xlsx-6-hol-10.1177_09596836241231450 for Small mammal records from Limay river basin (Northwestern Patagonia) in the Anthropocene from a taphonomical and paleoecological perspective by Fernando J Fernández, Ailín A Guillermo, José Agustín Cordero, Pablo Teta and Sara García-Morato in The Holocene

Footnotes

Acknowledgements

We are grateful to the archeological rescue and prehistoric research projects in the Alicurá area and Piedra del Águila directed by Amalia Sanguinetti de Bórmida and Eduardo Crivelli Montero, in agreement between Hidronor and the UBA. Mabel Fernández, part of the laboratory and excavation team, collaborated with the interpretation of the stratigraphy. Daiana Coll help us with the preparation of . The editor Gill Oliver, the guest editors Julian Mignino and José Manuel López and two anonymous reviewers made comments and corrections that improve this work.

Author contributions

FJF had the idea and structure of the paper. AG, FJF, JAC and PT worked with the LL newest samples. AG, FJF and JAC made the taphonomic analysis. FJF and PT provided background fossil and recent samples information and carried out the taxonomic identifications. FF and SGM performed and interpreted the statistical analyzes. AG, FJF, JAC and SGM prepared the images. All authors contributed to the clarification of the text.

Funding

The author(s) disclosed receipt of the following financial support for the research, authorship, and/or publication of this article: Laboratory work was funded by two grants (i- COOPB20589 / PID2021-126933NB-I00) of Consejo Superior de Investigaciones Científicas de Cooperación Internacional and Museo Nacional de Ciencias Naturales.

ORCID iDs

Fernando J Fernández

Ailín A Guillermo

Sara García-Morato

Supplemental Material

Supplemental material for this article is available online.

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