Herpetofaunal Remains from Khonkho ... - Wiley Online Library

International Journal of Osteoarchaeology Int. J. Osteoarchaeol. 23: 475–484 (2013) Published online 11 July 2011 in Wiley Online Library (wileyonlinelibrary.com) DOI: 10.1002/oa.1257

SHORT REPORT

Herpetofaunal Remains from Khonkho Wankane, an Urban and Ceremonial Centre in the Southern Lake Titicaca Altiplano: Unique Behavioural Correlates to Osseous Taphonomy JAMES T. POKINES* Boston University School of Medicine, Department of Anatomy and Neurobiology, Boston, MA 02118, USA

ABSTRACT

This research reports the herpetological (reptile and amphibian) remains from Khonkho Wankane, an urban and ceremonial centre in the southern Lake Titicaca region of the Bolivian Altiplano, which fluoresced during the Late Formative period. The total n=1710, including a minimum of six taxa and representing a significant portion of the relatively depauperate modern herpetofauna recorded for the Lake Titicaca area. A lizard of the genus Liolaemus (Tropiduridae) was identified. Amphibians are represented by Andean toad Rhinella spinulosa (Bufonidae) and by at least four taxa of frogs: [cf. Gastrotheca marsupiata (Hylidae) and Pleurodema marmorata or P. cinerea and Telmatobius spp. (Leptodactylidae)]. The latter genus does not include members of the giant variants (T. culeus) that dwell in Lake Titicaca. The question of how these remains were incorporated into the site deposits is examined taphonomically. No indicators of human or small carnivore predation were detected. Significant spatial clustering of amphibian remains was detected, with two clusters contributing n=1562 or 92.4% of the amphibian total (n=1690). These concentrations also are characterised by a depressed ratio of cranial versus postcranial elements, which cannot be explained by recovery methods. These concentrations of herpetofaunal remains likely represent in-burrow deaths of species sheltering for thermoregulatory benefits away from this harsh Altiplano environment, with the lowered recovery of cranial elements hypothesised to have resulted from a lack of ossification. The bulk of herpetofaunal remains recovered therefore likely were recent intrusions into the site deposits and do not represent the residue of human exploitation during the occupation of Khonkho Wankane. Copyright © 2011 John Wiley & Sons, Ltd. Key words: Altiplano; Bolivia; herpetofauna; Khonkho Wankane; paleoecology; taphonomy; Tiwanaku

Introduction Khonkho Wankane emerged as an urban and ceremonial centre in the southern Lake Titicaca region of the Bolivian Altiplano during the Late Formative I period by 1 AD and included the construction of a massive rectangular mound and sunken temple and plaza complex by the Late Formative II period, ca 250 AD ( Janusek, 2008). This site was later eclipsed with the rise of Tiwanaku, the major urban polity of * Correspondence to: Boston University School of Medicine, Department of Anatomy and Neurobiology, 72 E. Concord St., L1004, Boston, MA 02118, USA. e-mail: [email protected]

Copyright © 2011 John Wiley & Sons, Ltd.

the southern Lake Titicaca region, by 500 AD. Some residential occupation of Khonkho Wankane continued during the Tiwanaku period. Tiwanaku flourished from around 500 to 1100 AD (Bermann, 1994; Janusek, 2003, 2008; Stanish, 2003). The paleoecological adaptations of this major political and ceremonial centre and its subsidiary sites to this relatively harsh (ca 70cm annual rainfall), high-altitude (ca 3800m) environment included the creation of raised agricultural fields along the lake shore. These consisted of alternating rows of raised earth and small, linked canals. Extending for several square kilometres, this artificial system served to ameliorate the microenvironment around the agricultural plots to increase output and reliability Received 19 January 2011 Revised 7 May 2011 Accepted 19 May 2011

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476 of harvest, especially reducing frost damage and improving the moisture supply and fertilisation of crops (Argollo et al., 1996; Binford & Kolata, 1996; Binford et al., 1997). Alteration of landscape also included the formation of qochas, or artificial reservoirs or ponds to store rainwater (Janusek, 2008). These artificial systems therefore created or altered a large system of wetlands around Lake Titicaca, which no doubt had an effect upon the many vertebrate species inhabiting the region. Dietary resources included lake fish and birds, herded camelids (Webster & Janusek, 2003), cuy (guinea pigs) and other vertebrate taxa utilised throughout the Andes in antiquity (Marcus et al., 1999; Miller & Burger, 1995; Pérez Arias, 2007). Economic activities also included agriculture of quinoa and potatoes (Bermann, 1994; Janusek, 2008; Pérez Arias, 2007; Stanish, 2003; Webster & Janusek, 2003). Khonkho Wankane is located in the upper Desaguadero Basin in the Bolivian Altiplano, more distant than Tiwanaku from the (fluctuating) margin of Lake Titicaca (Figure 1). The immediate environs of Khonkho Wankane, however, also included multiple qochas, which may have partially altered the local environment to allow the habitation of more wetland species. In general, the paleoenvironment of this site is expected to have been drier than sites closer to the lake margin.

Khonkho Wankane is also expected to have had a lesser reliance upon natural resources from Lake Titicaca. Inclusion of vertebrate remains from this and other sources into the site deposits, however, cannot be assumed to have resulted from direct human action and must be examined taphonomically to determine the most likely vectors of deposition. This research reports the herpetological (reptile and amphibian) remains from Khonkho Wankane which come from deposits dated by ceramic seriation and radiocarbon assays from Late Formative I to Tiwanaku periods, approximately 150 to 500 AD (Janusek, 2008). As with all osteological remains, the question of how they were introduced into site deposits and by what taxon, human or nonhuman, is critical to their paleoecological interpretation (Pinto Llona & Andrews, 1999). In addition, their generally small size leads to lesser archaeological visibility, both through a lack of recovery and a lowered survivability or identifiability of skeletal elements. Analysis of these small faunas, however, allows for clearer palaeoecological modelling of the inhabitants of this locality, both as palaeoenvironmental indicators and as taxa which in some cases were harvested from the natural environment. Their presence in a site cannot be assumed to be the result of direct human capture for consumption. Human

Figure 1. Map of the south Lake Titicaca area, showing Khonko Wankane and other sites described in the text.


Int. J. Osteoarchaeol. 23: 475–484 (2013)

Herpetofaunal Remains from Khonkho Wankane actions potentially include ceremonial, medicinal or tool manufacture usage (see below). Taphonomic analysis of these remains is critical to these interpretations, because the manner in which these small faunal remains were introduced into the sites and modified after their introduction largely defines their interactions with the human inhabitants of Khonkho Wankane. Small faunal assemblages, especially herpetofauna, generally have gone underreported from Altiplano archaeological contexts, a situation which has been changing as archaeological recovery systems include finer screening and flotation of samples. This includes recent excavations at the primarily Formative period site of Chiripa on the shore of Lake Titicaca, which have yielded abundant microvertebrate remains indicating exploitation of multiple lake faunas (Kent et al., 1999; Moore et al., 1999). Recovery of these smaller taxa is compromised when these methods are not employed, and the scope of paleoenvironmental and paleoecological inference likewise decreases (Freeman et al., 1998; Pokines, 1998, 2000; Sampson, 2003). This recovery of smaller and more delicate fauna must also be accompanied by a taphonomic analysis of their modes of site entry because remains of this nature have multiple paths through which they can be incorporated into site deposits.

Materials and methods Standard archaeological methodology at Khonkho Wankane by the Proyecto Jach’a Machaca excavations employed at least 0.25inch (6.4mm) screens to all deposits, which allows adequate recovery of only the larger taxa/elements thereof among those discussed here. Some specialised contexts were screened using 0.125inch (3.2mm) mesh. The recovery of smaller individual elements and smaller taxa will be biassed against using the former size mesh (Pokines, 2000; Sampson, 2003). In each case, where these were encountered, visible clusters of small faunal remains (see succeeding text) were excavated en masse. All elements from these clusters were extracted manually and without screening: sediment was removed via direct manipulation under a low-power binocular microscope. Representative recovery of smaller taxa/elements thereof, in these instances therefore, is possible, and more analytical significance can be placed upon skeletal representation frequencies from these clusters (discussed succeedingly). All small faunal osseous remains were identified to species level where possible, using all elements. In many cases, skeletal remains could be identified only to family level or higher. The number Copyright © 2011 John Wiley & Sons, Ltd.

477 of identified specimens (NISP) and minimum number of elements (MNE) counts take into account bone siding, portion, taxon, age class and archaeological provenience. Osteological terminology for amphibian remains follows Maglia et al. (2007). Identifications were made using the faunal collections of the Field Museum, Chicago, IL, USA.

Taxonomic representation The herpetofaunal total is 1710 from all levels/areas excavated at Khonkho Wankane. The herpetofaunal remains include a minimum of six taxa (Table 1), representing a significant portion of the relatively depauperate modern herpetofauna recorded for the Lake Titicaca area. The harsh, arid alpine environment limits herpetofaunal taxonomic diversity, and the Andes form a natural barrier for species expansion and genetic mixing (Baudoin & Pachecho, 1991; Correa et al., 2007; De la Riva et al., 2000; Duellman, 1979; Ergueta Sandóval, 1991; Linares, 2005; Méndez et al., 2004; Padial et al., 2003; Rodríguez et al., 1993; Schulte et al., 2000; Vellard, 1992a, 1992b). These taxa are relatively unknown ecologically, including the frog genus Telmatobius, multiple species of which inhabit Lake Titicaca and its environs (Benavides et al., 2002). These include T. culeus, the giant frog endemic to Lake Titicaca. Only smaller members of this genus were detected among the herpetofaunal remains. Table 1. Khonkho Wankane herpetofaunal taxonomic representation. Taxon

Common Name

Reptile Tropiduridae Liolaemus Amphibian Bufonoidea Bufonidae Rhinella spinulosa (= Bufo spinulosus) Ranoidea Hylidae (cf. Gastrotheca marsupiata) Leptodactylidae Pleurodema marmorata/cinerea Telmatobius sp. (lg.) Telmatobius sp. (sm.) Total

Khonkho Wankane NISP

iguanas, etc. 22 529 toads Andean toad frogs tree frogs marsupial frog

192 5 19 99 703 1 140 1710

NISP, number of identified specimens.


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478 A single reptile of the genus Liolaemus (lizard, family Iguanidae) was detected at Khonkho Wankane (NISP=22). This genus is widespread and diverse, with over 160 species known from the Andes and surrounding lowlands (Frost, 2008; Schulte et al., 2000), and many recently described (Espinoza & Lobo, 2003; Hernandez& Espinoza, 2004; Lobo & Espinoza, 1999, 2004). Formerly described species present in the area include L. multiformis, which inhabits altitudes approximately 4300–4900m (Baudoin & Pachecho, 1991; Ergueta Sandóval, 1991). The skeletal remains recovered from Khonko Wankane may belong to this species or another Liolaemus. Amphibian remains are abundant among the Khonko Wankane faunal sample. Their exact identification is hindered by the multiple species present from the family Leptodactylidae, which includes multiple species of Telmatobius and two species of Pleurodema. The Pleurodema species present in the Lake Titicaca area today, P. marmorata and P. cinerea, may both be present at Khonkho Wankane, as they are similar in size: P. cinerea has a body length of 42mm (males)/45mm (females), whereas P. marmorata has a body length of 38mm (males)/46mm (females) (Duellman & Veloso, 1972). Ergueta Sandóval (1991) indicated that P. marmorata populations can be even smaller: 25mm (males)/28 mm (females). Pleurodema dominates the Amphibian sample, with a NISP=703. Osteological variation among the smaller amphibians therefore may also be explained by sexual dimorphism. In addition, a small species (much smaller than the giant lake frog, T. culeus and related forms) of Telmatobius (NISP=140) and one slightly larger species, also most consistent with Telmatobius (NISP=1), are present. Either of these may correspond to T. marmoratus or related species as discussed by Vellard (1992a), who noted the distinctive forms of small to medium Telmatobius found in nearly every bay of Lake Titicaca and intergradations between these forms. Additional description of these forms is necessary to establish clear species boundaries (Benavides et al., 2002; De la Riva et al., 2000). Although the giant forms of Telmatobius inhabit deep water, the smaller forms are not confined to the lake, inhabiting streams and shallow waters of the Lake Titicaca basin (Benavides et al., 2002). Pleurodema cinerea inhabits puna/semi-arid montane environments at elevations of 2900–4100m. They are not confined to the Lake Titicaca environs or similar environments and breed in temporary ponds (Duellman & Veloso, 1972:18). Pleurodema marmorata inhabits wet and dry puna (grassland) at elevations of 3000–5000m. They breed in permanent or temporary ponds. This species can endure environmental extremes and has been found Copyright © 2011 John Wiley & Sons, Ltd.

under snow-covered rocks and observed hopping across a snow field (Duellman & Veloso, 1972: 20–21). This behaviour of sheltering in hollows to avoid poor temperature conditions could act to introduce this species into archaeological sites. Another amphibian taxon, Gastrotheca, is likely present (NISP=19). This genus of tree frog, family Hylidae, has a single representative in the southern Peruvian and Bolivian Altiplano today, G. marsupiata (De la Riva et al., 2000; Duellman & Fritts, 1972). This species in Bolivia is sometimes referred to as G. boliviana (Vellard, 1992a). It is found at altitudes from 2560–4360m (Duellman & Fritts, 1972:19–20). Like other members of this genus, G. marsupiata broods its young in a pouch and releases tadpoles into shallow water sources (Duellman & Fritts, 1972), which is an adaptation allowing this otherwise lowland genus to colonise the arid Altiplano environment. The body length is up to 41.6 mm (males)/46.5mm (females) (Duellman & Fritts, 1972:17). Both larger and more distinguishable osteologically, Rhinella spinulosa (Andean toad) is likely thereby biassed upwards in representation (NISP=192). [Note that this species was and is still recorded frequently as Bufo spinulosus, and that R. spinulosa is a recent revision.] This species has a broad range through the Andes and can be found at altitudes from 1000–5000m (Duellman, 1979:457). It is highly adapted to the harsh Altiplano climate and open pastures and also will shelter in natural crevices and hollows as part of thermoregulatory behaviour (Sinsch, 1989). In addition, many skeletal remains could only be identified to a broader taxon such as Amphibian (n=529), either due to fragmentation or inherent unidentifiability of elements such as phalanges.

Taphonomic analysis None of the herpetofaunal remains displays the types of direct taphonomic alteration often indicative of human dietary usage, including gastric erosion, cut marks, or thermal alteration. Similarly, no markers of predation sometimes left by other species, which include small carnivores or raptors, were noted, including breakage and gastric erosion (Pinto Llona & Andrews, 1999). Because of their small size and delicate nature, it is unlikely that the skeletal remains of the amphibian and reptile species found in the Lake Titicaca region would survive mastication, gnawing or digestion by a large carnivore (including humans) in such a recognisable state. Burning, accidental or deliberate, would also tend to destroy their small, delicate bones. As with the Int. J. Osteoarchaeol. 23: 475–484 (2013)

Herpetofaunal Remains from Khonkho Wankane

479

mammalian microfauna, the introduction of significant amounts of herpetofaunal remains via predation by diurnal raptors is not supported by their taphonomic characteristics and recovery pattern. Introduction of some of these remains via owl pellet egestion, particularly by barn owl (Tyto alba), is possible (Pokines, 1998), as this species has been identified from this site and tends to leave behind osseous prey remains with little breakage or gastric erosion. No signs of weathering (sensu Behrensmeyer, 1978) were noted on any of the herpetofaunal remains. Weathering (with its inherent cracking and degradation) likely would have rendered remains of this nature too friable for recovery and/or subsequent identification.

Skeletal element representation and clustering Table 2 presents the overall amphibian MNE and NISP, categorised by element. Apparent is the predominance of postcranial elements over cranial elements (Figure 2). The ilium (MNE=225), femur (MNE=213), humerus (MNE=190), tibio-fibula (MNE=128) and radio-ulna (MNE=68) are all paired, relatively robust and identifiable postcranial elements that were recovered in large amounts. The urostyle Table 2. Khonkho Wankane amphibian skeletal representation. Total

Cluster B

Element

MNE

NISP

MNE

Ilium Femur Humerus Vertebra Tibio-fibula Urostyle Radio-ulna Scapula Pterygoid Mandible Parasphenoid Coracoid Sacrum Spenethmoid Clavicle Suprascapula Maxillary Frontoparietal Pubic/Ischium Cranial – other Exoccipital Foot element Fragment Total

225 213 190 171 128 72 68 39 33 28 19 22 18 16 15 9 8 4 4 3 1 N/A N/A

225 214 194 206 130 73 72 39 33 28 27 22 18 16 15 13 8 4 4 3 1 173 172 1690

199 188 150 98 91 57 51 29 28 23 11 21 12 16 5 5 5 2 4 0 1 N/A N/A

NISP 199 189 154 129 91 58 55 29 28 23 11 21 12 16 5 9 5 2 4 0 1 141 148 1330

Cluster C MNE

NISP

14 11 18 65 15 10 9 7 5 5 8 1 4 0 7 4 2 2 0 3 0 N/A N/A

14 11 18 69 15 10 9 7 5 5 16 1 4 0 7 4 2 2 0 3 0 16 14 232

MNE, minimum number of elements; NISP, number of identified specimen.


Figure 2. Khonkho Wankane overall amphibian skeletal representation by minimum number of elements (MNE).

(MNE=72) is a singular postcranial element and therefore was recovered in similar relative amounts. Other postcranial elements were recovered in lower amounts, probably due to their greater fragility and lesser identifiability. These include the scapula (MNE=39), coracoid (MNE=22), clavicle (MNE=15) and suprascapula (MNE=9). In contrast, the cranial elements are underrepresented by approximately an order of magnitude, with such paired elements such as the mandible (MNE=28), pterygoid (MNE=33), and maxillary (MNE=8) and singular elements such as the spenethmoid (MNE=16) recovered in small relative amounts. A portion of this disparity can be explained through recovery bias, where the generally smaller, less robust and potentially less identifiable cranial elements were selected using screen mesh size or lower visibility. The majority of the amphibian remains (Table 2) from Khonkho Wankane were discovered in one large cluster (Cluster B; total number of identified remains plus fragments=1330) and one smaller (Cluster C; n=232). Clusters B and C contributed n=1562 or 92.4% of the amphibian total (n=1690). Cluster B Int. J. Osteoarchaeol. 23: 475–484 (2013)

480

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includes all except one amphibian taxon [Telmatobius sp. (lg.)] identified at Khonkho Wankane. Also present in this assemblage of bone are Liolaemus (NISP=18) and mammalian microfaunal remains (NISP=37), which include Galea musteloides (common yellow-toothed cavy) and Oligoryzomys destructor (destructive pygmy rice rat). Cluster C includes only Pleurodema, and no other vertebrate remains. The skeletal patterning for Clusters B and C (Figures 3 and 4) also shows the large disparity of postcranial elements over cranial remains. The possible mechanisms that would act to cluster amphibians in these contexts therefore must also explain the clustering of other small vertebrates in these same contexts. Aggregation of live amphibians has been noted in the Altiplano. Espinoza & Quinteros (2008) found that diurnal, above-ground aggregation behaviour among Rhinella spinulosa inhabiting regions above 4300m provided thermal benefits to the toads. They noted clusters estimated to be up to ~200 subadult (recently metamorphed) individuals. As noted earlier, Rhinella spinulosa is known to shelter under damp stones and

Figure 4. Khonkho Wankane amphibian Cluster C skeletal representation by MNE.

Figure 3. Khonkho Wankane amphibian Cluster B skeletal representation by MNE.


natural crevices (Duellman, 1979; Sinsch, 1989). It is possible that underground aggregations also may benefit Altiplano amphibian species, to avoid dry or cold conditions (Espinoza & Quinteros, 2008). Subterranean aggregation die-offs need not have occurred as a single event, and instead they may have resulted from multiple uses of the same hole and by multiple species. Because the same Cluster (B) of small faunal remains includes mostly juvenile mammalian microfauna, the natural die-off scenario is supported. Multiple rodent species in the Altiplano shelter and raise their litters in burrows, whether excavated by their own species or taken over from another species, especially the avid burrowing species Ctenomys leucodon (white-toothed tuco-tuco) (Eisenberg & Redford, 1999). This species is present in the Khonkho Wankane area today. Clusters B and C are not consistent with predator accumulations. No predator in the Lake Titicaca area preys nearly exclusively upon amphibians and leaves highly intact, unaltered skeletal remains in such a concentrated area (Pinto Llona & Andrews, 1999). Although nocturnal raptors can concentrate bone in Int. J. Osteoarchaeol. 23: 475–484 (2013)

Herpetofaunal Remains from Khonkho Wankane relatively small areas and leave behind largely intact skeletal remains, much greater species diversity is expected, including an avifaunal component and a much higher mammal microfauna component. The decreased amphibian cranial MNE relative to postcranial MNE is not likely due in this case to recovery method, because Clusters B and C did not undergo any screening, which potentially could have biassed against smaller (in many cases cranial) skeletal elements. Excavation of these clusters proceeded en masse, with the entire visible concentrations of bone collected. The skeletal representation discrepancy may be due to the development of the amphibian skeleton. Maglia et al. (2007) found that cranial ossification in some small (miniature) amphibian species can be delayed, partial or nonexistent. Cartilaginous structures would not preserve under these taphonomic conditions. This developmental research, however, has not been carried out directly upon Altiplano amphibian species, so this explanation remains speculative regarding taxa such as Pleurodema. Taking into account among Altiplano amphibian species the known subterranean sheltering behaviour, aggregation of individuals for thermal and other benefits, and the possibility of delayed ossification among cranial elements, it is possible that the clustering of amphibian remains resulted from natural behaviour and skeletal development and is not the result of deliberate human action. All of these behavioural and biological development questions require specific research among these amphibian taxa. Because the vast majority of herpetofaunal remains recovered from Khonkho Wankane were concentrated in highly focal clusters, this scenario explains the likely taphonomic origin of the majority of these taxa into the site deposits and indicates that their origin is likely through recent burrowing intrusion.

Human utilisation

Ceremonial/Medicinal Human ceremonial and medicinal utilisation of amphibian species in the greater Andean region is still widespread (Cooke, 1989). Vellard (1992a) noted that amphibians are used by modern Altiplano inhabitants, both Aymaraspeaking and Quechua-speaking. These uses include dead and living specimens in medicine and ritual, with living members of the large Lake Titicaca species Telmatobius culeus often used in rituals performed to alleviate drought conditions. Toads (probably including Rhinella spinulosa because of its ubiquity) are used to Copyright © 2011 John Wiley & Sons, Ltd.

481 produce potions. Given possible uses such as these, difficulty of capture or low body mass yield may have been viewed as unimportant among Khonkho Wankane inhabitants. Ritual uses also would not have necessarily left behind visible taphonomic markers upon amphibian bone. It is unknown if smaller concentrations of amphibian bone are the result of deliberate burial by humans, because the skeletal and taphonomic pattern would be identical to natural mortality of animals sheltering in a burrow. Because this type of small animal offering is known from Khonkho Wankane in the form of falcon (likely aplomado falcon, Falco femoralis) burials, then it is possible that some amphibian remains were introduced to the site deposits in this manner.

Dietary The majority of amphibians and reptiles represented by these remains are relatively small taxa, so their inclusion in human diet is less likely on the logical grounds of efficient resource utilisation. Smaller taxa, unless easily harvested in groups, give a lower return on time, resources and energy expended relative to larger taxa. This can be ameliorated through technology, such as the harvesting of small fish using nets, traps or weirs, which was likely the case with the fish remains introduced into Khonkho Wankane and other Altiplano sites from Lake Titicaca. It is notable that no remains of the large form of lake frog, Telmatobius culeus, were recovered. If the desired product was amphibian meat, a potentially more productive resource apparently was not harvested despite clear use of other aquatic resources. Human exploitation of herpetofauna for food is known from the greater Andean region (Albino & Kligmann, 2007; Cooke, 1989), such as the hunting and consumption of lizards on the north coast of Peru, which has persisted into recent times (Holmberg, 1957). The use of long, portable barriers to direct the movement of these prey toward human hunters makes their harvest efficient. These lizards, called locally cañanes (Dicrodon holmbergi), are roasted whole, which would be expected to leave occasional thermal damage on osseous extremities. It is unknown if the bones of this species are consumed with the meat, but their direct consumption would be expected to leave clear taphonomic markers (breakage, chewing marks, and gastric corrosion) behind. Indeed, it is possible that very little would survive human consumption of small lizard taxa such as Liolaemus, if these were cooked and consumed whole. Processing methods used on herpetofauna in this region including drying/salting or boiling, however, need leave behind no grossly visible taphonomic signatures on microvertebrate osseous remains, unlike other Int. J. Osteoarchaeol. 23: 475–484 (2013)

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482 methods, which include butchery, roasting or skin/ gland removal to remove their contained toxins (Cooke, 1989). Dietary exclusion of small reptile taxa by humans due to their small size or cultural factors therefore cannot be assumed, nor would the food remains necessarily survive in archaeologically detectable quantities. No evidence of their exploitation for this purpose has been detected at Khonkho Wankane.

Paleoenvironment The use of this sample in paleoenvironmental reconstruction of Khonkho Wankane during its habitation is extremely limited, because the majority of them was recovered in Clusters B and C and were likely introduced into the site deposits intrusively at an unknown later time. The total NISP=128 for all contexts, excluding these two clusters. Taphonomic analysis such as this should serve as a cautionary note regarding the use of herpetofaunal remains from Altiplano contexts in paleoenvironmental interpretation, unless it can be demonstrated that their deposition was likely contemporaneous with site occupation. Given the ubiquity of burrowing rodent species in the area and the frequent reuse of their tunnel systems by multiple species, recovery context should be considered. No deviation from the modern herpetofauna of the region was noted, although the complexity of evolutionary variation within the genus Telmatobius and other taxa described here requires further examination in both modern and past contexts.

Summary and conclusions Taphonomic analyses specific to herpetofauna are rare in the taphonomic literature (Pinto Llona & Andrews, 1999), and the present research addresses one particular mode of entry of this type of skeletal remains into an Andean Altiplano site. Taphonomic and behavioural analysis of the herpetofaunal remains from Khonkho Wankane indicates that the majority of them were likely introduced to the site deposits recently via natural die-offs, while sheltering in re-used rodent burrows and were not a major component of the terrestrial vertebrate fauna of that site during its initial formation. The archaeological visibility of herpetofaunal remains in this size class, however, would be increased through fine screening of site deposits, so this source of vertebrate remains may become more prominent in future Altiplano excavations. This example illustrates the necessity for a critical analysis of the modes of entry of Copyright © 2011 John Wiley & Sons, Ltd.

osseous remains into a site to prevent mistaken interpretations of direct dietary or ceremonial usage of species, where a more mundane explanation is more plausible.

Acknowledgements This research was conducted in conjunction with the Proyecto Arqueologico Jach’a Machaca and also was sponsored by Vanderbilt University and the Joint POW/ MIA Accounting Command, Central Identification Laboratory, Hawaii. John Janusek of Vanderbilt University provided access to all materials and field support. The author also thanks Alan Resetar, Division of Amphibians and Reptiles, Field Museum, Chicago for his invaluable advice and access to comparative collections. The author also thanks the two reviewers of an earlier draft of this report for their highly valuable comments.

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