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... ecological knowledge, landscape use and natural resources management; Dilkes-Hall et al., 2020). Pollen assemblages preserved in coprolites reflects the composition and dynamics of the vegetation at a regional level (Bryant and Holloway, 1983;Reinhard and Bryant, 1992;Carrión et al., 2004;Caruso Fermé et al., 2018) while the changes observed in the microfragment record are of local relevance and would complement the regional inferences evidenced in the pollen record (Jørgenset et al., 2012;Wood et al., 2012;Rawlance et al., 2016;Mosca Torres et al., 2018;Llano et al., 2020;Petrigh et al., 2021). ...
Paleoenvironmental records in the eastern Andes (16–26° S), broadly indicate a gradual trend towards aridity from the Early to the Middle Holocene with certain chronological discrepancies. Even under this adverse climatic scenario, the archaeological record suggests the presence of hunter-gatherers in the southern Argentinian Puna associated with the availability of resources for their subsistence, even during drier than present periods. Herbivore coprolites contain multiple proxies that provide independent records of paleovegetation and paleoenvironments. Among them, plant tissue fragments and pollen grains, offer a powerful and complementary insight on past vegetation changes and their relationship with climatic conditions during the Holocene, particularly in the southern Argentine Puna. Thus, in this work camelid coprolites recovered from the archaeological site Quebrada Seca 3 (QS3) were analyzed using microhistological and palynological techniques to reconstruct the diet of these herbivores but also to infer vegetation and environmental changes from the Early to Middle Holocene (ca. 9600–6100 cal yrs BP) in the southern Argentine Puna (26°S). Results of both records show similar patterns in the percentages and species richness (especially those growing in wetlands) within and between coprolites dating to the Early Holocene (9600-8000 cal yrs BP) which suggest an environmental stability during this period. Around 8000 cal yrs BP, an increased variability in the richness of grasses, shrubs, dwarf shrubs, annual herbs and sedges species shows the gradual establishment of dry conditions, peaking around 6100 cal yrs BP. By this time, variability in the plant content of coprolites reflects seasonality in the assemblages due to the presence/absence of annual herbs either in the pollen or microfragment records. Therefore, it is likely that coprolites were from different animals that were hunted at different times of the year, supporting the evidence that QS3 has been occupied for short intervals of time after 7200 cal yrs BP when environmental conditions were extremely arid. The complementary analysis of the microfragments and pollen records preserved in coprolites provide valuable, complementary and alternative archive for herbivorous diet and for vegetation and environmental reconstructions, reinforcing that multiproxy approaches are necessary to obtain maximum information from coprolite studies.
... Coprolite palynology is often associated to cave sedimentary environments, providing a diversity of biogenic materials where pollen, spores and non-pollen palynomorphs (NPPs) can be relatively well preserved (Alcover et al., 1999;Carrión et al., 1995aCarrión et al., , 1995bCarrión et al., , 2004Carrión et al., , 2006Carrión et al., , 2007Carrión et al., , 2008Carrión et al., , 2018Daura et al., 2017;Gatta et al., 2016;González-Sampériz et al., 2003;Horwitz and Goldberg, 1989;Marais et al., 2015;Ochando et al., 2020a;Scott, 1987Scott, , 1994Yll et al., 2006). Evaluating the palaeoecological potential of coprolites requires more experimental research, especially on the taphonomy of the pollen assemblages as pollen transport, deposition, and preservation (Bottema, 1975;Carrión et al., 2009;Coles and Gilbertson, 1994;Davis, 1990;Fernández-Jalvo et al., 1999;Hunt and Fiacconi, 2018;Hunt and Rushworth, 2005;Navarro et al., 2000Navarro et al., , 2001. ...
Palynological investigations on Gravettian coprolites of Abrigo do Lagar Velho (Vale do Lapedo, Leiria, Portugal) dated between ca. 30 to ca. 27.3 ka cal BP are presented. The palaeoecological data shows a semi-forested landscape with Pinus, Poaceae, Erica, Artemisia, Juniperus and Quercus as the dominant taxa. A diversity of trees, shrubs, and herbs is also noticeable, including broad-leaf trees, Mediterranean woody shrubs and trees, conifers, xerothermophytes, indicators of saline substrates and heliophytes such as Erica, Asteroideae, Cistaceae, Ephedra fragilis and E. distachya. The overall reconstruction is a mosaic of different landscapes or types of vegetation around the rock-shelter. Indeed, we provided approach on the Gravettian vegetation of the Iberian Peninsula, connecting the palaeoecosystem with the biogeography of past human populations during the late Pleistocene.
... Coprolite palynology is often associated to cave sedimentary environments, providing a diversity of biogenic materials where pollen, spores and non-pollen palynomorphs (NPPs) can be relatively well preserved (Alcover et al., 1999;Carrión et al., 1995aCarrión et al., , 1995bCarrión et al., , 2004Carrión et al., , 2006Carrión et al., , 2007Carrión et al., , 2008Carrión et al., , 2018Daura et al., 2017;Gatta et al., 2016;González-Sampériz et al., 2003;Horwitz and Goldberg, 1989;Marais et al., 2015;Ochando et al., 2020a;Scott, 1987Scott, , 1994Yll et al., 2006). Evaluating the palaeoecological potential of coprolites requires more experimental research, especially on the taphonomy of the pollen assemblages as pollen transport, deposition, and preservation (Bottema, 1975;Carrión et al., 2009;Coles and Gilbertson, 1994;Davis, 1990;Fernández-Jalvo et al., 1999;Hunt and Fiacconi, 2018;Hunt and Rushworth, 2005;Navarro et al., 2000Navarro et al., , 2001. ...
... lakes sediments, peat or bogs. Particularly, coprolite pollen analyses give evidence about diet, seasonality and paleoenvironment (Carrión et al. , 2004(Carrión et al. , 2005Carrión 2002;Chaves and Reinhard 2006;Horrocks et al. 2002Horrocks et al. , 2003Horrocks et al. , 2008Reinhard and Bryant 1992;Riley 2008;Velázquez et al. 2010Velázquez et al. , 2014Velázquez et al. , 2015Velázquez and Burry 2012;Wood et al. 2012;among others). ...
... Pollen analyses of coprolites have been studied to infer paleodiet, home range and paleopharmacology. Relative to these topics, the value of coprolites to perform paleoenvironmental inferences has been neglected (Carrión et al. , 2004. However, research on coprolites is less common for paleoenvironmental reconstructions than analyses of lake deposits and peat bogs in part because it is difficult to find coprolites in a good state of preservation. ...
... The comparison between pollen spectrum of coprolites and contemporaneous sediments has shown that coprolites reflect a good representation of local and regional flora. In addition, coprolites have shown higher pollen diversity and abundance of entomophilous taxa than sediments (Carrión 2002;Carrión et al. 2004). ...
The aim of this study was to analyze pollen from the outer parts of camelid coprolites and contemporaneous sediments to evaluate the potential of coprolites for the reconstruction of the vegetation history of Patagonia during the Holocene. The pollen composition of camelid coprolites coming from archeological layers between 10,920 yrs. cal BP and 2823 yrs. cal BP found in the sites Cerro Casa de Piedra 5 and 7 (Perito Moreno National Park, Santa Cruz, Argentina) were analyzed. Coprolites were divided into outer and inner subsamples. Outer subsample pollen extraction was performed through acetolysis and determination and counting under optical microscope. Morisita-Horn index and Wilcoxon test were performed to evaluate similarities between pollen spectra of outer subsamples of coprolites and contemporaneous sediments. The dominant pollen types were: Nothofagus (Nothofagaceae), Empetrum (Ericaceae), Asteraceae subf. Asteroideae, Nassauvia (Asteraceae subf. Mutisioideae), Poaceae and Caryophyllaceae. The similarities found between outer subsamples of coprolites and contemporaneous sediments reflected the dominant pollen types of the vegetation and forest development, whereas significant differences could be attributed to coprolite pollen spectra that may be biased by season of coprolite deposition, selectivity of the diet and/or higher temporal resolution. Also, the presence of pollen types in outer subsamples of coprolites absent in sediments could give information for paleoenvironmental reconstructions by enriching information of the past flora. This study demonstrates that coprolites can be a valuable tool for paleoenvironmental reconstructions in arid and semiarid areas where depositional environments that support pollen preservation are typically rare.
... de los coprolitos también a la reconstrucción paleoambiental (Reinhard & Bryant, 1992;Carrión et al., 2001Carrión et al., , 2004Horrocks et al., 2003Horrocks et al., , 2008Velázquez et al., 2010;Velázquez & Burry, 2012;Wood et al., 2012). Varios autores señalan que un punto importante en el estudio de coprolitos es dilucidar la procedencia polínica (Alcover et al., 1999;Carrión et al., 2004;Fernández-Jalvo et al., 2010). ...
... de los coprolitos también a la reconstrucción paleoambiental (Reinhard & Bryant, 1992;Carrión et al., 2001Carrión et al., , 2004Horrocks et al., 2003Horrocks et al., , 2008Velázquez et al., 2010;Velázquez & Burry, 2012;Wood et al., 2012). Varios autores señalan que un punto importante en el estudio de coprolitos es dilucidar la procedencia polínica (Alcover et al., 1999;Carrión et al., 2004;Fernández-Jalvo et al., 2010). Por ejemplo, los granos de polen encontrados en los coprolitos de herbívoros y omnívoros tienen relación con su dieta y con la disponibilidad de las plantas en el ambiente, mientras que en carnívoros y también en omnívoros, se relacionan fundamentalmente a los granos que pudieran estar contenidos en el tracto digestivo de las presas consumidas (Carrión et al., 2001). ...
p> Introducción y objetivos: Una de las problemáticas de la copropalinología para estudios de paleodieta es identificar la procedencia del polen que podría deberse a la contaminación polínica del alimento, al beber agua con polen en suspensión, o a contaminación postdepositacional. El objetivo de este trabajo es analizar el espectro polínico de la superficie de tallos y hojas de Empetrum rubrum, Senecio filaginoides, Nardophyllum obtusifolium y Mulinum spinosum, que conforman la dieta de Lama guanicoe del Parque Nacional Perito Moreno, para evaluar la contaminación polínica del alimento.
M&M: Se recolectaron tallos y hojas de cada una de estas especies en el valle del río Roble. Hojas y tallos de cada ejemplar se observaron al microscopio electrónico de barrido y se realizó acetólisis y el recuento polínico bajo microscopio óptico.
Resultados: Los resultados evidencian que los tipos polínicos dominantes, salvo en las de N. obtusifolium, proceden de la misma especie. Sin embargo, es importante la presencia de otros tipos polínicos como Nothofagus, Nassauvia, Podocarpus, Poaceae y Rumex sobre la superficie de las plantas.
Conclusiones: Los resultados de este estudio se relacionan con las características epidérmicas de las plantas y aportan información referente a la procedencia del polen en los coprolitos de Lama guanicoe, permitiendo precisar la información para reconstruir paleodietas.</p
... They result from elements consumed by the animals, or from elements attached to the feces surface subsequent to their deposition. Analysis of these inclusions reveals some aspects of paleodiets, paleopharmacology, paleoenvironment and the times in which the beings that deposited the feces occupied the sites (Callen and Cameron, 1960;Reinhard et al., 1985;Reinhard et al., 1991;Reinhard and Bryant, 1992;Carrión et al., 2004;Chaves and Reinhard, 2006;Reinhard et al., 2006;Rivera et al., 2014;Velázquez et al., 2010Velázquez et al., , 2015Wang et al., 2015). ...
The analysis of coprolite inclusions is instrumental in exploring certain features of paleodiets and paleoenvironments, and in determining the time when the sites were occupied by those who deposited the feces. Plant remains found in palaeofeces can be recognized by means of optical microscopes and through microhistological techniques. The aim of this study is to perform a microhistological analysis of plant remains in probable human coprolites from the Cerro Casa de Piedra, Cueva 7 site, province of Santa Cruz, Argentina, with the purpose of identifying the plants included in the diet of the hunter - gatherers who lived in Patagonia during the Pleistocene - Holocene transition and the Holocene, of recognizing some of their cultural practices and of determining the seasonality of the shelters they used. Our microscope study of the samples revealed epidermal fragments of Ephedra sp. stem, leaf and fruit remains of Empetrum rubrum and Gaultheria mucronata, remains of vascular bundles of Azorella monantha leaves and of Armeria maritima epidermis. Only representatives of the Poaceae family were found among the monocots. Our microhistological study of coprolite provided evidence to endorse the consumption of fruits, namely of E. rubrum and G. mucronata. In addition, by determining the time of the year in which the mentioned species bear fruit, we were able to infer that humans used this cave in summer. Likewise, the identification of Ephedra sp., Armeria maritima and Azorella monantha also allowed us to infer that plants of these species were used as medicine and fuel.
... They result from elements consumed by the animals, or from elements attached to the feces surface subsequent to their deposition. Analysis of these inclusions reveals some aspects of paleodiets, paleopharmacology, paleoenvironment and the times in which the beings that deposited the feces occupied the sites (Callen and Cameron, 1960;Reinhard et al., 1985;Reinhard et al., 1991;Reinhard and Bryant, 1992;Carrión et al., 2004;Chaves and Reinhard, 2006;Reinhard et al., 2006;Rivera et al., 2014;Velázquez et al., 2010Velázquez et al., , 2015Wang et al., 2015). ...
... The coprolites are deposited at CCP sites with attributes (low exposition to solar radiation, relatively constant temperature and humidity) that have permitted preservation of organic remains, such as bone remains, charcoal, fleece and plant litters (De Nigris, 2004), and even coprolites. Studies on palaeoenvironments, paleodiets, resource use seasonality and parasitism status of ancient populations can be performed by means of coprolite microfossil content analyses (Reinhard and Bryant, 1992;Horrocks et al., 2003;Carrión et al., 2004;. In particular, paleoparasitological, palynological and microhistological studies of coprolites from the PMNP caves have been done (Fugassa, 2007;Fugassa et al., 2008;Taglioretti, 2008;Sardella et al., 2010;Yagueddú and Arriaga, 2010). ...
The aim of the present study was to develop a seasonality palynological model from modern feces of Lama guanicoe for the interpretation of pollen found in coprolites. For this purpose, fecal samples of L. guanicoe were collected during a year from dung piles located in the ecotone Subantarctic Forest–Patagonian Steppe, Perito Moreno National Park (PMNP) (NW Santa Cruz province, Argentina). For every season, the food items were characterized and the post-depositional pollen contamination of feces was evaluated. The results showed differences among the pollen sets of feces from every season and no post-depositional pollen contamination of the superficial fecal subsamples. Pollen concentration and percentage values evidenced high pollen abundance of: a) Asteraceae subfam. Asteroideae-Senecio type, Mulinum spinosum and Rumex in summer feces; b) Poaceae in autumn feces; c) Empetrum rubrum in winter feces; and d) Nassauvia, Plantago and Poaceae in spring feces. Seasonality variations in the feces were related to the phenology of plants, to variations in the diet and to the supply of palatable resources for L. guanicoe. This seasonal feces model will be useful as an analog in studies on Holocene camelid coprolites found in Cerro Casa de Piedra (PMNP) sites. It will allow clarifying the seasonality in site use. On the other hand, it will contribute data to the L. guanicoe's diet, which represent an important information for the species' regional sustainable management.
... The coprolites are deposited at CCP sites with attributes (low exposition to solar radiation, relatively constant temperature and humidity) that have permitted preservation of organic remains, such as bone remains, charcoal, fleece and plant litters (De Nigris, 2004), and even coprolites. Studies on palaeoenvironments, paleodiets, resource use seasonality and parasitism status of ancient populations can be performed by means of coprolite microfossil content analyses (Reinhard and Bryant, 1992;Horrocks et al., 2003;Carrión et al., 2004;. In particular, paleoparasitological, palynological and microhistological studies of coprolites from the PMNP caves have been done (Fugassa, 2007;Fugassa et al., 2008;Taglioretti, 2008;Sardella et al., 2010;Yagueddú and Arriaga, 2010). ...
The aim of the present study was to develop a seasonality palynological model from modern feces of Lama guanicoe for the interpretation of pollen found in coprolites. For this purpose, fecal samples of L. guanicoe were collected during a year from dung piles located in the ecotone Subantarctic Forest–Patagonian Steppe, Perito Moreno National Park (PMNP) (NW Santa Cruz province, Argentina). For every season, the food items were characterized and the post-depositional pollen contamination of feces was evaluated. The results showed differences among the pollen sets of feces from every season and no post-depositional pollen contamination of the superficial fecal subsamples. Pollen concentration and percentage values evidenced high pollen abundance of: a) Asteraceae subfam. Asteroideae-Senecio type, Mulinum spinosum and Rumex in summer feces; b) Poaceae in autumn feces; c) Empetrum rubrum in winter feces; and d) Nassauvia, Plantago and Poaceae in spring feces. Seasonality variations in the feces were related to the phenology of plants, to variations in the diet and to the supply of palatable resources for L. guanicoe. This seasonal feces model will be useful as an analog in studies on Holocene camelid coprolites found in Cerro Casa de Piedra (PMNP) sites. It will allow clarifying the seasonality in site use. On the other hand, it will contribute data to the L. guanicoe's diet, which represent an important information for the species' regional sustainable management.
... lakes sediments, peat or bogs. Particularly, coprolite pollen analyses give evidence about diet, seasonality and paleoenvironment (Carrión et al. , 2004(Carrión et al. , 2005Carrión 2002;Chaves and Reinhard 2006;Horrocks et al. 2002Horrocks et al. , 2003Horrocks et al. , 2008Reinhard and Bryant 1992;Riley 2008;Velázquez et al. 2010Velázquez et al. , 2014Velázquez et al. , 2015Velázquez and Burry 2012;Wood et al. 2012;among others). ...
... Pollen analyses of coprolites have been studied to infer paleodiet, home range and paleopharmacology. Relative to these topics, the value of coprolites to perform paleoenvironmental inferences has been neglected (Carrión et al. , 2004. However, research on coprolites is less common for paleoenvironmental reconstructions than analyses of lake deposits and peat bogs in part because it is difficult to find coprolites in a good state of preservation. ...
... The comparison between pollen spectrum of coprolites and contemporaneous sediments has shown that coprolites reflect a good representation of local and regional flora. In addition, coprolites have shown higher pollen diversity and abundance of entomophilous taxa than sediments (Carrión 2002;Carrión et al. 2004). ...
The study of coprolites permits to deal with ecological and cultural aspects of past populations, such as palaeoenvironmental reconstructions, palaeodiets, inferences related to the use of resources and parasitism status. The objective of this work is the recognition of the contribution of studies of pollen in coprolites to the palaeoenvironmental reconstructions, and the identification of probable items in the diet of guanacos living during the Holocene in the area near the site Cerro Casa de Piedra 7 (Perito Moreno National Park, Santa Cruz, Argentina). The pollen content of 15 coprolites of guanacos coming from seven archaeological layers dated by 14C and resulting in ages between 9.640 ± 190 years BP and 5.400 ± 64 years BP were analysed. Pollen extraction was performed through acetolysis, and determination and counting under optical microscope. The presence of plant tissue and parasite remains was recorded. The coprolite pollen spectrum was compared with the sediment spectrum, and statistical analyses were conducted. The dominant pollen types were: Nothofagus, Empetrum rubrum, Asteraceae subfam. Asteroideae and Nassauvia, Poaceae and Caryophyllaceae. Pollen sets of contemporaneous coprolites and sediments show similarity in some cases, and different ratio or composition in some others. The similarities found between coprolites and sediments allow to infer that pollen in coprolites reveals the dominant vegetation taxa; on the contrary, a greater abundance of certain pollen types in coprolites than in sediments could be due to the fact that pollen in coprolites is biased with over-representations of the items preferred by guanacos, whereas a greater abundance in sediments than in coprolites could be due to the fact that the ingested plants were not in their flowering period. The results of this work show the value of coprolites as palaeoenvironmental indicators, and it contributes data to the palaeoenvironmental reconstruction and to the guanacos` diet.
