Bone Mineral Density of Wild Turkey (Meleagris gallopavo) Skeletal Elements and its Effect on Differential Survivorship
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Journal of Archaeological Science (2001) 28, 817–832
doi:10.1006/jasc.2000.0600, available online at http://www.idealibrary.com on
Bone Mineral Density of Wild Turkey (Meleagris gallopavo)
Skeletal Elements and its Effect on Differential Survivorship
Frank J. Dirrigl Jr*
Department of Anthropology, 344 Mansfield Road, U-2158, University of Connecticut, Storrs, CT 06269-2158,
U.S.A.
(Received 9 November 1999, revised manuscript accepted 14 August 2000)
Zooarchaeologists recognize that density-mediated attrition is a bias that demands consideration in the analysis of
archaeofaunal samples. This paper presents bone mineral density values (aereal and volumetric) for the wild turkey
(Meleagris gallopavo) and represents the first application of Dual-energy X-ray Absorptiometry (DEXA) to birds. The
relationship between density and the survivorship of avian skeletal elements and portions is demonstrated by examining
20 assemblages from the archaeological record of the northeastern United States. The results suggest that density may
account for the differential survivorship of least 35% of the samples tested. Further analysis of a single site demonstrates
the importance of examining bone mineral density when interpreting past bird hunting and use.
2001 Academic Press
Keywords: BONE MINERAL DENSITY, DENSITOMETER, GALLIFORMES, TAPHONOMY, WILD
TURKEY, ZOOARCHAEOLOGY.
Introduction that the last approach is preferable (see Lyman,
Houghton & Chambers, 1992 re-examination of
A
rchaeologists share a special interest in study- Grayson, 1989). Most recent measurements of bone
ing bone mineral density to assess its affect on mineral density focus on methodology, improving
the survivorship and representation of bone analysis, and application (Pavao, 1996; Lam et al.,
elements and portions. For years, researchers have 1998a,b; Higgins, 1999; Pavao & Stahl, 1999; Stahl,
recognized that bone mineral density is one tapho- 1999). Dirrigl (1998: 16–20) presents a review of clini-
nomic bias affecting the survivorship of the remains cal and archaeological studies of bone mineral density
of vertebrate animals (e.g., Brain, 1967, 1969; and the different methods used.
Behrensmeyer, 1975; Binford & Bertram, 1977; There is no doubt that bird skeletal elements and
Lyman, 1982, 1984, 1993, 1994; Butler, 1990, 1996; portions differ in their survivorship potential in both
Nicholson, 1991, 1992; Elkin & Zanchetta, 1991; paleontological and archaeological contexts (Mayr,
Stewart, 1991; Chambers, 1992; Kreutzer, 1992a, 1946; Dawson, 1969; Ericson, 1987; Bjordal, 1987;
1992b; Lyman, Houghton & Chambers, 1992; Van Livingston, 1989). The decomposition and survivor-
Houten, 1992; Butler & Chatters, 1994; Elkin, 1995; ship of bird remains tends to follow recognizable
Pavao, 1996; Farquharson, Speller & Brickley, 1997; patterns (Schäfer, 1972; Bickart, 1984) affecting
Hindeland & MacLean, 1997; Dirrigl, 1998; Higgins, archaeological visibility and identification (Ericson,
1999; Pavao & Stahl, 1999; Stahl, 1999). These studies 1987). Not surprisingly, zooarchaeologists suggest that
of bone mineral density appear in several forms. In bone mineral density may be a potential bias resulting
early studies, archaeologists assumed that the bones of in the patterning of bird bone assemblages (Rich, 1980;
one vertebrate animal class were denser than another Livingston, 1989; Nicholson, 1991, 1996). Further-
(e.g., Gifford, 1981). Next, archaeologists used bone more, Livingston (1989: 540) suggests that the bone
mineral values of animals within the same vertebrate density of bird elements may play ‘‘an overriding
class (e.g., Grayson, 1989). Later, archaeologists exam- role in element survivorship, regardless of agents of
ined bone mineral values for a number of skeletal deposition, movement, or attrition.’’
elements of a single animal or a closely related group No longer can zoologists and archaeologists assume
(e.g., Lyman, 1982, 1984). Zooarchaeologists agree that bird skeletons are lighter than those of mammals
(Cubo & Casinos, 1994), and that their survival poten-
*Correspondence to: Frank J. Dirrigl Jr, E-mail:
fdirrigl@teikyopost.edu. Present address: Environmental Theories
tial is therefore low. Although bird appendicular skel-
and Applications Program, Teikyo Post University, 800 County etal elements exhibit positive allometry (i.e., the mass
Club Road, PO Box 2540, Waterbury, CT 06723-2540, U.S.A. of bone increases with animal size), the avian humerus,
817
0305–4403/01/080817+16 $35.00/0 2001 Academic Press818 F. J. Dirrigl Jr
ulna-radius, and tibiotarsus-fibula are known to have a examination to four skeletal specimens of wild turkey.
greater mass than insectivore or rodent counterparts However, this small sample size is applicable because
(Cubo & Casinos, 1994). For example, avian femora the purpose of taking the measurements was not to
have a mass similar to mammalian femora (Cubo & develop definitive measures, but to obtain ordinal
Casinos, 1994). These observations merit a new measures that would allow the ranking of skeletal
assessment of bird bone survivability. elements and portions.
At present, the literature contains only limited The dermestid beetle method of skeletal preparation
clinical or archaeological applications of assess- (see Dirrigl, 1988 for a review) is useful in studies of
ing bird bone mineral density (e.g., Nicholson, 1991; bone mineral density. Although bones fixed in formalin
Leznicka, 1992; Dirrigl, 1998; Higgins, 1999). Whereas can be cleaned with dermestids (Dirrigl, Dubos &
Nicholson (1991) examined organic:mineral ratios for Rusch, 1993), the fixation will affect bone hardness and
a single domestic pigeon (Columba livia), analysis of measures of bone mineral content or density (Kann,
bone mineral density for a group of closely related Piepkorn & Beyer, 1997). The use of dermestids to
birds using multiple specimens is preferable (Dirrigl, remove the flesh of fresh specimens does not affect
1998; Higgins, 1999). Higgins (1999) used water dis- measurements (Hefti et al., 1980) or produce errors
placement to measure density values in specimens of associated with the presence of intraosseous fat after
Anatidae and Podicipedidea. museum specimens have thoroughly dried (Lahtinen,
Because gallinaceous birds (turkey; grouse; hen; Vaananen & Karjalainen, 1980; Lahtinen, pers.
pheasant; quail) are archaeologically important taxa comm.). In vitro DEXA measurements may also
(Senior & Pierce, 1989; MacDonald, 1992; MacDonald result in less reproducibility error than with in vivo
& Edwards, 1993), my study will be useful in providing (Rozenberg et al., 1995).
scales to determine when density-mediated attrition is Bone mineral density varies intra- and inter-
among the non-cultural factors responsible for the specifically with age, sex, nutrition, and genetics
patterning of skeletal elements and portions among (Lyman, 1982). For birds, sex is an important consid-
archaeological samples of wild turkey in Mexico, the eration because of the presence and absence of medul-
United States, and Canada. The purpose of this paper lary bone by females. The medullary bone of females
is fourfold. First, I review a modified method for occurs in all birds of breeding age, and its occurrence
assessing the bone mineral density of bird skeletons (only observed internally in bone cross-sections) varies
using a dual energy X-ray bone densitometer (DEXA). with reproductive activity and eggshell formation
Second, I present the first baseline information for (Simkiss, 1975; Dacke et al., 1993). Because the forma-
bone mineral density of the wild turkey (Meleagris tion and reabsorption of medullary bone during egg
gallopavo). Third, I use this data to develop a rank laying periods potentially affects mineral density, I
order scheme and evaluate how bone mineral density examined only male specimens. Furthermore to avoid
effects the differential survivorship of wild turkey any age (immature versus mature) or pathological
remains in archaeological assemblages from the north- (diseased versus healthy) biases, I examined only adult
eastern United States. Lastly, I use a single site assem- specimens (following Hargrave (1972) based on the
blage to demonstrate the utility and implications of the sternum, furcula, coracoid, pelvis, and tarsometa-
density data to understanding the past behaviour of tarsus) showing no indication of pathological disease.
Native Americans. Dual-energy X-ray absorptiometry (DEXA) is be-
coming more popular among zooarchaeologists to
examine bone mineral content (BMC) and density
Material and Methods (BMD) (Kreutzer, 1992a, b; Butler & Chatters, 1994;
Pavao & Stahl, 1999). Lyman (1982, 1984, 1994)
Using DEXA to measure bone mineral density reviews the ambiguity among different measures of
At the Osteoporosis Center, University of Connecticut bone density applied to archaeological bone remains.
Health Center, I used a Lunar DPX-L Dual Energy Critiques of using DEXA in archaeological applica-
X-ray Bone Densitometer (Lunar Corporation, Madi- tions question: its inability to distinguish flat, mineral
son Wisconsin) to measure bone mineral density. I rich from thick, mineral poor bone; ambiguity among
scanned museum appendicular skeletons of wild turkey elements of varying shape; and, dependency on bone
(Meleagris gallopavo) (Total examined, N=4. New orientation during scanning (Butler & Chatters, 1992;
York (N=3): Cornell University (CU) 48300, 8528, Kreutzer, 1992b). Although Sievanen, Kannus &
48560; South Carolina (N=1): American Museum of Jarviners (1994) reports that precision is good even
Natural History (AMNH) 5091), free of muscle and without modifying for small animal measurements
ligaments, and prepared with dermestid beetle larvae. (mammals), thin bird skeletons offer unique challenges
Due to the limits imposed on article length by JAS, the and require the use of Lunar’s Small Animal Software
data for other gallinaceous birds, bobwhite (Colinus Version 1.0e, August 1994. Lunar developed this
virginianus), ruffed grouse (Bonasa umbellus), and prai- software for laboratory animals (e.g., rats) of
rie hen (Tympanuchus cupido), may be found in Dirrigl lengthsBone Mineral Density of Wild Turkey 819 The following procedure is condensed from that pre- I placed ROIs over the scanned images, and manu- sented in Dirrigl (1998), and a future paper will provide ally placed each vertex to produce the largest specific details on using DEXA to measure the bone quadrilateral within the boundaries of the bone. Alter- mineral density of birds. Before measurements were natively, the shape of the ROI was altered manually to taken, the DPX-L was calibrated daily with the Quality fit the contours of problematic bones such as the Assurance Test (% c.v.
820 F. J. Dirrigl Jr
Figure 1. a–f.Bone Mineral Density of Wild Turkey 821
Figure 1. g–k.
Figure 1. Regions of Interests examined for gallinaceous bird skeletal elements. Drawn from Tympanucus cupido (YPM 349). (a) Femur; (b)
Tibiotarsus; (c) Tarsometatarsus; (d) Humerus; (e) Ulna; (f) Radius; (g) Carpometacarpus; (h) Coracoid; (i) Scapula; (j) Pes phalanx; (k) Manus
phalanx. (From Dirrigl, 1998).822 F. J. Dirrigl Jr
Table 1. Bone mineral density of Meleagris gallopavo
BMDa (g/cm2) BMDv (g/cm3)
Fossil class Min Max Mean .. Min Max Mean ..
Proximal femur 0·359 0·528 0·477 0·076 0·126 0·272 0·226 0·065
Medial femur 0·446 0·632 0·568 0·081 0·413 0·555 0·486 0·065
Distal femur 0·390 0·505 0·466 0·050 0·233 0·274 0·255 0·017
Proximal tibiotarsus 0·422 0·564 0·498 0·059 0·193 0·246 0·222 0·022
Medial tibiotarsus 0·489 0·683 0·594 0·083 0·527 0·763 0·627 0·102
Distal tibiotarsus 0·480 0·692 0·623 0·094 0·274 0·439 0·352 0·068
Proximal tarsometatarsus1 0·472 0·587 0·530 0·058 0·366 0·427 0·397 0·030
Medial tarsometatarsus1 0·496 0·791 0·644 0·148 0·719 0·904 0·812 0·092
Distal tarsometatarsus1 0·327 0·422 0·375 0·047 0·569 0·590 0·580 0·011
Pes, Phalanx 1, Digit 31 0·328 0·462 0·395 0·067 0·297 0·548 0·423 0·126
Proximal humerus 0·296 0·420 0·374 0·054 0·179 0·234 0·213 0·024
Medial humerus 0·394 0·566 0·491 0·075 0·335 0·496 0·419 0·072
Distal humerus 0·306 0·438 0·396 0·059 0·220 0·353 0·292 0·055
Proximal ulna 0·342 0·477 0·408 0·055 0·200 0·281 0·241 0·034
Medial ulna 0·420 0·644 0·517 0·100 0·437 0·708 0·557 0·115
Distal ulna 0·353 0·499 0·433 0·060 0·276 0·391 0·326 0·053
Proximal radius 0·269 0·353 0·321 0·038 0·439 0·510 0·471 0·030
Medial radius 0·301 0·435 0·361 0·061 0·528 0·737 0·628 0·089
Distal radius 0·233 0·316 0·283 0·035 0·272 0·365 0·320 0·041
Proximal carpometacarpus 0·374 0·521 0·468 0·065 0·331 0·445 0·402 0·050
Medial carpometacarpus 0·470 0·675 0·568 0·092 0·558 0·722 0·628 0·069
Distal carpometacarpus 0·286 0·484 0·405 0·086 0·515 0·787 0·680 0·122
Manus, phalanx 1, Digit 2 0·196 0·262 0·223 0·028 0·272 0·341 0·297 0·030
Proximal coracoid1 0·438 0·454 0·446 0·008 0·370 0·389 0·380 0·009
Medial coracoid1 0·428 0·472 0·450 0·022 0·340 0·375 0·358 0·018
Distal coracoid1 0·268 0·304 0·286 0·018 0·282 0·292 0·287 0·005
Proximal scapula1 0·253 0·298 0·276 0·022 0·230 0·311 0·271 0·040
Medial scapula1 0·252 0·292 0·272 0·020 0·654 0·664 0·659 0·005
1
Based on two specimens.
Stahl, 1999; Stahl, 1999). Because a ROI represents a of density-mediated attrition (Rapson, 1990; Kreutzer,
region of varying bone thickness (BT), I calculated the 1992a, b; Lyman, 1993, 1994). Lyman (1992, 1993)
mean BT of a skeletal element and portion as the proposed that the greatest measured density value for
greatest thickness and least thickness (measured with an anatomical region is a better choice. I found that
dial calipers) divided by two. My approach conforms the relationship between maximum and mean values
to Lyman (1982, 1984) and Kruetzer (1992a, b) while of BMDa (r=0·98, P=0·001) and BMDv (r=0·98,
recognizing the contributions of previous researchers P=0·001) for wild turkey to be a high positive and
(Elkin, 1995; Pavao, 1996; Galloway, Willey & Snyder, significant correlation. These results suggest that either
1997; Pavao & Stahl, 1999; Stahl, 1999). the mean or maximum values of BMDa, and BMDv
are applicable in developing ranking schemes for my
study of density-mediated attrition.
Results and discussion Because the DPX-L measures areal bone density
(BMDa), only approximations of true physical density
Measures of bone mineral density or volume bone mineral density (BMDv) are possible
This study compiled data for 28 fossil classes of M. (Blake & Fogelman, 1996). Thus, researchers also
gallopavo and is drawn from my examination of gall- debate over whether BMDa or BMDv provides the
inaceous birds that included 420 analysed ROIs from best measure of bone mineral density (Kellie, 1992;
161 scanned images saved to computer files (Dirrigl, Carter, Bouxsein & Marcus, 1992; Cummings et al.,
1998). Table 1 provides both mean and maximum 1994). Although measurements of BMDa values can be
values of BMDa and BMDv. I found the mean of compared between animals of the same taxa and
BMDa means to be 0·443 g/cm2 with a range of means different sizes (Ho et al., 1990; Mitlak, Schoenfeld &
from 0·223 (manus, phalanx 1, digit 2) to 0·644 g/cm2 Neer, 1994), archaeologists tend to prefer using BMDv
(medial tarsometatarsus). The mean of volume bone measures to compare different skeletal elements or
mineral density means (BMDv) was 0·427 g/cm3 with a portions and predict survivorship. This view originates
range of 0·213 (proximal humerus) to 0·812 g/cm3 from the positions that: (1) areal determinations
(medial tarsometatarsus). (BMDa) underestimate bone mineral density; and (2)
Archaeologists debate the utility of using mean or intra- and inter-taxonomic comparisons of areal densi-
maximum values of BMDa and BMDv in their studies ties between ROIs, skeletal elements, or portions (e.g.,Bone Mineral Density of Wild Turkey 823 mid shafts) are inappropriate (Lyman, 1982, 1984; bird bone (Dirrigl, 1998; Higgins, 1999). Perhaps the Marean & Spencer, 1991; Kreutzer, 1992a; Pavao, best solution to examining survivorship among bird 1996; Galloway, Willey & Snyder, 1997; Marean & skeletons involves assessments that consider both Frey, 1997; Lam et al., 1998a, b; Marean, 1998; BMDa and BMDv measures rather than relying on Marean & Kim, 1998; Pavao & Stahl, 1999; Stahl, only one. 1999). Kreutzer (1992a: 282–283) took the strongest stand and described linear density (i.e., areal density or BMDa) as ‘‘misleading,’’ ‘‘at best ambiguous and at Assessment of the effect of bone mineral density on worst invalid,’’ and ‘‘of dubious value.’’ differential survivorship Alternatively, researchers are beginning to prefer Four assumptions are fundamental in assessing the methods using visual imaging which do not underesti- effect of bone mineral density on the differential survi- mate areal bone mineral density: computer aided tom- vorship between wild turkey samples. First, denser ography (CAT), dual-energy quantitative computer bone, larger bones, and those bones with the greatest tomography (QCT), computer mapping and optical tensile strength possess a greater potential for survival density, radiographic microdensitometry, spectropho- (Grayson, 1979; Klein & Cruz-Uribe, 1984; Klein, tometery, using a modified EMI brain scanner, and 1989; Marean, 1991). Second, ‘‘mechanical and chemi- low angle x-ray scattering techniques (LAXS) (Pullan cal attrition should have a greater effect on bones with & Roberts, 1978; Phillips, Owen-Jones & Chandler, low bulk density (high porosity)’’ (Lyman, 1994: 239). 1978; Lees, Healy & Cleary, 1979; Turnlund & Third, bones of similar density, volume, and surface Margen, 1979; Kapila et al., 1994; Karkhanehchi area decay similarly (Binford & Bertram, 1977). Based et al., 1996; Farquharson et al., 1997; Farquharson & on these assumptions, I adopt Lyman’s model for Brickley, 1997; Lam et al., 1998a, b). The new prospect Artiodactyla (Lyman, 1982, 1984) which proposes that of using broadband ultrasound attenuation (BUA) to the most dense skeletal elements and portions have the measure BMD also holds promise (Duquette et al., greatest potential for surviving attritional processes 1997). Because these methods generate 3D visual (resulting in the variability of skeletal remains images of scans and measure areal and volumetric reported). density concurrently, potential errors produced by One workable, approach for assessing the variability physical measurements of bone thickness and the of wild turkey remains is to assess the effects of bone DEXA underestimating BMDa values, especially mineral density on differential survivorship by consid- across cross-sections, are avoided. ering both BMDa and BMDv measures. If denser Regardless of the proposed methods, inherent prob- bones of wild turkey have the greatest potential for lems still exist and there is no single solution to the survivorship, then fossil classes with the highest ranks BMDa versus BMDv debate. Although the benefits have the greatest potential for surviving. For example, from these different methods of compensation are the medial tibiotarsus of M. gallopavo could be consid- appealing or desirable, areal (BMDa) and volume ered to survive decomposition better than the proximal (BMDv) measures of bone mineral density possess humerus, based on its BMDa and BMDv ranks similar value in predicting clinical studies of bone (Table 2). deterioration in living skeletons (Cummings et al., Table 2 provides scales of BMDa and BMDv rank- 1994). In my study of bone mineral density of wild ing the bone mineral density of 28 fossil classes for M. turkey skeletons, the results question the applicability gallopavo. These scales depend on the mean bone of BMDv to measuring several specific bird bones, mineral density values scanned for their proximal, especially those skeletal elements or portions that are medial, and distal portions. The medial tarsometatar- long, moderately wide, and thin (e.g., medial scapula; sus and distal and medial tibiotarsus ranked highest in distal coracoid). The inflated BMDv values of the BMDa, whereas the proximal and medial scapula and medial scapula originate from dividing values of areal manus ranked lowest. For BMDv, the medial tar- bone mineral density by values of low thickness sometatarsus, distal and medial carpometacarpus (Dirrigl, 1998: 67–70) rather than the inability of ranked highest, whereas proximal femur, proximal DEXA to discriminate high from low-density bones. tibiotarsus, and proximal humerus ranked lowest. My For example, the visual DEXA scan of a wild turkey results support Rich’s (1980) presumption that bird scapula (Dirrigl, 1998: 45 figure 2.4) shows how bone tarsometarsi are of high structural density. density decreases (displayed as dissipating pixels) from Overall, the ranking of skeletal elements and por- the proximal to distal portion of the scapula. This tions differed between BMDa and BMDv. I found the result is repeated for Colinus virginianus, Bonasa relationship (Figure 2) between mean values BMDa umbellus and Tympanuchus cupido (Dirrigl, 1998). and BMDv to be weak positive and insignificant Therefore, I consider the BMDv values of the medial (r=0·27, P=0·19). However, several bones were either scapula to be suspect, and these ROIs should not be close in rank (e.g., proximal coracoid and medial ulna) considered in examining differential survivorship of or equal in rank (e.g., medial tarsometatarsus and these gallinaceous birds. Additional biases result from medial carpometacarpus). When I removed the poten- the pneumaticity and presence of fossa and foramen in tial effects of the medial scapula, the relationship
824 F. J. Dirrigl Jr
Table 2. Rank-ordering and pooled frequency (f) of BMDa and BMDv from least (1) to most (28) dense for
Meleagris gallopavo fossil classes
Rank BMDa F BMDv F
Least dense
1 Manus 5 Proximal humerus 1
2 Medial scapula 2 Proximal tibiotarsus 1
3 Proximal scapula 0 Proximal femur 1
4 Distal radius 1 Proximal ulna 1
5 Distal coracoid 2 Distal femur 1
6 Proximal radius 0 Proximal scapula 0
7 Medial radius 2 Distal coracoid 2
8 Proximal humerus 1 Distal humerus 2
9 Distal tarsometatarsus 2 Manus 5
10 Pes 28 Distal radius 1
11 Distal humerus 2 Distal ulna 0
12 Distal carpometacarpus 1 Distal tibiotarsus 4
13 Proximal ulna 1 Medial coracoid 9
14 Distal ulna 0 Proximal coracoid 4
Subtotal 47 32
Most dense
15 Proximal coracoid 4 Proximal tarsometatarsus 0
16 Medial coracoid 9 Proximal carpometacarpus 1
17 Distal femur 1 Medial humerus 4
18 Proximal carpometacarpus 1 Pes 28
19 Proximal femur 1 Proximal radius 0
20 Medial humerus 4 Medial femur 3
21 Proximal tibiotarsus 1 Medial ulna 8
22 Medial ulna 8 Distal tarsometatarsus 2
23 Proximal tarsometatarsus 0 Medial tibiotarsus 9
24·5 Medial femur 3 Medial radius 2
24·5 Medial carpometacarpus 4 Medial carpometacarpus 4
26 Medial tibiotarsus 9 Medial scapula 2
27 Distal tibiotarsus 4 Distal carpometacarpus 1
28 Medial tarsometatarsus 12 Medial tarsometatarsus 12
Subtotal 61 76
between BMDa and BMDv remained similar (r=0·36, contexts in mulitcomponent sites (Grayson, 1984). One
P=0·087). inherent problem with assessing density-mediated attri-
To examine the regional effect of density-mediated tion from reports of archaeofaunal samples is that
attrition on the representation of birds, I compiled data skeletal elements and portions are quantified rather
from archaeological samples (Table 3) with a prefer- than the presence of scan sites (Lyman, 1982, 1984,
ence towards the northeastern United States (Dirrigl, 1993; Pavao & Stahl, 1999). My use of regions of
1998). These data are drawn from the literature and my interest (ROI) for the proximal, medial, and distal
personal database of over 140 samples. Zooarchaeolo- portions of bird skeletal elements avoided this prob-
gists differ in their reporting of animal remains (Butler lem. For inventories of complete elements or for those
& Lyman, 1995), and therefore a limitation of this specimens in which researchers did not identify the
study is the exclusion of samples for which data skeletal portion, I followed Lyman (1982, 1984) by
regarding skeletal elements and their quantification are applying the medial ROI. Because the medial portions
absent, unavailable for analysis, or incomplete. I agree of long bones of both mammal (Marean & Spencer,
with Lyman’s (1993) assertion that studies of bone 1991; Lam et al., 1998a, b) and birds (Dirrigl, 1998)
mineral density attrition are plagued by the lack of this tend to be more dense and less likely to be affected by
information. Nonetheless, I believe the samples I se- density-mediated attrition, I recognize the potential
lected for examination are representative broadly of bias in the ranking of fossil classes by adopting
the variability that can occur in the archaeofaunal Lyman’s convention. Additionally, the anonymous
record of the Northeast (Dirrigl, 1991, 1998). reviewer of my paper cautioned: (1) that there may be
Whenever possible, I recorded the number of ident- less of a chance at recovering a ROI than a scan site;
ified specimens (NISP) (Payne, 1975; Grayson, 1979) and (2) that the recognition of a ROI as recovered in
by skeletal element and portion for preserved wild an assemblage must account for the percent region
turkey remains (see Dirrigl, 1998). Whenever feasible, I surviving (see Marean, 1991). All of these concerns
separated aggregated samples to correct for the effects instill the need for future studies of bird bone mineral
of ‘‘lumping’’ the faunal material found from several density to examine how different methodologicalBone Mineral Density of Wild Turkey 825
0.7 samples of wild turkey, because most fossil classes were
represented by fewer than three specimens or had an
equal numbers of surviving fossil classes (Dirrigl,
0.6 1998). I considered a fossil class to be ‘‘dense’’ if their
mean BMDa or mean BMDv values were greater or
0.5 equal to the value at which the frequency distribution
BMDa
skewed to the higher values. A stem and leaf plot of
BMDv values for M. gallopavo exhibited a median
0.4
value of 0·389, and the distribution of the values
skewed to the right, decreasing in interval frequency
0.3 after BMDv=0·397 (proximal tarsometatarsus). The
proximal tarsometarsus ranks 15th among fossil
classes, and therefore I considered values to represent
0.2
0 1 2 3 4 high density bones if their rank were greater or equal to
BMC
15.
Likewise, I compared the occurrence of the fossil
classes to the average rankstandard deviation of the
0.9 average ranks. The average rank is
0.8 = f X/n.
0.7
The variance of these grouped data is
0.6
BMDv
0.5
0.4 where the standard deviation equals
0.3 s=ss2.
0.2
0 1 2 3 4 This method allowed me to examine the number of
BMC fossil classes ranking above and below the average
rank. I considered a sample to be affected possibly by
bone mineral density, BMDa or BMDv, if the number
0.9 of fossil classes ranking above the average rank was
higher than the number below.
0.8 Table 2 provides the frequency occurrence of wild
0.7
turkey bones recovered from 20 archaeological samples
examined in this study. Archaeological samples tended
0.6 to be dominated by medial tarsometatarsus remains
BMDv
that ranked highest in BMDa and BMDv measures.
0.5 However, pes (toe) remains also tended to occur more
frequently than other fossil classes. The high frequency
0.4 of pes remains may be explained biologically (i.e., if
BMDv is considered to be the best measure of bone
0.3
mineral density) or culturally, (e.g., resulting from the
deposition of feet after butchering and consumption
0.2 0.3 0.4 0.5 0.6 0.7 into features or disposal pits). Ericson (1987) also
BMDa found archaeological samples to be dominated by
Figure 2. Scatter-plots for the relationships between mean values of extremities.
BMC, BMDa, and BMDv of M. gallopavo. BMDa and BMDv may account for 45% (N=9) to
74% (N=14) respectively of the bird assemblages being
affected by differential survivorship. Although, the
approaches affect the results of measurements and rank order scale based on volumetric measure results in
interpretations. a greater number of assemblages associated with
I began this examination by adopting both graphical BMDv, I previously proposed that perhaps the best
and statistical methods to assess regional and site solution to examining the differential survivorship of
specific differential survivorship. I found graphical birds involves assessments considering both BMDa
representations to be appropriate for Northeastern and BMDv. This conservative procedure involved826 F. J. Dirrigl Jr
Table 3. List of assemblages and overall effect of BMDa and BMDv on survivorship of wild turkey remains
Overall effect on survivorship
May or
Sample Local Reference NISP May may not May not
Sylvan Lake New York det. F. Dirrigl 1 X
Van der Kolk New York det. F. Dirrigl 1 X
RI 1428 Rhode Island det. F. Dirrigl 2 X
Dogan Point New York Whyte, 1994; Claassen, 1995 4 X
Mon City Pennsylvania Church, 1994 14 X
Fish Club Cave New York det. F. Dirrigl 15 X
Woodruff Rockshelter Connecticut Swigart, 1987 15 X
Bates New York det. F. Dirrigl 1 X
Blundee Rockshelter Connecticut McBride, 1984 1 X
Bronck House Rockshelter New York det. F. Dirrigl 1 X
Hoffman Hideaway Connecticut det. N. Bellantoni 1 X
Ostungo New York Socci, n.d. 3 X
Mohantic Fort Mashantucket det. F. Dirrigl 3 X
Coudart Ledge Connecticut det. N. Bellantoni 4 X
Old Lyme Shell Heap Connecticut Amorosi, 1991 5 X
Slackwater Pennsylvania Custer et al., 1995 7 X
Greenwich Cove Rhode Island Bernstein, 1987 27 X
Elwood New York Socci, n.d. 1 X
Rabuilt Cave New York Vargo & Vargo, 1983 1 X
72–30 Connecticut det. N. Bellantoni 1 X
Total 7 10 3
applying three categories: (1) may (>50% of the fossil Although early excavations of New York sites did
classes surviving rank>15 for BMDa and BMDv); (2) not include screening in their recovery techniques,
may or may not (the surviving fossil classes for BMDa William Ritchie and Robert Funk paid special atten-
and BMDv show widespread differences among ranks); tion to recover as much bone as possible for analysis.
and (3) not (15 for BMDa and BMDv). Table 3 and their reports detailing the different taxa of inver-
presents these categories for 20 bird assemblages and tebrate and vertebrate animals represented (e.g., terres-
shows that the survivorship of over a third (35%, N=7) trial snail versus deer) in their assemblages, the
of the assemblages may be affected by bone mineral different sizes of animal bones recovered (e.g., frag-
density biases. This result demands that analyses of ments of freshwater mussel to complete bear mandi-
archaeological samples from the northeastern United bles), and their recognition of the value of faunal
States consider differential survivorship before cultural remains to their reconstruction of prehistoric lifeways
interpretations of bird use are developed. in New York (Ritchie, 1965; Ritchie & Funk, 1973;
Funk, 1976, 1993).
The assemblage of wild turkey remains (NISP=15)
Archaeological implications were identified by myself from the Archaeological
The relationship between bird bone mineral density Collection, New York State Museum, Albany. Because
and the survivorship of skeletal portions and elements the transportation of birds from kill to camp sites
and the implications of interpreting past bird hunting may not be an issue with birds (i.e., the entire carcass
and use can be demonstrated with a single archaeologi- can be carried easily), researchers begin by examin-
cal assemblage. Fish Club Cave (Funk, 1976) is located ing the representation and abundance of bird skeletal
in Coeymans Township, Albany County, New York. elements and portions to determine between natural
The cave exists on a hill 15 feet above and overlooking and cultural deposited remains and food or non-food
Hannacrois Creek, which flows into the Hudson River. use of birds (Schäfer, 1972; Hargrave, 1965, 1970;
William Ritchie and R. Arthur Johnson excavated a Rich, 1980; Ericson, 1987; Livingston, 1989; Senior &
total area of 36·6 m2 in 1962. The bone remains, that Pierce, 1989; Serjeantson, Irving & Hamilton-Dyer,
included wild turkey, were recovered from the human 1993; Serjeantson, 1997; Higgins, 1999). However,
occupation of Zone II. The definitive lithic identifica- Serjeantson, Irving & Hamilton-Dyer (1993) note
tions from this zone included the following points: that they found conflicting interpretations in their re-
Otter Creek, Vosburg, Normanskill, cf. Madison, view of aviafaunal studies which focus on bone distri-
Levanna, and Fox Creek stemmed. This cultural butions in their assessments.
sequence represents a Hudson Valley occupation of the The skeletal distribution of wild turkey remains
cave from the Late Archaic to Middle Woodland from Fish Club Cave represent a cultural deposition
(c. 4000 to 1000). resulting from food use for the following reasons.Bone Mineral Density of Wild Turkey 827 First, hindlimb elements (e.g., femora, tarsometatarsi, Conclusion and tibiotarsi) were more abundant than the fore- limb elements that survived (Ericson, 1987), however The bone mineral density of archaeological bone can Livingston (1989) and Serejeantson, Irving & be altered through variable diagenesis (Stout, 1978). In Hamilton-Dyer (1993) question the utility of this indi- humid climates, the loss of organic and inorganic cator. Second, tibiotarsi splints were absent, and all material from bone specimens is greater than speci- bones were disarticulated (Hargrave, 1965). Third, the mens recovered from arid climates (Salomon & Haas, humerus, ulna, and radius occurred together suggest- 1967). Dirrigl’s (1998) examination of wild turkey ing perhaps that the use of wings was non-ornamental paleontological samples (Steadman, 1980) concurs (Hargrave, 1970). Although cut mark, butchering, or with Salomon & Haas (1967). For example, bone preparation evidence were absent, which tends to mineral content and density correlated significantly imply non-food use (see Senior & Pierce, 1989; Dirrigl, with 43% (N=3) of the samples from Florida (Dirrigl, 1998: 166–167; Higgins, 1999), my examination of 1998: 117, table 3.5), perhaps the most humid of the bird remains in the northeastern United States finds locations reported. Studies combining bone anatomy this situation common. Even in the largest North or histology and the measurement of bone mineral American assemblages of wild turkey, the evidence density offer new and exciting areas of research of cultural modification may appear low. For (Nicholson, 1992, 1996; Higgins, 1999). Additionally, example, Corona (1997) found only 16% of the turkey while the bones of older animals may possess a greater remains (N=120) he examined to exhibit any cultural potential for survival; some bones (e.g., the scapula; modification. distal humerus; mandible; distal tibia of mammals) Previous archaeologists could not use DEXA values survive better regardless of age (Binford, 1981). In my to assess density-mediated survivorship of wild turkey archaeological study, I only examined, whenever poss- and to determine if this bias accounts for the represen- ible, the bones of adult male wild turkeys occurring in tation and abundance of bird bones. Traditionally, the the Northeast and surrounding areas to avoid the effect relationship between percent survivorship and struc- of the environment or age. tural density of bone is examined statistically (Lyman, Nicholson (1996) assessed the decomposition of 1982, 1994). It is possible to compute BMDa (r=0·35, domestic pigeon bone (Columba livia) in her exper- P=0·392) and BMDv (r=0·17, P=0·688) correlations imental burials. She found that bird bones had an for wild turkey bones from Fish Club Cave, and overall high survivorship, as suggested by mean skel- interpret these results as low positive and non- etal completeness at four sites. In contrast, Bickart significant. However, it is important to consider the (1984: 534) reported an anecdotal case where bird frequency distribution of skeletal elements before bones were ‘‘reduced from fresh to chalky, crumbly generating any conclusions (Hartwig & Dearing, bones within three weeks.’’ An experimental study in 1979; Drennan, 1996). Because the distribution is which wild turkey bones were measured for bone skewed with most skeletal elements or portions repre- mineral content and bone mineral density, buried for sented by two or fewer specimens, the application of several periods, and then reexamined would prove rank-order statistics to evaluate this assemblage is useful in assessing the speed and potential causes of erroneous. This scenario is common for assemblages of bone loss (see Davis, 1997). bird bones in the Northeast, and therefore the in- Ricklan (1986) warns that not all samples of bones terpretation of graphical representations is preferable are affected by density-mediated attrition. Studies dem- (Dirrigl, 1998). onstrate that archaeofaunal samples of birds would Graphical representation of eight fossil classes (Fig- be affected greatly by animal predation and scaveng- ure 3) shows that seven fossil classes ranked d15 for ing of bird carcasses (Rosene & Lay, 1963; Bickart, BMDa and sixd for BMDv. Because >50% of the 1984; Balcomb, 1986; Bramwell, 1987; Tobin & fossil classes for both BMDa and BMDv ranked d15, Dolbeer, 1990; Linz, 1991; Serjeanston, Irving & bone mineral density may account for the survivorship Hamilton-Dyer, 1993; Oliver & Graham, 1994). of the skeletal portions and elements represented. Potential scavengers of wild turkey include raccoon Further support is provided by presence of medial (Procyon lotor), grey (Urocyon cinereoargenteus) and tarsometarsi (BMDa and BMDv rank=28) and the red (Vulpes vulpes) foxes, and domestic dogs (Canis frequency (Figure 3) of medial ulnae (BMDa rank=22; familiaris) (Bickart, 1984). Balcomb (1986) reports the BMDv rank=21). Additionally, animal scavenging most striking example of bird bone loss resulting from may not be a bias affecting the frequency of bones at scavenging. Over five days, he found a 92% removal Fish Club Cave because tooth marks are absent. The of songbird carcasses in a sample of 78 carcasses. implication of my observations and findings is that, Removal of birds included either complete disappear- although the deposition of wild turkey remains results ance of the entire carcass or movement within five from cultural behaviour, the pattern of surviving skel- metres of initial deposition. Rosene & Lay (1963) etal elements most likely reflects density-mediated reported a similar situation for bobwhite quail (Colinus attrition rather than any cultural selection, use, or virginianus). In recognition of how predation and disposal of body parts. scavenging can effect samples of mammal and bird
828
F. J. Dirrigl Jr
Figure 3. Frequency of fossil classes recovered from Fish Club Cave (Funk, 1976, Dirrigl, 1998), Each ‘‘X’’ represents a single specimen.Bone Mineral Density of Wild Turkey 829
bones and mimic density-mediated attrition (Brain, (Peabody Museum of Natural History, Yale Univer-
1981; Livingston, 1989; Marean & Spencer, 1991), I sity); and Paul Sweet (American Museum of Natural
avoided samples in which archaeologists reported evi- History). Additional assistance with archaeological
dence of scavenging. In all samples I examined, I found collections was provided by Lisa Anderson (New York
no evidence of damage to wild turkey bones resulting State Museum), Susan Bruce (Peabody Museum of
from scavenging. However, this bias demands as much Archaeology and Ethnology, Harvard University),
attention as differential survivorship and differential Eric Johnson (Massachusetts Historical Commission),
identifiability in any future studies of Native American Barbara Leudtke (University of Massachusetts, Bos-
use of gallinaceous birds. ton), and Stephen Warfel and Janet Johnson (State
My recent review of the archaeofaunal record of the Museum of Pennsylvania, Section of Anthropology).
Northeast provides additional support that gallina- For the mentoring provided throughout this project, I
ceous birds played an important role in the prehistoric wish to express thanks to Robert Dewar, George
use of birds by Native Americans (Dirrigl, 1998). In my Clark, Robert Bee, and Nicholas Bellantoni. I would
database of 140 archaeological samples, wild turkey also like to thank Jane O’Donnell for drawing Figure
occurs in 38% of the samples. Less likely to be found 1. This paper benefited from the critical review and
are the remains of other gallinaceous birds (ruffed editing provided by Peter Stahl and an anonymous
grouse followed by bobwhite and prairie hen). Clearly, reviewer for the Journal of Archaeological Science;
the development of a regional characterization of however, I accept responsibility for any errors or
Native American bird use necessitates adopting a omissions.
taphonomic approach to examine the cultural and
non-cultural factors resulting in patterns of skeletal
remains. References
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