Abstract
We present the results of an interdisciplinary study conducted at the Pedernales-5 site, situated in the Salar de Pedernales basin (26°S; 3356 masl), dated from 10,510−10,749 to 11,201−11,612 cal. BP. Unlike other Late Pleistocene and Holocene sites in the Andean Puna, where vicuñas (Vicugna vicugna) were the primary prey, at Pedernales-5, the zooarcheological assemblage consists almost exclusively of flamingo (Phoenicopteridae) and bird remains. Through the analysis of lithic and bone artifacts, archeobotanical remains, pigment composition, and paleoenvironmental data, we propose that the emphasis on flamingo exploitation cannot be solely explained by alimentary consumption but also served cultural and symbolic purposes related to the acquisition of feathers, hides, and bones. The unique archeological context of Pedernales-5 offers a distinctive perspective on human dynamics in the Andean highlands during the Early Holocene.
Introduction
In recent years, the investigation of several archeological sites in the highlands of the Chilean southern Andean Puna has facilitated the reconstruction of an occupational sequence spanning from 12,500 cal. BP to the recent historical periods (López Mendoza et al., 2023a; Mendoza et al., 2023b). Throughout these contexts, vicuña (Vicugna vicugna) remains predominate, aligning with the prevailing pattern observed in most archeological sites across the Andean Puna. However, the case of Pedernales-5 (hereinafter PE-5) diverges from this trend. Situated as a small rock shelter near the Quebrada de Pedernales outflow (26°S-69°W; López Mendoza et al., 2023a), PE-5 yields osteofaunal evidence primarily comprising flamingo (Phoenicopteridae) and birds (Aves), and to a lesser extent, vicuña, puma (Puma concolor), lizards (Liolaemus sp.), and rodents.
Why flamingos rather than vicuñas? According to economic utility analysis, flamingos offer a reduced income of fat and meat compared to vicuñas, a crucial consideration in the challenging environment of the Andean Puna. With substantial temperature fluctuations both daily and seasonally, a considerable daily caloric intake is necessary for survival. Pre-Columbian Andean societies capitalized on the abundant meat supply of vicuñas and their high-quality skins, rendering them a principal prey species. This preference is evident throughout the chronological and cultural sequence of the southern Puna, where vicuñas dominate the zooarcheological record alongside guanacos (Lama guanicoe) (Cartajena et al., 2005; López et al., 2021b, 2022; López Mendoza et al., 2021a, 2023a; Mendoza et al., 2023b).
After the Darwin’s rhea (Rhea pennata) and the condor (Vultur gryphus), the flamingo boasts the third-largest body size among the numerous bird species inhabiting Andean salt flats. These birds are characterized by their colorful plumage, and their gregariousness makes them an easy prey to capture. Moreover, their feathers, eggs, and bones remain highly valued resources to date serving diverse economic, technological, medicinal, and ritual purposes. Particularly, their feathers play an integral role in the enduring rites and traditions of the central-southern Andes, as well as other regions of South America and beyond (Bárcena, 2017; Cartajena et al., 2005, 2010; Hesse, 1984; Urquiza and Aschero, 2014; Urquiza and Echevarria, 2018; Velardez-Fresia, 2023).
The archeological assemblages of PE-5 present a unique record, diverging from the predominant trends observed in both the study area and other archeological regions within the Andean Puna. Additionally, it prompts intriguing inquiries into the early phases of highland occupation. In this study, we analyze various lines of evidence, including lithic remains, bone industry, archeobotanical assemblages, faunal remains, and mineral pigments recovered from recent excavations at the Pedernales-5 site. Our aim is to construct a parsimonious interpretation of this distinctive archeological context, shedding light on the human dynamics of the southern Puna region during the Pleistocene-Holocene transition. This region has historically received limited scholarly attention and is presently threatened by extractive activities.
Study area and environmental background
The Salar de Pedernales (26°13′59″S-69°07′16″W) is located at an average altitude of 3,356 masl within the Atacama Region of Chile, covering an area of 3620 km2 (Figure 1). The prevailing climate is characterized by a cold mountain desert climate, with rainfall primarily concentrated during January and February, influenced by the South American Summer Monsoon. Throughout the rest of the year, rainfall is scarce, averaging 150–200 mm annually, with temperatures maintaining an average of 4°C. Only a few plant species have adapted to this arid environment, with grasslands and high-altitude wetland species predominating in the zonal vegetation. The fauna includes species such as guanacos, vicuñas, foxes (Lycalopex culpaeus and Lycalopex griseus), pumas (Puma concolor), rodents (Ctenomys spp., and Lagidium viscacia), and lizards (Liolaemus spp., and Callopistes palluma).

(a) Location of the study area; (b) location of the PE-5 site; (c) PE-5 rock shelter section view; and (d) view of the site and the ravine to the west.
Among the avifauna, flamingos such as Phoenicoparrus andinus, Phoenicoparrus jamesi, and Phoenicopterus chilensis (the focus of this article) seasonally nest on the salt flat. Additionally, various bird species inhabit the area, including Wilson’s Phalarope (Phalaropus tricolor), Andean Gull (Chroicocephalus serranus), Puna Plover (Charadrius alticola), Ochre-naped Ground-Tyrant (Muscisaxicola flavinucha), Andean Condor (Vultur gryphus), Ornate Tinamou (Nothoprocta ornata), Baird’s Sandpiper (Calidris bairdii), various species of Ducks (Anas spp.), Diademed Sandpiper-Plover (Phegornis mitchellii), Mountain Caracara (Phalcoboenus megalopterus), Hooded Grebe (Podiceps gallardoi), Black-capped Hemispingus (Kleinothraupis atropileus), Rufous-bellied Seedsnipe (Attagis gayi), Aplomado Falcon (Falco femoralis), Crested Duck (Lophonetta specularioides), Horned Coot (Fulica cornuta), and Barn Swallow (Hirundo rustica), among others (Martínez and González, 2017).
The Pedernales-5 site
The PE-5 site is situated in the Quebrada de Pedernales, which flows into the southwest sector of the Salar de Pedernales basin. Several archeological sites, characterized by surface evidence predominantly comprising lithic and ceramic scatters, are concentrated within a Quaternary fluvial deposit bed (López Mendoza et al., 2023a; Mendoza et al., 2023b). Some of these sites have been excavated, providing insight into the early occupation of Quebrada Pedernales. PE-38 contained multiple occupational events dating from 9733−10,158 to 2349−2,699 cal. BP associated with projectile point production and camelid hunting near a rock shelter on an ignimbrite outcrop. The PE-1 site, adjacent to the ignimbrite outcrop, comprises successive discrete occupational events involving the consumption of vicuñas, egg collection, and lithic knapping, dated from 2357−2705 to 552 to 653cal. BP (López et al., 2021b; López Mendoza et al., 2021a).
PE-5, situated on the same ignimbrite outcrop as PE-38, comprises an occupation adjacent to a rock shelter with a small interior space measuring 2 m from the back to the drip line, serving as a shelter. This area was filled with wind-blown and colluvial sediments. Eight stratigraphic units (SUs) were distinguished during the excavation (Figure 2a). SU-1 through SU-7 primarily originated from a combination of aeolian action and sediment input from colluvium located in the upper part of the ignimbrite wall. Throughout the sequence, silty sands ranging from fine to coarse, occasionally containing fine gravel with moderate to poor sorting, and exhibiting a high frequency of quartz and fragments of igneous and volcanic origin (lavas and ignimbrite) were observed. Differences between each SU are attributed to variations in sand grain size and the color of dry deposits. SU-8 and SU-9 show a higher frequency of ignimbrite fragments, accompanied by increasing compaction of the matrix sediment as depth progresses, ultimately reaching the base of the Miocene ignimbrite outcrop (Mendoza et al., 2023b).

(a) Stratigraphic profiles of the PE-5 site; (b) distribution of artifacts and biofacts, including 14C-dated samples; (c) PE-5 site plan; and (d) Early Holocene occupational floor.
Human occupation is concentrated below a depth of 35 cm (levels 7–12; SU-4 and SU-5), surrounding several extended hearths (Features 2, 3, and 4, Figure 2b). The obtained dates indicate an initial occupation at 11,201−11,612 cal. BP, followed by another dated to 10,501–10,749cal. BP (López Mendoza et al., 2023a). These represent some of the earliest human occupational events in the area to date, preceded only by the Infieles-1 site, with an initial occupation dating between 11,842 and 12,429 cal. BP (Mendoza et al., 2023b). However, a Vicugna vicugna bone sample taken from the excavation level 1 (0–5 cm) yielded a date of 10,880–11,241cal. BP (Table 1), attributed to the actions of fossorial rodents inducing vertical artifact and ecofact displacement. Fossorial rodent activity extended to the base of SU-3, with no such disturbance observed in the higher concentration of artifactual or ecofactual evidence below this stratigraphic unit (Figure 2c and d).
Radiocarbon dates of the PE-5 site. Abbreviations: SU. Stratigraphic Unit. Dates were calibrated with the OxCal 4.4.4 software, Interface Build 170 (Bronk Ramsey, 2021), using the SHCal20 curve (Hogg et al., 2020).
Flamingos
Flamingos inhabit a range of environments, from sea level to approximately 4500 masl, typically found in lagoons and shallow waters. They form nesting colonies and undertake extensive travels between feeding and breeding areas (Caziani et al., 2007). Among the species found in Pedernales, the Andean flamingo (Phoenicoparrus andinus) stands out, reaching a maximum height of 136 cm. Its plumage is predominantly whitish with vibrant pink hues, while its tail feathers are black, and its upper chest displays violet coloration. Breeding primarily occurs during the summer months, with preferred nesting areas being the salt flats on the western slope of the Andes, situated below 3000 masl. Typically, the Andean flamingo lays only one egg, although in rare instances it may lay two (Parada et al., 1990; Valqui et al., 2000).
The Chilean flamingo (Phoenicopterus chilensis) is an endemic South American bird, ranging from southern Brazil to Patagonia. It inhabits a variety of water environments, from highlands to marine wetlands (Araya and Millie, 2005). With a height ranging from 110 to 130 cm, it features light-pink plumage with darker patches near the tail, and a distinctive curved beak that transitions from light to dark coloring along its length. Lastly, James’s flamingo (Phoenicoparrus jamesi) is a gregarious species found in saltwater environments. It also nests in colonies and undertakes extensive migrations for feeding and breeding (Caziani et al., 2007). James’s flamingo is the smallest of the three species, with an average height of 90–92 cm, a wingspan ranging from 100 to 160 cm, and a weight of approximately 2 kg. It is characterized by its red legs and scarlet coloration at the base of the neck and back, with black coloring on the posterior third of the body. Its neck, legs, and beak are shorter compared to other species, with a less curved beak that features an orange-yellow color transitioning to a sharp black apex. Reproduction among all three species occurs across seven stages, encompassing various events from pair formation to chick dispersal throughout the annual cycle (Parada et al., 1990). The detailed chronology of these events is illustrated in Figure 3.

(a) Dates of pair formation until dispersal of the species Phoenicoparrus andinus, Phoenicoparrus jamesi, and Phoenicopterus chilensis (figure taken from Rodríguez, 2006); (b) illustrations of the species (1) P. andinus, (2) P. jamesi, and (3) P. chilensis (illustrations by Patricio López Mendoza); and (c) view of Phoenicoparrus jamesi colony in Pedernales (photograph taken by Sergio Arancibia).
Material and methods
A total of 2071 bone specimens were recovered from PE-5. Phoenicopterus chilensis collections were referenced to identify Phoenicopteridae samples. However, classification at the genus and species level was not feasible due to limited osteological study of these species. As a proxy, the osteometric record from Cartajena et al. (2010) was utilized, and quantification was conducted using the Number of Identified Specimens (NISP). Taxonomic abundance analysis relied on the Minimum Number of Individuals (MNI). Anatomical unit frequency was determined by the Minimum Number of Skeletal Elements (MNE), Minimum Number of Animal Units (MAU), and %MAU. Immature and adult individuals were differentiated based on cortical tissue ossification degree (Lefèvre and Pasquet, 1994).
Richness, Diversity, and Evenness indices were calculated to compare sites across the cultural sequence. Richness (NTaxa) reflects the number of used taxa in a site, calculated as the sum of non-overlapping taxonomic categories. The Diversity Index (H’) assesses species relative importance and variety, using the Shannon Index formula H’ = −∑ (pi) (ln pi), where pi is the number of specimens of category i divided by the sample size. The Evenness Index (J’) gages abundance distribution and species variety, based on equal/unequal species use and resource exploitation dependence. Values range from 0 to 1, where values closer to 1 imply taxa are equally abundant. J’ = H’/ln S calculates this index, with H’ as the Shannon Index and S as the number of taxa (NTaxa). For the latter two indices, MNI was multiplied by each living taxon’s average weight to estimate biomass from animal resources (Del Papa et al., 2011). Mammalia, Rodentia, and Aves taxonomic categories were excluded due to quantification challenges with fragments of varying sizes. Taphonomic analyses focused on weathering indexes, root, rodent, or carnivore marks on bones, and chemical alterations or sedimentological particle movement.
X-Ray Fluorescence (XRF) analysis was conducted on three samples of mineral pigment. The first corresponds to a rock fragment from level 10 with red pigment adhered to its surface. The second sample is a fragment of a bone fragment from level 11 (feature 1), entirely covered with red pigment. The third sample is a fragment of red mineral pigment found in the deflated zone of the site, 2 m east of Excavation Unit 1. Energy Dispersive X-Ray Fluorescence analyses were performed at the Electron Microscopy Laboratory of the Universidad de Santiago de Chile, with three points analyzed for each sample. The lithic study focused on the evidence recovered from levels 7 to 12 (35–60 cm depth). The analysis encompassed the classification of the assemblage according to dimensional, technological (class and type), presence of cortex and raw material criteria (Andrefsky, 1998).
Results
Zooarcheological study
The zooarcheological evidence from PE-5 is summarized in Table 2. The highest concentration of faunal remains was found in the Early Holocene occupational floor, spanning levels 10, 11, 12, and 19. The few vicuña remains identified (MNI = 2) are primarily represented by skull, mandible, radius, ulna, femur, and calcaneus, with a generally high degree of fragmentation (Supplemental Material 1). The Aves category mainly consists of long bone diaphyseal fragments smaller than 5 cm (Supplemental Material 2). The sole taxonomic category identified within the bird group corresponds to Phoenicopteridae; thus, it is probable that the majority of the evidence in the Aves category pertains to this taxonomic group. Measurements taken from adult Phoenicopteridae individuals align with the size of Phoenicopterus chilensis. However, as depicted in Figure 4, variations in the measurements of the proximal femur and coracoid are observed between the analyzed samples and the reference collections used. Nevertheless, variability in body size across species does not necessarily indicate intraspecific differences. Environmental factors related to carrying capacity may also influence variation in body size (Silva-Martínez et al., 2023), such as the increased productivity and humidity during the Pleistocene-Holocene transition. Therefore, the osteometric data for PE-5 (Supplemental Material 3 and 4) suggests that the measured adult samples correspond to large Phoenicopteridae specimens, considering the size gradient discussed by Cartajena et al. (2010).
Taxonomic frequency (NISP) of the PE-5 site distributed by excavation levels and stratigraphic units. Samples with adhered pigment are detailed.

(a) Measurements of proximal femur; and of (b) coracoids from Phoenicopteridae samples from PE-5, compared to specimens of Phoenicopterus chilensis and Phoenicoparrus jamesi (measurements taken from Cartajena et al., 2010). Dp: Greatest depth of the proximal end; Bp: Greatest breadth of the proximal end: Caput femoris-most lateral point of the Trochanter major; GL: Greatest (diagonal) length; and BF: Breadth of the facies articularis basalis (Von Den Driesch, 1976).
Flamingos represent the most abundant species within the archeofaunal assemblage, comprising 12 individuals. The majority are adults, or individuals nearing full maturity within a few weeks or months. Consequently, the frequency of anatomical units was recorded by considering all remains within a single assemblage (Supplemental Material 5). The most prevalent Phoenicopteridae remains include distal epiphyses of the tarsometatarsi, proximal epiphyses of the femora, distal tibiotarsi, second phalanges, quadrates, sternum fragments, skull fragments, scapulae, and coracoids. Missing bones include the synsacrum, carpometacarpus, and nail phalanges (Figure 5a–k). The incidence of Aves bones exhibiting signs of exposure to fire is minimal, with only 14 such remains identified in this category, along with one Phoenicopteridae bone (Table 3).

Phoenicopteridae remains recovered from the PE-5 site: (a) coracoid; (b) skull fragment; (c) phalanges; (d) cervical vertebrae; (e) quadrate; (f) distal condyle of tarsometatarsus; (g) proximal epiphyses of femora; (h) distal (left) and proximal (right) epiphyses of tibiotarsi; (i) diaphysis and distal epiphyses of humeri; (j) eggshell fragment; and (k) %MAU representation of Phoenicopteridae remains from PE-5.
Frequencies of bone remains exposed to fire (expressed in NISP) according to taxonomic category.
Among the remains, a possible fragment of a bead made from a long bone of an Aves was identified. Obtaining this piece involved the segmentation of the diaphysis by sawing the two ends, as evidence by the transverse grooves. The final detachment of the piece was carried out by bending, resulting in a distinctive tongue fracture plane (languette) (Figure 6a).

(a) Diaphysis of a long bone of Aves with sawing grooves at its ends; (b) present-day pellets collected at Quebrada de Pedernales.
The occurrence of taphonomic agents at the site is minimal, with only a few specimens displaying weathering and scarce evidence of carnivore marks. Manganese imprints on the bone surface are highly prevalent, likely attributable to the characteristics of the rock shelter, which retain water and snow almost year-round. Despite this, phenomena such as stratigraphic deposit flooding and subsequent bird bone flotation (Mameli, 2003: 210), as observed in excavated sites in Pedernales, Infieles, and Maricunga, have not been documented. The occupational floors have primarily been affected by fossorial rodents. The presence of taxa such as Liolaemus sp., cricetids, and passerines may be linked to predatory or raptor birds, consistent with the discovery of contemporary pellets at the ignimbrite wall base, containing mainly lizard, cricetid, and passerine remains (Figure 6b). Based on this evidence, the impact of natural agents on the accumulation and degradation of Phoenicopteridae bone remains is minimal, suggesting their presence is predominantly due to human activity.
The predominance of flamingos in PE-5 undoubtedly marks a distinctive pattern. Throughout the entire sequence from the Late Pleistocene to the Late-Holocene (refer to radiocarbon dates in Supplemental Material 6), vicuñas emerge as the most prevalent species in the zooarcheological record of the area (Table 4; Supplemental Material 7). Richness (NTaxa), Diversity (H’), and Evenness (J’) indices across these sites exhibit variations within the same period (Figure 7), which can be attributed to factors such as site functionality, seasonality, and the intensity of occupations.
Calculation of Minimum Number of Individuals (MNI) for each site and divided by periods. The weight is detailed for each taxa. LP: Late Pleistocene; EH: Early Holocene; MH: Middle Holocene; LH: Late-Holocene. *Weight calculated from the average between Lama guanicoe and Vicugna vicugna.

Graphic representation of the Richness (NTaxa), Diversity (H’), and Evenness (J’) indices of the sites studied to date in Infieles, Pedernales, and Maricunga. The shadows indicate the taxa recorded at each site, where the most abundant (MNI multiplied by the weight of each animal) are found at the top, and the least abundant at the bottom. LP: Late Pleistocene; EH: Early Holocene; MH: Middle Holocene; LH: Late-Holocene; Vi: Vicugna vicugna; Ph: Phoenicopteridae; Ch: Chinchillidae; Pa: Passeriforme; Ca: Camelidae; Pu: Puma concolor; Li: Liolaemus sp.; Cr: Cricetidae; La: Lama sp.; Ly: Lycalopex culpaeus; and Lv: Lagidium viscacia.
Pigment
Regarding the pigment analysis, traces of iron oxide were identified in all three samples (Supplemental Material 8, 9, and 10). This oxide is abundant in eroded strata of the Domeyko Mountain Range (Supplemental Material 11), located 7 km west of the site, although it also appears in specific areas within the Pedernales Ravine. The clast sample also exhibits the peculiarity of containing a quantity of exogenous phosphorus (absent in the natural rock), which may be explained by the dissolution of bone contents and/or by bone grinding together with the pigment as a means of binding the iron oxide dust.
Lithic assemblage
The lithic assemblage recovered from PE-5 is relatively small, totaling 56 pieces exclusively from levels 7 to 12 (35–60 cm), with the majority concentrated in level 10 (n = 24; SU-4, SU-5, features 2–4). The assemblage is primarily composed of siliceous rocks (n = 41, 73.21%), which are abundant raw materials found in the ravine in the form of nodules (Figure 8c, Soto, 2023), with smaller proportions of andesite (n = 10, 17.86%), granitic rocks (n = 2, 3.57%), and quartz crystal (n = 1; 1.79%) among others. Most artifacts consist of non-diagnostic knapping debris, such as flakes and flake fragments (n = 40). Additionally, core flakes, mainly of andesite (n = 13), one resharpening flake of a unifacial retouched tool (n = 1), and a bifacial fragment in siliceous rocks (n = 1), suggests the production and use of retouched tools at the site (Table 5). Furthermore, the lithic assemblage includes a pebble (>11 cm maximum length) with red pigment from feature 1 (hearth) and a large tabular block from SU-3, both displaying surfaces impregnated with red pigment (Figure 8a and b). Approximately 37.5% of the lithic pieces exhibit remnants of cortex.
Distribution of lithic evidence recovered from the excavations of PE-5 according to excavation level and Stratigraphic Unit.

(a) Tabular block recovered with evidence of red pigment on its exposed face; (b) subrounded fragment with red pigment (feature-1); (c and d) non-diagnostic flakes; (e) bifacial fragment.
Archaeobotanical record
In relation to the macrobotanical record, 40% (n = 80) are charred and 60% (n = 122) are non-charred remains. Among the latter, the majority corresponds to indeterminate fragments (Indet-1), which represent close to 27% of the total. Regarding taxonomic identification within the charred assemblage, 24% (n = 19) of the pieces were identified, 23% (n = 18) were non-determined, and 54% (n = 43) were indeterminable. Within the non-charred assemblage, 42% (n = 51) could be determined and 58% (n = 71) could not be determined. As a result, 12 of the taxa detailed in Table 6 were identified.
Taxonomic determination of PE-5 macrobotanical remains. (*) Charred samples.
Discussion
The role of the flamingo in the economic and technical system
We could tentatively dismiss the possibility that bird exploitation played a more predominant role in the human diet due to the decline in the population of other animal species. Paleoclimatic records do not indicate adverse conditions for vicuña proliferation. On the contrary, evidence suggests that during the Late Pleistocene and Early Holocene, the southern Puna experienced higher rainfall than present-day conditions (Grosjean, 2001; Lamy et al., 1999; Tiner et al., 2018). Further north (22°S), indicators from rodent deposits suggest significantly wetter conditions during the Central Andean Pluvial Event II compared to the present (de Porras et al., 2017; López Mendoza et al., 2023a; Mendoza et al., 2023b). Climatic phenomena such as snowstorms may cause mass vicuña mortality, particularly in ravine habitats rather than highland plains and wetlands. However, despite the mass nature of these mortalities, they only account for 4.5% of the total census population (Cajal and Ojeda, 1994).
Given the prevalent long-term exploitation of wild camelids in the region, the prominence of birds in Pedernales-5 poses an intriguing question. For the southern salt flats, the feather record is extremely limited; however, feather headdresses may have been depicted in rock paintings (González Godoy et al., 2023). This could indicate ornamental use. It is also possible to discuss potential technological uses. In the lithic assemblages of PE-38, large, stemmed points measuring 11–13 cm were recovered (Loyola et al., 2023). These points feature elongated triangular blades, barbs, convergent stems, and convex bases, suggesting their use as spear-throwers or thrusting spears due to their size and weight. Conversely, for Late-Holocene occupations, both surface and stratigraphic deposits yielded smaller stemmed points measuring less than 5 cm, likely intended for use in bow and arrow systems. Feathers could have been utilized as stabilizers for both propelled and thrown weapons, with flamingo feathers, given their size and shape, being particularly suitable for this purpose (Urquiza and Echevarria, 2018). Regarding the bones, we believe that the diaphysis of the limbs, such as the humerus and tibiotarsus, would have been used as raw material; this is consistent with the discovery of a possible bead fragment. Similar short segments obtained by sawing bird long bones have been commonly described in other archeological sites in this and other regions, where they were interpreted as beads (Beretta and Zubimendi, 2017; Legoupil, 2003; Orquera and Piana, 1999; Schiappacasse and Niemeyer, 1964, 1965).
For now, it is difficult to precisely ascertain the role of the flamingo in the technological, economic, and symbolic system of the hunter-gatherer societies of Pedernales during the Early Holocene. However, the patterns observed in the archeological record seem to suggest that the capture of flamingos did not solely serve subsistence purposes.
The capture, transportation, and processing of flamingos at PE-5
The presence of nearly intact birds in the archeological record of Pedernales-5 suggests that they were transported fully intact, with adult-sized prey prioritized, even including anatomical units with no economic value. The short distance to the actual flamingo nesting areas (currently 500–700 m from the site) was certainly a key factor. The scarcity of bones such as carpometacarpi and anterior phalanges may be related to the extraction of relatively complete skins in the wing area, where these bones were not removed due to their anatomical location (see discussion in Blasco et al., 2019; Bovy, 2002, 2012). The absence of cut marks may be due to a removal practice similar to that used by taxidermists, where a cut is made along the chest to then manually remove the skin, making cuts in the skull area, where there is greater adherence (Penalba and Albe, 2023).
The lithic evidence indicates the maintenance and use of retouched tools at the site made from presumably local raw materials. Although the assemblage is not highly diagnostic, there are notable differences from other records from contemporary sites such as PE-38, where a bifacial industry intended for hunting wild camelids predominates (Loyola et al., 2024). Furthermore, the macrobotanical remains present at the site, mainly taxa with forage and fuel value (Aldunate et al., 1981), indicate that their collection coincided with the flamingo nesting season between December and April. This evidence, together with the remains of eggshells, is consistent with at least one summer occupation of the site but does not exclude longer-term occupations during the annual cycle. At PE-5, egg collection was apparently much less important than at PE-38 and other contexts in the Andes Puna (see Cartajena et al., 2010).
In some lithic (n = 2) and osteological (n = 7) remains, we recorded small spots of red pigment. In the case of the latter, the presence of pigment may be due to contact with oxide particles. However, in a fragment of mammal (possibly camelid) analyzed by XRF, all its faces show a red coloration (Supplemental Material 9). One plausible scenario is the grinding of pigment and marrow with bone fragments still attached. The use of marrow as a binder for mineral pigment has been ethnographically reported (Malainey, 2010) and in archeological evidence for neighboring areas (Sola et al., 2013). The presence of exogenous phosphates in the clast analyzed by XRF seems to point in the same direction.
Iron oxides contain ideal physio-chemical and antiseptic qualities for tanning hides (Rifkin, 2011) as well as decorative properties. Evidence in Northwestern Argentina indicates that pigments were used to paint leather, cloaks, pouches, and tobacco pouches (Sola et al., 2013). In the archeological locality of Río Doncellas (22°S-66°W), Casanova (1967) mentions red-painted leather and rattles, including a red-painted leather mask associated with a funerary context. At the Inca Cueva site (23°S-65°W), bags, headbands, leather thongs (Aguerre et al., 1973), and a vicuña leather with red paint in a stratigraphic unit dated to 10,620 ± 140 years BP (Aschero, 1984; Aschero and Podestá, 1986; Sola et al., 2013). The processing of vicuña hides with red pigment has solid evidence in Andean highland areas since the Early Holocene (Sola et al., 2013). However, in the case of birds, evidence tends to be scarcer and related to the use of feathers and bones (Velardez-Fresia and Urquiza, 2024).
In summary, the various lines of evidence lead to an interpretation of PE-5 as a specialized task site, mainly dedicated to the capture and processing of flamingos. This is consistent with ethnographic data of hunter-gatherers, which indicate that rock shelters and caves are mainly used for brief occupations and limited activities (Binford, 1978; Nicholson and Crane, 1991). The evident differences with other relatively contemporary sites could be related to a functional differentiation of space and the organization of hunting activities. This could have been influenced by the restrictive nature of the ignimbrite outcrops in the ravine, where the available rock shelters were greatly limited in space for overnight stays for more than three to four people due to their shallow depth and interior space. In this context, camps, specialized sites, sporadic shelters, hunting structures, and even ritual spaces were integrated into the highlands landscape.
Comparing the archeological and ethnographic record
It is evident from the evidence that the capture and processing of flamingos were central activities of the groups that occupied PE-5 during the Early Holocene. However, the means and objectives of flamingo exploitation are less clear. Bovy (2002) has proposed different archeological hypotheses regarding the use of flamingo wings, including processing and consumption, the production of tools and decorative objects using wings and feathers, and differential transportation, where undesired parts were discarded in middens regardless of the reason for bird capture. A later study suggested that the higher frequency of wing distal ends could be due to the transport of skins obtained from discarded post-consumption birds or because these segments were separated during bird cooking (Bovy, 2012). This pattern is not observed in cases such as those in Chilean Patagonia during the Mid and Late-Holocene, where entire birds were transported. The low frequency of bones such as the radius and ulna has been attributed to differential preservation, especially in smaller species like swallows. Additionally, in species such as cormorants and petrels, these bones tend to fracture more easily due to their thinner structure (Lefèvre and Laroulandie, 2014).
The techniques and means used to capture flamingos are challenging to infer due to the limited evidence of tools. This may be attributed to the fact that their capture likely involved simple techniques and equipment. Bird capture offers various advantages, including high visibility and species diversity, seasonal and spatial predictability, gregariousness, easy transportation of many individuals, and straightforward processing (Estevez and Mameli, 2004). Ethnographic data from Uru-Chipaya groups in the Bolivian highlands provide relevant insights into flamingo hunting. Among these groups, bird hunting, including flamingos, occurs during the cold season (may-september), when flamingos gather at night in saline water bodies. Traps, known as tckalkauni, are set up based on observations of flamingo flight paths and meeting places. Each tckalkauni consists of a system of aerial snares suspended from a 100 m-long rope, stretched about 2 m high with the help of several stakes. The rope is equipped with a 100 of these snares evenly spaced along its length (Figure 9a and b). Groups of three or four men install multiple tckalkauni during the new moon phase. Flamingos are ensnared, with 10 or more individuals often captured in this manner (Ríos and Rocha, 2002; Sáenz, 2006; Wachtel, 1985).

(a) Detail drawing of traps (tckalkauni) used by Uru-Chipayas groups to capture flamingos; (b) detail of the snares used in the traps (figures taken and modified from Sáenz, 2006); and (c) sequence for the use of liwi or squñi to capture flamingos among Uru-Chipayas groups.
According to Acosta (2008), Uru-Chipaya groups used instruments such as liwi or squñi (a three-cord boleadora made from sheep wool, llama fiber, and small stones, Figure 9c), chalalawa or llipi (possibly the tckalkauni), and juarul (lantern), for hunting Phoenicopterus chilensis. Flamingo chick hunting is infrequent and often unsuccessful due to the inaccessibility of their breeding areas, such as islands or muddy shores. Flamingo egg collection occurs in late january and february, providing a source of fatty acids and vitamins. However, evidence at PE-5 is limited to a few fragments (n = 85), compared to thousands of eggshells remains at neighboring site PE-38, suggesting that this was a secondary activity at PE-5 (López Mendoza et al., 2023a). The evidence suggests that skins, feathers, and bones were the main targets in flamingo capture at the site. According to ethno-historical data from the southern Puna of Argentina, fragments of feathers were utilized in shafts, headdresses, headbands, necklaces, loincloths, camouflage, and other items. Flamingo feathers, in particular, were employed in various rituals associated with rain and the earth (Urquiza and Echevarria, 2018; Velardez-Fresia and Urquiza, 2024). Additionally, flamingo feathers were burnt and used to treat hemorrhages and fevers (Wachtel, 1985).
Flamingo hunting also involves symbolic and ritual aspects. These birds are considered symbols of fertility and prosperity during the rainy season (Flores and Velárdez Fresia, 2018; Sigl and Mendoza, 2012), and their hunting holds complex symbolism for highland groups. Among the Uru-Chipaya, during hunting using tckalkauni, hunters perform certain rites prior to setting up snares and traps. One of them is the placement of red mineral powder in cups used for prayers prior to the hunting of flamingos (Wachtel, 1985). In the town of Peine, in the Antofagasta Region, flamingo feathers are offered on the surface of springs and water wells during ceremonies, symbolizing different members of the family (Cartajena et al., 2010). This ethnographic information highlights the various roles of flamingos in rituals, economy, technology, and medicine, as well as the simple techniques used in their capture. However, it should not be used as an argument by analogy, but rather as a reference point to discuss possible scenarios based on the archeological record.
Conclusion
Pedernales-5 presents a unique pattern among other Early Holocene occupations in the Andean Puna, with a notable predominance of flamingos in the zooarcheological record. There are no ecological, environmental, or taphonomic explanations for the low frequency of vicuñas compared to the high frequency of flamingos at PE-5. The occupation of the site aligns with periods of increased humidity, sustaining a biomass comparable to present-day conditions. Despite the ignimbrite outcrops being areas frequented by carnivores, the scarcity of fox or puma marks on the bones, along with their arrangement in clearly cultural floors, suggests minimal influence from natural agents. Consequently, cultural factors likely played a predominant role in shaping the osteofaunal record of the site.
The evidence provides a unique perspective on the economic, social, and symbolic dynamics of Early Holocene human groups. We suggest that the capture and processing of flamingos at Pedernales-5 served not only as a means of sustenance but also potentially involved the exploitation of their skins, feathers, and bones for various purposes. The use of pigment appears to be related to these tasks as well. The potential extraction and preparation of flamingo skins with adhered feathers at PE-5 is a scenario that warrants further exploration. Additionally, the technological equipment and hunting strategies associated with both the capture of these birds and the processing of their carcasses should be investigated using new lines of evidence and analytical approaches. This will enable a deeper understanding of the social organization of hunting and exploitation of these birds, as well as their multiple economic, technical, and symbolic dimensions for the past societies that occupied Quebrada de Pedernales during the Early Holocene.
Supplemental Material
sj-xlsx-1-hol-10.1177_09596836241254481 – Supplemental material for Pedernales-5: An enclave for flamingo (Phoenicopteridae) exploitation during the Early Holocene in the Chilean southern Puna (26°S-69°W)
Supplemental material, sj-xlsx-1-hol-10.1177_09596836241254481 for Pedernales-5: An enclave for flamingo (Phoenicopteridae) exploitation during the Early Holocene in the Chilean southern Puna (26°S-69°W) by Patricio López Mendoza, Rodrigo Loyola, Carlos Carrasco, Constanza Roa Solís, Daniel Varas, Francisca Santana-Sagredo, Víctor Méndez, Gabriela Bravo, Elvira Latorre, Alina Sáez, Vanessa Orrego and Ariel Sperling in The Holocene
Footnotes
Acknowledgements
We would like to express our gratitude to archeologist Catalina Soto for her invaluable assistance in interpreting the data. Additionally, we extend our thanks to archeologists Pablo Mariani, Bárbara Neumann, Josefina De La Barra, Diego Mayorga, and Francisca Vera for their support throughout the excavation process. Finally, we acknowledge Gladys Olivares from the Electron Microscopy Laboratory at the Universidad Santiago de Chile for her assistance in identifying the pigment samples.
Author contributions
Data availability statement
All specimens in this study are curated in the Museo de Historia Natural y Cultural del Desierto de Atacama, Calama, Chile. They are available prior consultation to the collection curator: Osvaldo Rojas,
Declaration of conflicting interests
The author(s) declared no potential conflicts of interest with respect to the research, authorship, and/or publication of this article.
Funding
The author(s) disclosed receipt of the following financial support for the research, authorship, and/or publication of this article: Research financed through Fondecyt Projects 1190197 and 1240193, ANID R20F0002 granted by the Agencia Nacional de Investigación y Desarrollo (ANID), Chile.
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References
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