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International Journal of Osteoarchaeology Int. J. Osteoarchaeol. 23: 746–756 (2013) Published online 10 November 2011 in Wiley Online Library (wileyonlinelibrary.com) DOI: 10.1002/oa.1299

SHORT REPORT

Avian Remains from Khonkho Wankane, an Urban and Ceremonial Center in the Southern Lake Titicaca Altiplano J. T. POKINESa* Boston University School of Medicine, Dept. of Anatomy & Neurobiology, 72 East Concord St., L1004, Boston, MA 02118, USA

ABSTRACT

The urban and ceremonial center Khonkho Wankane flourished in the southern Lake Titicaca Altiplano region during the Late Formative prior to the rise of Tiwanaku as a dominant urban center. This site yielded an avifaunal number of identified specimens (NISP) = 631, with a minimum of 26 taxa represented among 539 skeletal elements or fragments. A total of 92 egg shell fragments also were recovered. The avifauna inhabited two major ecological zones: Lake Titicaca, its margins/wetlands and other water margins; and the dry Altiplano grassland (or puna). Lake/wetland taxa include Phoenicopterus chilensis (Chilean flamingo), Nycticorax nycticorax (night heron) and multiple taxa in the Order Anseriformes (ducks, geese, etc.) including Chloephaga melanoptera (Andean goose), Anas georgica (yellow-billed pintail), A. flavirostris (speckled teal) and Oxyura jamaicensis (Andean ruddyduck). Also present are Fulica ardesiaca (Andean coot), Gallinago andina (puna snipe), Himantopus mexicanus (black-necked stilt) and Charadrius alticola (puna plover). Puna taxa include Nothoprocta cf. ornata (ornate tinamou), Nothura cf. darwinii (Darwin’s nothura), Metropelia sp. (ground doves), Athene cunicularia (burrowing owl), and multiple Passeriformes (songbirds, etc.). Taxa with a cosmopolitan distribution include Falco cf. femoralis (aplomado falcon) and Tyto alba (barn owl). Taphonomic analysis indicates that the avifauna were used in ceremonial contexts, including multiple Falco cf. femoralis burials, and for dietary and possibly tool-making purposes. Other sources of introduction into the site deposits likely include natural mortality of taxa directly inhabiting the site area. These findings are compared to the avifauna recovered from the Formative site of Chiripa along the Lake Titicaca margin, which included a much higher proportion of lake bird fauna. At Khonkho Wankane, the importance of wetland avifauna may have been enhanced by the alteration of the local environment to include qochas, or artificial reservoirs/ponds, and continued despite reliance upon multiple domesticated plant and mammal species. Copyright © 2011 John Wiley & Sons, Ltd. Key words: Avifauna; Khonkho Wankane; Tiwanaku; Bolivia; Altiplano; Paleoecology; Taphonomy

Introduction Khonkho Wankane emerged as an urban and ceremonial center 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 the southern Lake Titicaca region, by 500 AD. Some residential occupation of Khonkho Wankane continued during * Correspondence to: James Pokines, BU School of Medicine. e-mail: [email protected]

Copyright © 2011 John Wiley & Sons, Ltd.

the Tiwanaku period. The site of Tiwanaku itself flourished from around 500 to 1100 AD (Bermann, 1994; Janusek, 2003, 2008; Stanish, 2003). The paleoecological adaptations of this major political and ceremonial center and its subsidiary sites to this relatively harsh (ca. 70 cm annual rainfall), high-altitude (ca. 3800 m) 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 kilometers, this artificial system served to ameliorate the microenvironment around the agricultural plots in order to increase output and reliability 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). Received 19 January 2011 Revised 5 October 2011 Accepted 7 October 2011

Avian Remains from Khonkho Wankane Alteration of the landscape also included the formation of qochas, or artificial reservoirs or ponds, to store rainwater (Janusek, 2008; Craig et al., 2011). These artificial systems therefore created or altered a large system of wetlands around Lake Titicaca, which no doubt had an effect upon the many species of wetland bird taxa inhabiting or migrating through the region. This resource, along with lake fish (Capriles et al., 2008), formed a portion of southern Lake Titicaca Altiplano diet less visible archaeologically than the remains of large camelids (Webster & Janusek, 2003; Gladwell, 2010) and other vertebrate taxa utilised throughout the Andes in antiquity (Miller & Burger, 1995; Marcus et al., 1999; Pérez Arias, 2007). Economic activities also included quinoa and potato agriculture (Bermann, 1994; Bruno & Whitehead, 2003; Stanish, 2003; Pérez Arias, 2007; Janusek, 2008). 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 paleoecological adaptation of Khonkho Wankane to this marginal, high-altitude environment therefore took place with lesser immediate access to natural wetland resources than Tiwanaku and with far less direct access than Chiripa, a primarily Formative period (ca. 1500–100 BC) site adjacent to the lake shore (Hastorf, 2003). The area surrounding Khonkho Wankane, however, also included multiple qochas, the closest of which were approximately 5 km distant. These artificial ponds may have attracted lake birds somewhat closer to the human habitations in an area that was otherwise dry grassland and agricultural land with only a few areas of natural wetlands interspersed at the base of the nearby Kimsachata range

747 to the northeast. This natural aquatic resource therefore may have existed in addition to the direct agricultural and pastoral benefits of water management and storage within an area both low and highly seasonal in rainfall. It is noteworthy that small qochas excavated in the Khonkho Wankane area by local farmers today often attract avifauna, including multiple species of duck (Anseridae). The avian remains from Khonkho Wankane ceremonial and domestic contexts analysed here come from deposits dated by ceramic seriation and radiocarbon assays from Late Formative I up to early Tiwanaku periods at Khonkho Wankane, approximately 150 to 500 AD (Janusek, 2008). Avian remains typically are of a smaller size and less overall durability, lessening their archaeological visibility through a lack of recovery and a lowered survivability. Analysis of avifaunas, however, allows for broader paleoecological modeling of the inhabitants of this locality who harvested these taxa from the Altiplano environment. In some cases, remains were introduced independently of human action, including through bird occupation of the site independent of humans, or consumption and deposition of any small species by nonhuman predators including carnivores and raptors. In addition, human actions are not limited to dietary consumption and in some cases included ceremonial, medicinal or tool manufacture usage. Taphonomic analysis of these remains is critical to these interpretations, since the manner in which these avifaunal remains were introduced into the sites and modified after their introduction defines a large portion of the human interactions with this zoological portion of their environment. This interaction also extended to portrayals in iconography (Janusek, 2008), which further illustrates the importance of avifauna to the inhabitants of Khonkho Wankane and other southern Lake Titicaca sites. Small faunal assemblages 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 Chiripa, which have yielded abundant small fauna remains indicating exploitation of multiple lake taxa (Kent et al., 1999; Moore et al., 1999; Hastorf, 2003; Capriles et al., 2008) as well as evidence for early quinoa cultivation (Bruno & Whitehead, 2003).

Materials and methods

Figure 1. Map of the southern Lake Titicaca region, showing the sites described in the text.


Standard archaeological methodology at Khonkho Wankane by the Proyecto Jach’a Machaca excavations employed at least 0.25-inch (6.4-mm) screens to all deposits, which allows adequate recovery of the larger Int. J. Osteoarchaeol. 23: 746–756 (2013)

J. T. Pokines

748 taxa/elements thereof among the avifauna. Some specialised contexts were screened using 0.125-inch (3.2-mm) mesh. Recovery of these smaller taxa increases significantly when fine-mesh screening methods are employed, and the scope of paleoenvironmental and paleoecological inference likewise decreases (Freeman et al., 1998; Pokines, 1998, 2000). The recovery of individual elements, particularly, will be biased against using 0.25-inch mesh. In each case where encountered, visible clusters of small faunal remains were excavated en masse, and all elements were extracted manually. Representative recovery of smaller taxa or elements thereof in these instances therefore is possible, and more analytical significance can be placed upon skeletal representation frequencies from these contexts. All avifaunal osseous remains were identified to species level where possible. In many cases, remains could be identified only to genus level or higher (Superfamily or Order). Where it was not possible to identify an element or portion thereof to a taxonomic grouping, these were assigned to a general size class, from A to D. Size Class A is the smallest and includes small Passerines (songbirds, etc.); Size Class B includes individuals roughly in the size of doves/pigeons; Size Class C includes individuals the size of most ducks, falcons, etc.; and Size Class D includes the largest species found in the Lake Titicaca area (geese, herons, etc.). Not including these remains would otherwise bias taphonomic analyses based upon skeletal part representation. The NISP counts for all taxa will be lower than these inclusive amounts. Discussions will also include minimum number of individual (MNI), calculated as the minimum number of skeletons necessary to produce a given assemblage. These counts take into account bone siding, portion, taxon, age class and archaeological provenience. Minimum number of element (MNE) counts also take into account bone siding, portion, taxon, age class and archaeological provenience and are the per-part MNI. Identifications were made using the faunal collections of the Field Museum, Chicago, IL, USA.

Taxonomic representation The total number of avifaunal remains from Khonkho Wankane is 631. A minimum of 26 taxa are represented among 539 skeletal elements or fragments thereof identified as avifauna (Table 1). A total of 92 egg shell fragments also were recovered. No remains of historically introduced species were identified. More recent avifauna such as these could have been introduced to the site deposits through mixing. Due to the greater biodiversity among the Lake Titicaca avifauna (Fjeldså, Copyright © 2011 John Wiley & Sons, Ltd.

1985; Mitchell, 1985; Remsen & Traylor, 1989; Fjeldså & Krabbe, 1990; Dejoux, 1992; Kent et al., 1999; Erize et al., 2006; Herzog, 2006; Schulenberg et al., 2007) relative to other small fauna classes and the inherent fragility of avian bones, identification in many cases could not proceed to the species level. Included in this total are a large number of passerine (songbirds, etc.) species, the exact identification of which often is hindered by their small size and multiple closely related forms inhabiting the same area. All identified taxa are present in the Lake Titicaca region today (Fjeldså & Krabbe, 1990; Dejoux, 1992; Kent et al., 1999; Schulenberg et al., 2007). The avifauna present at Khonkho Wankane inhabit two major ecological zones: (i) Lake Titicaca, its associated margins/wetlands and other wetlands, and (ii) the dry Altiplano grassland (or puna). Species with cosmopolitan distributions also are present. These habitat preferences largely hold by avian Order, and ecological accounts follow Dejoux (1992), Fjeldså and Krabbe (1990), Kent et al. (1999) and Schulenberg et al. (2007). Khonkho Wankane is distant enough from the Lake Titicaca margin that the presence of waterfowl remains is most likely due to direct human exploitation of the majority of these taxa. Avifauna native to the dry grassland environment surrounding the site also potentially were introduced to the deposits through natural mortality. In some cases, direct human action is indicated through additional contextual and taphonomic evidence.

Lake/Wetland taxa The Order Phoenicopteriformes (flamingoes) is represented by a single species, Phoenicopterus chilensis (Chilean flamingo; NISP = 3; MNI = 1). Multiple flamingo species inhabit the Lake Titicaca region. These large birds wade shallow lake and salt water flats, straining small organisms with their specialised bills. They are highly gregarious, congregating in large flocks. The Order Ciconiiformes (storks, herons, egrets, etc.) is also represented by a single species, Nycticorax nycticorax (night heron; NISP = 9; MNI = 7), although an additional member of this order may be present (NISP = 1). This more solitary species is associated with water margin and shallow wetland habitats. The presence of P. chilensis and N. nycticorax in Khonkho Wankane deposits is most likely due to direct predation and transport by a large carnivore, human or otherwise, from the nearby lake habitat. The Order Anseriformes (ducks, geese, etc.) is represented abundantly, and all taxa are members of the family Anatidae (total NISP = 62). These include a single element most consistent with Chloephaga melanoptera Int. J. Osteoarchaeol. 23: 746–756 (2013)

Avian Remains from Khonkho Wankane

749

Table 1. Khonkho Wankane avifaunal taxonomic and habitat representation Taxon Tinamiformes Nothoprocta cf. ornata cf. Nothoprocta ornata Nothura cf. darwinii cf. Nothura darwinii Phoenicopteriformes Phoenicopterus chilensis Ciconiiformes cf. Ciconiiformes Nycticorax nycticorax Anseriformes Anatidae (lg.) cf. Chloephaga melanoptera cf. Anas Anas Anas flavirostris Anas cf. flavirostris Anas cf. georgica Oxyura jamaicensis cf. Oxyura jamaicensis Falconiformes Falco cf. femoralis Gruiformes Fulica ardesiaca Charadriiformes Charadriiformes, small Himantopus mexicanus cf. Himantopus mexicanus Charadrius cf. alticola Scolopacidae Gallinago cf. andina Tringa flavipes cf. Tringa flavipes Tringa solitaria Columbiformes cf. Metropelia Strigiformes Athene cunicularia Tyto alba Piciformes Colaptes cf. rupicola cf. Colaptes rupicola Passeriformes Passeriformes, small Geositta tenirostris Geositta cunicularia Fringillidae cf. Sicalis uropygialis Agelaius thilius Total Size Class A Size Class B Size Class C Size Class D Total Size Class C Size Class D Total Total Primary Habitat Wetlands Wetlands/dry grassland Dry grassland Cosmopolitan Total (a)

Habitat(s)(a)

Common Name tinamous ornate tinamou

NISP

MNI

9 3 37 1

7 3 18 1

shallow lake

3

1

lake/water margins lake/water margins

1 9

1 7

9 1 1 8 32 2 1 6 2

8 1 5 9 2 1 3 1

103

16

2

2

3 4 2 16 2 32 1 1 3

4 2 5 2 3 1 1 2

dry grassland

4

2

open dry habitats Cosmopolitan

2 1

1 1

dry grassland/roosts in rocks, houses dry grassland/roosts in rocks, houses

3 1

3 1

16 1 4 3 10 1 340 29 80 64 26 199 2 90 92 631

7 1 1 2 2 1 128

dry grassland/scrub dry grassland/scrub dry grassland dry grassland

Darwin’s nothura flamingoes Chilean flamingo storks, herons, egrets, etc. night heron ducks, geese, etc.

lake/wetlands lake margin/wetlands lake/wetlands lake/wetlands lake/wetlands lake/wetlands lake/wetlands lake/wetlands lake/wetlands

Andean goose

speckled teal yellow-billed pintail Andean ruddyduck falcons, hawks, etc. aplomado falcon rails, coots, cranes, etc. Andean coot gulls, stilts, sandpipers, etc.

cosmopolitan/open lake/wetlands lake margin/wetlands lake margin/wetlands lake margin/wetlands lake margin/wetlands lake margin/wetlands lake margin/wetlands lake margin/wetlands lake margin/wetlands lake margin/wetlands

black-necked stilt puna plover snipes, sandpipers, etc. puna snipe lesser yellowlegs solitary sandpiper pigeons, doves ground doves owls burrowing owl barn owl woodpeckers, flickers, etc. Andean flicker songbirds, etc. slender-billed miner common miner siskins bright-rumped yellow-finch yellow-winged blackbird

dry grassland dry grassland dry grassland/forest edge/forest dry grassland/rural habitations wetlands/dry grassland

(sparrows, etc.) (doves, etc.) (ducks, falcons, etc.) (geese, herons, etc.) (egg shell fragments) (egg shell fragments)

NISP 141 1 94 104 340

% 41.5% 0.3% 27.6% 30.6% 100.0%

% 50.4% 0.8% 34.7% 14.0% 99.9%

MNI 61 1 42 17 121

Following: Dejoux (1992); Fjeldså (1985); Fjeldså & Krabbe (1990); Kent et al. (1999); Schulenberg et al. (2007).


Int. J. Osteoarchaeol. 23: 746–756 (2013)

750 (Andean goose) and a single element most consistent with Anas georgica (yellow-billed pintail); multiple elements (NISP = 34; MNI = 11) identified to/most consistent with Anas flavirostris (speckled teal), and multiple elements (NISP = 8; MNI = 4) identified to/most consistent with Oxyura jamaicensis (Andean ruddyduck). Additional remains of unknown Anatidae are present, including an additional large species (NISP = 9; MNI = 8) and other remains ascribable to Anas. The primary habitat of all of these taxa within this region is Lake Titicaca, its shoreline and other wetlands. These taxa would not have been normally frequent inhabitants of the immediate Khonkho Wankane area, although the creation of artificial wetlands near the urban core may have attracted these species closer to humans. The Order Gruiformes (rails, coots, etc.) is represented by one species, Fulica ardesiaca (Andean coot). This lake and wetlands species is highly gregarious and occurs in large flocks, yet occurs only in a small amount (NISP = 2; MNI = 2) at Khonkho Wankane. The Order Charadriiformes (gulls, stilts, sandpipers, plovers, etc.) has the highest diversity representation among the Khonkho Wankane avifauna. All of these taxa are associated with the Lake Titicaca margin or other types of wetlands. The most abundant (NISP = 32; MNI = 3) is Gallinago, with the species represented most likely G. andina (puna snipe). Two additional elements could only be identified to the family Scolopacidae, which includes the genera Gallinago and Tringa. The latter genus is represented by elements (NISP = 2; MNI = 2) identified as/most consistent with T. flavipes (lesser yellowlegs) and elements (NISP = 3; MNI = 2) identified as T. solitaria (solitary sandpiper). Multiple skeletal remains (NISP = 6; MNI = 6) are identified as or are most consistent with Himantopus mexicanus (black-necked stilt). The genus Charadrius is also highly represented (NISP = 16; MNI = 5), with the species represented most likely C. alticola (puna plover).

Puna taxa The Order Tinamiformes (tinamous) is represented by two taxa, Nothoprocta cf. ornata (ornate tinamou; NISP = 12; MNI = 10, including also those remains most consistent with this taxon) and Nothura cf. darwinii (Darwin’s nothura; NISP = 38; MNI = 19). Nothoprocta ornata and Nothura darwinii are grassland species found in the puna zone of the Andes, with the latter species also found at lower altitudes. Each species is primarily solitary and would have inhabited the immediate area around Khonkho Wankane. Of the Order Columbiformes (pigeons and doves), skeletal elements most consistent with the genus Copyright © 2011 John Wiley & Sons, Ltd.

J. T. Pokines Metropelia (ground doves) are present (NISP = 4; MNI = 2). Multiple species of Metropelia inhabit the Lake Titicaca region today, including M. melanoptera (black-winged ground dove), M. ceciliae (bare-faced ground dove), and M. aymara (golden-spotted ground dove). All of these species are primarily open-habitat dwelling and would have inhabited the area around Khonkho Wankane. Both M. melanoptera and M. ceciliae are also associated with human habitations. The Order Strigiformes (owls) is represented by one species associated with open, dry habitats: Athene cunicularia (burrowing owl; NISP = 2; MNI = 1). As this species is associated with the dry, open habitat surrounding Khonkho Wankane, its skeletal remains could have been introduced post-occupation into the site by its habit of roosting underground. The Order Piciformes (woodpeckers, flickers, etc.) has a single taxon represented, with a small amount of elements (NISP = 4; MNI = 4) most consistent with Colaptes rupicola (Andean flicker). This puna/grassland species would have inhabited the area around Khonkho Wankane and will sometimes nest in abandoned houses, so it is possible that its remains entered the site deposits through natural morality. The Order Passeriformes (songbirds, etc.; which includes about half of all extant bird species described worldwide) is well represented among the Khonkho Wankane avifauna. Multiple skeletal remains (NISP = 16; MNI = 7) could be identified only to the level of this Order. The identified taxa include two species of Geositta: G. tenirostris (slender-billed miner; NISP = 1) and G. cunicularia (common miner; NISP = 4; MNI = 1). The family Fringillidae (siskins) has multiple members present today in the Lake Titicaca region; multiple elements were most consistent with this family (NISP = 3; MNI = 2). Multiple remains most consistent with Sicalis uropygialis (bright-rumped yellow-finch; NISP = 10; MNI = 2) were identified, and a single element was identified as Agelaius thilius (yellow-winged blackbird). All identified taxa are primarily dry grassland inhabitants. Sicalis uropygialis is common on the Altiplano and also can be found among dispersed human habitations (Schulenberg et al., 2007:592). Agelaius thilius is also associated with open wetlands, and some members of the family Fringillidae also are associated with forest/ forest edge habitats (Fjeldså & Krabbe, 1990). All of the passerine taxa present are small enough to have been present around humans and therefore could have been introduced into the site deposits through natural mortality. They are also small enough to have been the prey of multiple common raptor species (Pokines, 1998; Reboledo & Lartiga, 1998), including two recovered from Khonkho Wankane (see below). Int. J. Osteoarchaeol. 23: 746–756 (2013)

Avian Remains from Khonkho Wankane

751 cf. rupicola; and distal tibiotarsus portions Nothura cf. darwinii and Nycticorax nycticorax. All of these taxa are large enough (Size Class B or larger) to have been plausible contributions to human diet. In the case of the tibiotarsus of N. nycticorax (night heron), the entire circumference has regular cut marks where it has been severed from the shaft, probably for the production of a bone tube or other tool from the remaining shaft. The overall pattern of taphonomic alteration indicates only a small degree of direct human action upon the avifaunal remains. Due to the relatively delicate nature of these remains, it is probable that elements undergoing gnawing, consumption, burning and processing were more likely to be destroyed and removed from the archaeological record.

Cosmopolitan taxa The most abundantly represented avian species (NISP = 103; MNI = 16) is Falco cf. femoralis (aplomado falcon; Order Falconiformes which includes falcons, hawks, etc.), a species slightly smaller than F. peregrinus (peregrine falcon). Although found in many habitats throughout its Neotropical range, F. femoralis usually hunts invertebrates and small vertebrates (primarily birds) in open habitats, especially dry grasslands (Keddy-Hector, 2000; Macías-Duarte et al., 2004; InfoNatura, 2007). They typically roost in arboreal nests abandoned by other species. The total number of remains identified to this taxon is highly inflated due to their introduction into the deposits as deliberate burials, along with some scattered remains (see below). Barn owl (Tyto alba; Order Strigiformes) is represented by a single element. This species is one of the most widely distributed bird species and is found throughout the world except in Arctic and boreal regions. It is found in multiple habitats, and as its name implies, it often roosts in human habitations, abandoned or otherwise. It commonly hunts in open environments, preying upon most small animals present within its territory. Tyto alba is not common in the Titicaca region today (Reboledo & Lartiga, 1998). Its presence in Khonkho Wankane indicates that some of the small vertebrates in the site deposits could have been introduced through egested pellets, which contain large amounts of identifiable bone (Pokines, 1998).

Skeletal element representation The Khonkho Wankane avifaunal skeletal representation is presented in Table 3. The overall recovery pattern likely has been shaped partially by recovery methods, since smaller but still relatively durable elements such as pedal (MNE = 27) and alal (MNE = 8) phalanges are underrepresented. This factor also may tend to underrepresent smaller taxa, especially passerines. Thin elements including the sternum (MNE = 19) are relatively underrepresented, even given its singular versus paired skeletal representation. Smaller but relatively delicate elements, especially vertebrae (MNE = 10) and ribs (MNE = 12), are highly underrepresented. Density-mediated destruction through the processes of consumption, trampling and compaction likely have affected this assemblage. The most highly represented elements are the most relatively large, durable and identifiable (Serjeantson, 2009), such as the humerus (MNE = 67), tibiotarsus (MNE = 62), ulna (MNE = 46), carpometacarpus (MNE = 33), femur (MNE = 31), tarsometatarsus (MNE = 30) and coracoid (MNE = 27). The ratio of anterior/wing element (humerus, ulna and carpometacarpus) versus posterior/leg element (femur, tibiotarsus and tarsometatarsus) total NISP has been

Taphonomic analysis The Khonkho Wankane avian remains exhibit some direct taphonomic alteration (Table 2). Carnivore gnawing (0.6%), rodent gnawing (0.2%), gastric corrosion (0.6%) and burning (0.6%) were detected in low amounts. No elements displaying weathering (sensu Behrensmeyer, 1978) were detected. A total of six elements (1.1%) display cut marks: two Size Class C partial ulnae; one nearly complete humerus of Oxyura jamaicensis and one partial humerus of Colaptes

Table 2. Khonkho Wankane avifaunal taphonomic indicators Faunal Component Khonkho Wankane Avifauna (n = 539)

Carnivore Gnawing

Rodent Gnawing

Gastric Corrosion

Cut Marks

Burnt

n=

%

n=

%

n=

%

n=

%

n=

%

3

0.6

1

0.2

3

0.6

6

1.1

3

0.6



J. T. Pokines

752 Table 3. Khonkho Wankane avifaunal skeletal representation Element

NISP

MNE

Humerus Tibiotarsus Ulna Carpometacarpus Femur Tarsometatarsus Coracoid Pedal phalanges Radius Sternum Pelvis Synsacrum Rib Scapula Mandible Vertebra Alal phalanges Furcula Maxilla Cranial – other Fragments Total

67 69 56 33 31 30 27 27 20 20 19 16 16 13 10 10 8 6 4 23 34 539

67 62 46 33 31 30 27 27 20 19 19 16 12 13 10 10 8 6 4 N/A N/A

postulated by multiple researchers as an indicator of avifaunal assemblage origin (summarised in Serjeantson, 2009). This ratio at Khonkho Wankane is 156:130 or 1.2:1, indicating a slight predominance of wing elements across all size classes and contexts. Expressed as a percentage with an assumed starting contribution of 50% for each category, anterior/wing elements contributed 55% of the total to 45% for the posterior/leg elements. Avifaunal assemblages which are of known anthropogenic origin tend to have ratios in favour of anterior/wing elements as opposed to assemblages produced by raptor or carnivore accumulation. Other factors which may contribute to this preponderance include greater robusticity of these elements due to the taxa represented, such as the large contribution made by waterfowl including the Anatidae identified in the Khonkho Wankane sample (Serjeantson, 2009:157). The members of this taxon tend to have large, powerful wings relative to their hindquarters, which may contribute to relative skeletal preservation. Egg shell fragments of large birds are represented in small amounts in the Khonkho Wankane deposits. The total NISP = 92, consisting of small to medium fragments. A minimum of two taxa are represented. Two fragments found together are cream colored and had a thickness ranging from 0.44 to 0.48 mm. This range of thicknesses is found among the Anserinae (geese), among other bird taxa, and is larger than those typical of the Anatinae (ducks) (Serjeantson, 2009:172–174). The remaining fragments come from multiple contexts, Copyright © 2011 John Wiley & Sons, Ltd.

in one to four fragments, one cluster of 11 fragments and one cluster of 68 fragments. All of these latter egg shell fragments retain a light green tint, range in thickness from 0.61 to 0.89 mm and appear to be from the same taxon in terms of overall appearance and morphology of shell arc. This range of thicknesses is at or beyond those found among Cygninae (swans) (Serjeantson, 2009:172–174), indicating their origin from a large avifaunal taxon. The sizes of all of these fragments therefore place them within the range of possible human exploitation (Sidell, 1993).

Site distribution Except for some isolated individuals (see below), no significant clustering of avifaunal remains was detected at Khonkho Wankane. In one context, a largely complete skeleton of Gallinago cf. andina was recovered. This Charadriiformes species inhabits the Khonkho Wankane area, and the presence of these remains could represent an isolated instance of natural mortality combined with rapid burial to allow such excellent preservation. This individual placement also could have resulted from deliberate human action. This behavior more likely would involve ritual burial instead of dietary usage, due to the completeness of these remains.

Human utilisation Dietary and raw material usage As described above, a Nycticorax nycticorax distal tibiotarsus was severed circumferentially from its shaft, possibly for the creation of a bone tube. Such bone tubes would have had several possible uses, including storage of small implements such as needles, portions of musical instruments and storage of lime for coca chewing (Webster & Janusek, 2003). The leg elements of large wading birds are highly suitable for use as a raw material in this manner, as they are long, straight and have a large interior space. There is of course no reason why the same individual prey item could not have been used first for its meat and plumage in addition to raw material usage. Five other avian elements bear cut marks, which may be interpreted as signs of butchery/ dismemberment. Each of the other avian elements with cut marks is from a taxon of sufficient size to have been a plausible contributor to human diet: Colaptes cf. rupicola, Oxyura jamaicensis, Nothura cf. darwinii and unknown Size Class C. The elements involved (two humeri, two ulnae and an additional tibiotarsus) are associated with meaty portions of the avian skeleton, so their involvement in butchery activities is likely. Int. J. Osteoarchaeol. 23: 746–756 (2013)

Avian Remains from Khonkho Wankane The necessary transport of at least the majority of the lake/wetlands avifauna from Lake Titicaca to Khonkho Wankane indicates probable human action and the inclusion of this resource in human diet. These avifauna are also relatively large and would provide a significant dietary resource relative to the energy expended to acquire them. The hunting techniques employed in taking these species are unknown, but projectile or netting techniques are plausible options (Moore et al., 1999). Human dietary or other direct usage relating to the egg shells discovered at Khonkho Wankane is probable due to their size: where a nesting passerine may have gone unnoticed, it is unlikely that any bird species producing such large shells could have nested among human habitations. Since the eggs were so large, they are most likely from the lake/wetland environment, since the avifauna in this area are of sufficient size to produce these eggs. Ceremonial usage Evidence of direct ceremonial use of birds is present at Khonkho Wankane in the form of multiple (probable aplomado) falcon (Falco cf. femoralis) burials. These two concentrated (NISP = 10, MNI = 1 and NISP = 81, MNI = 2), largely complete and partially articulated clusters of Falco cf. femoralis elements account for 91 of its total NISP of 103. These were all adult individuals with no visible signs of perimortem trauma to their skeletons, so the manner of their capture and/or dispatching is unknown. Neither of these concentrations (see Figure 2 for an example) was associated with human burials, but they were associated with structures,

753 indicating their possible origin as ceremonial offerings. Additional, mostly isolated elements of this taxon were found in multiple other contexts, contributing to the total MNI of 16. This predator species would have proven difficult to hunt, but it may have been taken by projectile weapons while roosting. It is unknown if juveniles were captured alive and then raised to adulthood for ritual purposes, but this possibility cannot be ruled out. Falco femoralis is a relatively common raptor throughout its Neotropical range, and it would have been seen hunting small prey commonly throughout the open areas around Khonkho Wankane. It is noteworthy that the Portugal Stela at Khonkho Wankane has one of the few Formative period avian depictions in stone and depicts an upward-facing predatory bird of unknown taxon. Other recognisable predatory bird motifs are common throughout southern Lake Titicaca iconography during this period and the Tiwanaku period, appearing in three-dimensional representations in ceramic vessels and in other artistic contexts (Janusek, 2003). Some are close enough in design that they may represent Falco femoralis or similar species, such as eagles. Given the prevalence of diurnal raptors in the iconography, the inclusion of falcons as burial offerings is of probable high ritual significance. Waterfowl (apparently ducks) were also important enough to show up in Tiwanaku iconography (Janusek, 2003). This motif was commonly portrayed on vasijas (small pitchers or similar ceramic vessels) as were llamas during Tiwanaku IV and later. Sometimes, vasijas were fashioned in effigy shapes, which included ducks. Since fish from the lake also were exploited in significant amounts, these line evidence indicate that the fauna of Lake Titicaca were an important dietary resource to the inhabitants of Khonkho Wankane and played a significant cultural role as well.

Paleoenvironment and paleoecology: Khonkho Wankane and Chiripa

Figure 2. Falco cf. femoralis burial at Khonkho Wankane.


Comparison with another site in the Lake Titicaca region in terms of relative lake resource utilisation is possible. Chiripa is a primarily a Formative period (ca. 1500–100 BC) site adjacent to Lake Titicaca, excavated in the 1990s by a multidisciplinary team (Kent et al., 1999; Moore et al., 1999; Hastorf, 2003). These excavations yielded a large, diverse avifaunal sample (Table 4). Figure 3 shows the proportions of avifaunal NISP from Khonkho Wankane and Chiripa. The Khonkho Wankane avifauna shows a broader, more diverse spread among habitat types of wetlands, dry grasslands and cosmopolitan taxa. Unsurprisingly, Int. J. Osteoarchaeol. 23: 746–756 (2013)

J. T. Pokines

754 Table 4. Chiripa avifauna (Kent et al., 1999) grouped by primary habitat type Taxon

NISP

Podicipedidae 110 Phalacrocorax brasilianus 5 Nycticorax nycticorax 2 Phoenicopteridae 19 Chloephaga cf. melanoptera 1 Anas flavirostris 3 Anas puna 1 Anas georgica 7 Anas sp. 10 Oxyura jamaicensis 18 Rallidae 340 Himanotopus mexicanus 1 Larus serranus 2 519 Vanellus resplendens 6 Tinamidae Metropelia ceciliae Metropelia aymara Metropelia sp. Colaptes rupicola Passeriformes cf. Bolborhyncus aurifrons Ciccaba cf. albitarsus Trogon sp. cf. Buteo sp. Tyto alba Total Primary Habitat Wetlands Wetlands/dry grassland Dry grassland Forest Cosmopolitan Total

Primary Habitat lake/wetlands lake/wetlands lake/water margins shallow lake lake margin/wetlands lake/wetlands lake/wetlands lake/wetlands lake/wetlands lake/wetlands lake/wetlands lake margin/wetlands lake margin/wetlands wetlands/dry grassland

34 dry grassland 5 dry grassland 9 dry grassland 5 dry grassland 5 dry grassland 21 dry grassland 79 1 forest/scrub 5 Forest 2 (probable lowland import) 8 13 Cosmopolitan 4 Cosmopolitan 17 629 %NISP 82.5% 1.0% 12.6% 1.3% 2.7% 100.1%

Figure 3. Khonkho Wankane and Chiripa avifaunal NISP by habitat type.


given its location on the lake shore, the Chiripa avifauna is dominated by wetland taxa. The large cosmopolitan habitat component at Khonkho Wankane, however, is dominated by Falco cf. femoralis remains which were introduced into the deposits by deliberate burial, which inflates the NISP total for this species (Table 1). It also is not indicative of a single habitat type. Excluding cosmopolitan species, those inhabiting both wetlands and dry grassland, and the minor forest component from Chiripa, the following proportions are obtained: Khonkho Wankane, 141 wetlands to 91 dry grassland (1.5:1 ratio); Chiripa, 519 wetlands to 79 dry grassland (6.6:1 ratio). These ratios also may reflect in part the effects of natural mortality of avian species living adjacent to these two sites. They also reflect at least in part the variation in exploitation by humans of birds. Despite a reliance upon camelids as a dietary mainstay at Khonkho Wankane (Gladwell, 2010) and its distance from the Titicaca wetlands environment, wetland bird species remained a significant component of dietary exploitation. The proportions of wild fauna in Chiripa diet may also be elevated due to its earlier occupation and a lack of greater reliance upon the agricultural developments (Bruno & Whitehead, 2003) of the later Tiwanaku state. The lake environment also yielded fish as an important resource at Khonkho Wankane and in much higher recovered amounts at Chiripa (Moore et al., 1999; Capriles et al., 2008).

Summary and conclusions The Khonkho Wankane avifaunal remains constitute a diverse community with a minimum of 26 avifaunal taxa represented, reflecting both the surrounding dry puna environment and taxa imported from the Lake Titicaca and its surrounding wetlands ecosystem, natural or artificial in the form of raised fields. The wetland avifaunal taxa also may have been more direct visitors to the Khonkho Wankane environs due to the creation of artificial wetlands in the form of multiple qochas in the otherwise dry puna surrounding this site. Human dietary usage is likely for the majority of the large species, which come largely from the lake or artificial wetlands environment. Bird skeletons also served as a useful raw material. One species with a cosmopolitan distribution, Falco cf. femoralis, is highly represented (NISP = 103) due to its inclusion in the site deposits of at least three deliberate burials of largely complete skeletons. Additional scattered remains of this species were recovered, indicating that other burials may have been present. Raptorial and wetland birds figure prominently in Tiwanaku iconography, Int. J. Osteoarchaeol. 23: 746–756 (2013)

Avian Remains from Khonkho Wankane and the cultural importance of this relationship is enhanced by the findings here. These analyses must also expand to other sites in the Altiplano region, so that more comparisons can be made among sites to understand the paleoecological relationships of each site to its surroundings, natural and artificial. What is clear is that despite the urbanised setting of Khonkho Wankane and its other contemporaneous Altiplano sites, wild fauna continued to play an important role, and Lake Titicaca resources were acquired even when other domesticated resources that would have filled the same role were not (Bermann, 1994; Kent et al., 1999; Moore et al., 1999; Pérez Arias, 2007; Janusek, 2008; Pokines et al., 2009; Pokines, 2011). How much of this was due to alterations of the landscape through the creation of artificial wetlands (Binford et al., 1997) that in turn increased the biomass of wetland resources (including both fish and waterfowl) requires additional research.

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 thanks Julian Kerbis Peterhans and William Stanley (Division of Mammals) and John Bates, Tom Gnoske, and David Willard (Division of Birds), Field Museum, Chicago for their invaluable research advice and access to comparative osteological collections. The author also thanks the three anonymous reviewers of this manuscript for their valuable comments.

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