Journal of Archaeological Science (1998) 25, 973984

Article No. as980276

Gear Selectivity Models, Mortality Profiles and the

Interpretation of Archaeological Fish Remains: A Case Study
from the Harney Basin, Oregon
Ruth L. Greenspan
Heritage Research Associates, Inc., 1997 Garden Avenue, Eugene, Oregon 97403, U.S.A.
(Received 7 July 1997, revised manuscript accepted 14 January 1998)
Size selectivity of fishing gear and catastrophic mortality profiles have been used as models to test hypotheses regarding
(1) the natural or cultural origin of archaeological fish assemblages, and (2) the types of fishing gear that may have been
employed in harvesting fish. Although such models provide a useful framework within which to interpret archaeological
fish remains, it is important that they be applied with careful attention to fish population structure and behaviour, and
to archaeological and environmental factors. A case study from the Great Basin of North America is used to illustrate
both the strengths and weaknesses of gear selectivity models and catastrophic mortality profiles in the interpretation of
archaeological fish bone assemblages.
 1998 Academic Press
Keywords: TUI CHUB, FISH, MORTALITY PROFILES, GEAR SELECTIVITY, SIZE RECONSTRUCTION,
GREAT BASIN, HARNEY BASIN.

Introduction

conducted many studies on the selectivity characteristics of various types of fishing gear (e.g. Rollefsen,
1953; Hamley, 1975, and references cited therein;
Jensen, 1986, 1990; Millar, 1995). Such studies tend to
be highly technical and concerned with precision in
modelling gear selectivity.
Some archaeologists have used generalized and
greatly simplified models of gear selectivity to test
hypotheses regarding archaeological fish assemblages,
including whether an assemblage is cultural or natural
in origin, and what types of fishing gear may have
been used in harvesting the fish (e.g. Balme, 1983;
Noe-Nygaard, 1983; Wing & Scudder, 1983; Butler,
1996). The basic assumption underlying the use of size
distribution or gear selectivity models is that many
types of fishing gear do not result in the capture of a
random or representative sample of the available fish
at a given fishing location, but rather are selective for
particular types and size ranges of fish, and that this
selectivity is reflected in the population structure of an
archaeological assemblage of fish remains.
For example, gill nets are more selective with respect
to size than most other methods of fishing. Gill nets are
staked out vertically in the water, and catch fish by
entangling them by the gills when they try to swim
through the mesh. Fish that are sufficiently smaller
than the gauge of the mesh can swim through the net
without being caught, and those whose girth is sufficiently larger than the mesh can back away unharmed.
Thus a gill net with a relatively uniform gauge mesh
will capture fish in a fairly limited size range. A
generalized, hypothetical gill net selectivity curve

n many archaeological contexts, it is not uncommon for fish remains to be present, often in very
large quantities, but for no direct evidence of
fishing technology to be observed. In addition, many
fish bones in archaeological assemblages exhibit no
signs of cultural modification, such as burning or cut
marks. Thus, interpretation of fish remains is often
dependent upon such assemblage characteristics as
species composition, skeletal part representation and
the size range of the fish represented.
This paper focuses on the use of gear selectivity or
size distribution models to test hypotheses regarding
(1) whether an assemblage was accumulated by cultural or natural means, and (2) what type of fishing
gear may have been employed in harvesting the fish,
assuming the assemblage is a cultural one. I examine
the assumptions underlying the use of such models,
and point out a number of pitfalls to be avoided,
using an archaeological assemblage of tui chubs (Gila
bicolor) from the Harney Basin in the northwestern
Great Basin of North America as an example.

Gear Selectivity Models

Choice of fishing gear is dependent upon what kind of
fish is being targeted. Factors to be considered include
the size, feeding habits and behaviour of the fish being
sought, as well as conditions at a particular fishing site
(e.g. Lagler, 1978). Fisheries biologists and managers
concerned with stock assessment or gear efficiency have
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 1998 Academic Press

974

R. L. Greenspan

what species and size range of fish might be caught by

various types of gear.

Frequency

Hypothetical total population

Population Structure and Mortality Profiles

Seine net
Gill net

Baited hook

Fish size
Figure 1. Generalized gear selectivity curves for a gill net, hook and
line, and a seine net, compared to a hypothetical total population of
fish.

(Figure 1) is unimodal and roughly normal in distribution (Hamley, 1975: 1946). The particular size range
included under the curve depends upon the mesh size
of the net. Baited hooks may also have unimodal
selectivity curves (Figure 1), although they are less
strongly selective than gill nets, and thus typically have
a broader curve (Kenchington, 1993; Millar, 1995).
Unlike gill nets, the selectivity of bag-type nets such
as seine and dip nets is primarily a function of small
fish escaping through the mesh. Among the fish encountering the net, those small enough to escape
through the mesh are not caught, while larger fish
are caught. Thus a hypothetical selectivity curve for
seine nets shows a sharp cut-off on the left side of the
curve (Figure 1), while the remainder of the curve
should follow the size frequency distribution of the fish
encountering the net.
These hypothetical curves are also affected by aspects of fish ecology and behaviour. Fish encountering
the net may not be a cross-section of the total population of the lake, stream, or other fish habitat. For
example, the size distribution of fish encountering a
beach seine or a dip net operated from shore or in
shallow water might be very different from that of fish
encountering a seine or dip net operated in deeper
water from a boat. Net avoidance or escape on the part
of the fish are also relevant considerations. In addition,
it is important to take into account the fact that some
fish may have different seasonal distributions that
could affect gear selectivity models. In the case of
lacustrine tui chubs, adult fish tend to move offshore
into deeper water in the wintertime, and to move
inshore in the spring, while young fish remain in the
shallows most of the time. Thus in the summer months,
particularly during spawning activity, there may be
very dense aggregations of adult tui chubs in shallow,
nearshore water (e.g. Snyder, 1917), while during the
winter months, the adults are far more dispersed, and
many of them are not in nearshore waters at all
(Kimsey, 1954: 399; Vigg, 1980: 53). Such changes in
distribution need to be taken into account in predicting

The application of gear selectivity models to archaeological assemblages typically involves comparing the
size distribution of an assemblage not only to various
selectivity curves, but also to curves that represent
the hypothetical total population or a catastrophic
mortality profile. This is particularly important in
lacustrine settings, where natural mass mortality may
result in accumulations of fish carcasses along shorelines. Comparison of the population structure of the
archaeological assemblage with that expected for a
natural fish kill can aid in distinguishing between
natural and cultural accumulations of fish remains in
lakeside archaeological sites.
An example is a study by Noe-Nygaard (1983), in
which she analysed an assemblage of pike remains
from a Mesolithic lakeside site in Denmark. She compared the population structure of the archaeological
assemblage to specimens from an ecologically similar
modern lake that was poisoned with rotenone. The size
structure of the two assemblages differed markedly,
which contributed to Noe-Nygaards conclusion that
the archaeological assemblage exhibited signs of having been deliberately and selectively fished by humans.
A number of archaeologists and vertebrate palaeontologists have used similar models, typically age profiles derived from theoretical population dynamics, in
interpreting mammalian fossil assemblages. A catastrophic age profile, also known as a living structure
age profile, shows the structure or age distribution of a
living population. An attritional age profile, or true
mortality profile (Klein, 1982a: 58) shows the age
distribution of individuals that die each year of noncatastrophic, or attritional causes, such as accidents,
disease, predation, or old age (Figure 2(c)). Archaeologists have used such models of population structure
to help distinguish between cultural and non-cultural
accumulations of bones, or to determine the circumstances under which carcasses were obtained, such as
whether animals were scavenged or deliberately hunted
(e.g. Klein, 1982a, b; Stiner, 1990, 1995; Savelle &
McCartney, 1991).
Studies that have used theoretical catastrophic mortality models in interpreting fish remains include one
by Balme (1983) and one by Butler (1996). Balme
compared the reconstructed size range of golden perch
from sites along the Lower Darling River in Western
New Wales to selectivity curves for gill nets and traps,
and to a total population curve. Butler compared
the size distribution of an archaeological assemblage
of tui chubs from Stillwater Marsh, Nevada, to a
theoretical catastrophic mortality profile.
The catastrophic morality profiles used by both
Balme and Butler (Figure 2(a), (b)) are diagonal in

Interpretation of Archaeological Fish Remains

(a)

% Fish

Hypothetical total
population
Trap
Gill net

Fish length

Frequency

(b)

Fish length

Number of individuals

(c)

Increasing age

Increasing age

Figure 2. (a) Generalized selectivity curves for a drum net trap and a
gill net, compared to a hypothetical total population curve (redrawn
after Balme, 1983: 29). (b) Theoretical catastrophic mortality profile
used in Butlers study (redrawn after Butler, 1996: 704). (c) Theoretical catastrophic and attritional age profiles for mammalian population not characterized by multiple births (redrawn after Klein,
1982a: 152).

shape and imply a constant rate of mortality throughout life; neither Balme nor Butler provided any justification of why such a profile should be appropriate for
modelling fish population structure. Balme does not
discuss the theoretical derivation of her hypothetical
total population curve at all (Balme, 1983). Butler
(1996: 702703) cites other archaeological and palaeontological studies that have made use of theoretical
models of population structure in explaining the
derivation of the model she uses.
One of the studies Butler cites (Stiner, 1991) shows a
catastrophic mortality profile similar to Butlers (1996:
704) with a diagonal, or half-pyramid shape, implying
a constant rate of mortality. In fact, such a form
probably never occurs in natural vertebrae populations (MacArthur & Connell, 1966; 124). Both Butler
(1996) and Stiner (1990, 1991) acknowledge that the

975

corresponding attritional profile illustrates a high rate

of juvenile mortality, low mortality rates for prime age
adults and increased mortality for old individuals.
However, neither of them incorporates these variations
in mortality rate into the catastrophic or living structure profile. As pointed out by Klein, such a treatment
seems to assume that catastrophic and attritional age
structures are independent of one another, when in fact
they are related in essentially the same way as the fullness and emptiness of a container (Klein, 1995: 1844).
Although she does not explicitly say so, Butler
(1996) seems to assume that the shape of catastrophic
and attritional profiles would be the same for all
populations and that theoretical models of population
structure derived for large mammals would be equally
applicable to fish. However, Klein (1982a: 5254) has
demonstrated that age structure and mortality patterns
are not even the same for all large mammals, and that
an increase in reproduction rate will result in increased
juvenile mortality (assuming a stable population), and
corresponding changes in the shape of catastrophic
and attritional profiles. Many fish, including tui chubs
(Moyle, 1976: 167), may lay thousands of eggs per
spawning season; fish typically have a commensurately
high rate of juvenile mortality (Deevey, 1947: 301).
Given that the patterns of reproduction and mortality
differ widely between fish and mammals, the appropriateness of using a single model of population structure
for both of them is highly questionable.
An additional difficulty with Butlers model is that
she appears to assume that fish age profiles can be used
as a proxy for fish size profiles. The age of an organism
increases at a constant rate throughout its life. Increase
in size, on the other hand, is not constant, and
most vertebrates that live past adulthood reach their
maximum size before they reach their maximum age.
Despite this, Butler (1996: 704) compares the reconstructed length frequencies of her archaeological specimens to a catastrophic mortality profile, which is a
model based on age structure.
Many fish populations have a restricted spawning
season and relatively rapid growth during the first few
years. This combination of factors results in a distinctly
multimodal size profile (Sheldon, 1965; Craig & Oertel,
1996a, b). Like many other fish taxa, tui chubs grow
rapidly during their first few years, and then growth
tapers off (e.g. Kimsey, 1954: 405). In a tui chub size
and growth study conducted in Eagle Lake California,
Davis (1986) found that some individuals appeared to
reach their maximum length of roughly 340 mm by
approximately 8 years of age; some of these individuals
lived for more than 20 years. Although Davis sample
may not be representative of the entire population of
Eagle Lake, the small amount of size overlap between
the first four age classes (Figure 3) indicates that the
size structure of the entire population would still
exhibit a multimodal profile. Comparison between the
size distribution and the age distribution of Davis
Eagle Lake tui chub specimens (Figure 4) illustrates the

976

R. L. Greenspan
60
Age
10+
4
3
2
1
0

Frequency

45

30

15

100

200
Length (mm)

300

400

Figure 3. Size frequency distribution of tui chubs from Eagle Lake.

Each age class is represented by a different fill pattern (raw data
provided by Lawrence Davis).

60
(a)
Frequency

45
30
15
0
100

200
Length (mm)

300

400

60
(b)

Frequency

45
30
15
0
0

10

15
20
Age class

25

30

35

Figure 4. Histograms of size frequency (a) and age class frequency

(b) from the same sample of tui chubs from Eagle Lake (raw data
provided by Lawrence Davis).

point that fish size and age profiles may have very
different shapes and should not be substituted for one
another.
As with any taxon that exhibits a very high rate of
juvenile mortality, the age or size structure of a fish
population will vary depending upon what time of year
the census is taken. A population census taken shortly
after the fry hatch will show a much higher percentage
of the young of the year, or the 0 age class, than will a
census taken 6 months later. In addition, the likelihood

of the tiny, fragile bones of age class 0 fish being

recovered in an archaeological assemblage seems very
slim. Perhaps, therefore, models of fish size or age
structure that are to be useful for archaeological
studies should exclude age class 0 altogether. Admittedly, excluding age class 0, and only age class 0, is
both arbitrary and lacking in empirical support. Unfortunately, there is a lack of good population structure data for many fish taxa of archaeological interest
and lack of experimental data on element survivorship
for very young fish. Until these gaps in the data are
addressed, omitting age class 0 from fish population
structure models may increase the usefulness of such
models for archaeological interpretations.

The Harney Dune Site

The Harney Dune site (35HA718) is an extensive lithic
scatter situated on the lakeward side of Harney Dune
on the north shore of Harney Lake in south-central
Oregon (Raymond, 1994) (Figure 5). The Harney
Basin is the northwestern-most basin in the Great
Basin and, like the rest of that region, is characterized by internal drainage. The site is on land that
is administered by the Malheur National Wildlife
Refuge, U.S. Fish and Wildlife Service.
Cultural materials, including chipped and ground
stone tools, chipped stone debitage, fire-cracked rock,
stone net weights, bone barbs and faunal remains have
been exposed at the Harney Dune site in the beach
zone between the lake and the dune (Figure 5). Radiocarbon dates and temporally sensitive projectile points
place the occupation of the site within the last 500
years (Raymond, 1994).
In the summer of 1992 a systematic surface survey
was made of 764 1010 m units at the site as part of
a research programme for talented and gifted high
school students sponsored by Earthwatch, Inc. and the
Malheur National Wildlife Refuge. During the course
of this survey, a number of features were identified,
including concentrations of artefacts, debitage and/or
faunal remains. Several of the features contained concentrations of fish bones, which, in many instances,
were burned. Subsurface testing of some of these
features was accomplished during the summers of 1992
and 1994 (Raymond, 1994).
Fish remains recovered from three of the features
were selected for preliminary analysis. All of the fish
specimens that could be identified to species represent
tui chubs. Many of the specimens were burned, and an
inventory of skeletal elements indicates that all portions
of the fish were represented (Greenspan, 1993).
In addition to tui chub bones, a number of artefacts
were recovered that are interpreted as having been
associated with fishing activity. Flat, notched stones
identified as net weights (Figure 6) have been found at
a number of lowland archaeological sites in southeastern Oregon. At the Harney Dune site, several features

Interpretation of Archaeological Fish Remains

977

Basalt Uplands

126

5m

Gr

ea

1260
m

HA
RN

se

wo

od

Basalt Uplands

Fl

at

Mud
Lake

NE
U
D

Harney Lake
35HA718

Study Area
s

nd

a
pl

at

OREGON

ry

Fl

a
nt

e
as

od

im

re
G 1
265 m

ou

f
Tu

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ac

d
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2 km

Figure 5. Map showing the location of Harney Dune and the Harney Dune site on the northeast shore of Harney Lake; inset shows the location
of the study area in southeastern Oregon (after Greenspan & Raymond, 1996).

contained groups of notched stones found clustered

together (Figure 7); these features are interpreted to
represent locations at which a net was discarded or
abandoned and eventually decomposed, leaving only
the net weights behind. Both seine nets and gill nets
require the use of weights, and both were used by
various Northern Paiute groups who were inhabiting
the Northern Great Basin at the time of historic contact
(Fowler, 1989, 1992). In addition to net sinkers, small
bipointed bone artefacts that may have served as gorgetype fish hooks (Figure 8) have been found at the
Harney Dune site and at other sites in the Northern
Great Basin (Jenkins, 1994: 223, 227228, 232).
Although there is little ethnographic information on
fishing practices of the Wadatika Northern Paiute who
lived, and continue to live, in the Harney Basin, there
are detailed descriptions of fishing for other Northern
Paiute groups. According to Willard Parks informants
among the Northern Paiute groups of western Nevada,
the primary method of taking fish in Walker Lake was
by gill-netting (Fowler, 1989: 33). Other types of nets
reported to Park for use in river or lake settings
included dip nets, bag nets and seine nets (Fowler,
1989: 3035; 1992: 6162). The gauge of a net depended upon the type and size of fish being targeted.

Gauges made of wood were sometimes used when

making nets to achieve the desired mesh size; often,
fingers were used as gauges (Fowler, 1989: 87; 1992: 33,
116117). In the Lower Carson Basin, Northern Paiute
groups obtained tui chubs with gill nets, dip nets, open
twine basket trays, basket traps and hook and line
(Fowler, 1992: 6162).

Size Reconstruction of Tui Chub Specimens

The hypothesis
Common sense, experience and experimental data all
indicate that mass capture of fish that are aggregated
when spawning or schooling is more efficient than
individual capture techniques (e.g. Raymond & Sobel,
1990; Lindstrom, 1996). Lindstrom (1996) presents
diet breadth models of the prehistoric Truckee River
fishery. Her research indicates that small fish, including
tui chubs, are most efficiently exploited by mass capture with a basketry scoop, with gill nets and traps
being second. Larger fish, when aggregated, are most
optimally exploited by either gill nets or traps. The use
of bag, dip and lift nets rank next, and are followed
by various methods of obtaining non-spawning fish
(Lindstrom, 1996).

978

R. L. Greenspan

Although Lindstrom (1996) does not distinguish

between large tui chubs and small ones, but rather
considers all tui chubs in the category of small fish,

Figure 6. Notched stone net weights from the Harney Dune site.

there is reason to believe they should be considered

separately, both with respect to capture technique and
with respect to processing and consumption. Parks
ethnographic data indicate that the Cattail-Eater
Northern Paiute people at Stillwater Marsh fished for
large tui chubs with gill nets, traps and with hook and
line, while smaller tui chubs were most frequently
caught with dip nets and basket scoops (Fowler, 1992:
6162). A study of tui chubs in modern Eagle Lake
resulted in the capture of young of the year tui chubs
through the use of seine nets, while older (larger) tui
chubs were most effectively caught with gill nets
(Davis, 1986).

Figure 8. Two examples of composite fish hooks using single-pointed

barbs. The drawing on the left shows a greasewood trout hook
collected by S. A. Barrett in western Nevada (drawing by Kevin
McCornack from a photograph in Fowler (1989: 34)). The drawing
on the right is one of the 17 fish hooks on a trot line collected from
a cave on the east side of Pyramid Lake (drawing by Kevin
McCornack from a photograph in Tuohy (1990: 136)).

Figure 7. Feature 106 at the Harney Dune site consists of clustered net weights and is interpreted to be a location where a net was discarded
and eventually disintegrated, leaving only the stone weights.

Interpretation of Archaeological Fish Remains

There are both ethnographic and archaeological

data to suggest that small tui chubs were processed
differently from large ones. The Cattail-Eater Northern
Paiute dried small tui chubs whole by simply laying
them out in the sun along the shoreline or on an area of
cleared ground. When dried, the fish were placed in
woven tule bags and stored either by hanging them in
the shade or by burying them in pits lined with cattail
leaves. Small fresh fish were wrapped in cattail leaves
and buried in hot ashes to bake. The bones were
sufficiently softened by this process that there was no
need to remove them and they could be eaten along
with the meat. At least one Paiute informant stated a
preference for small fish because the bones did not need
to be removed (Fowler, 1992: 6263).
Large tui chubs were split lengthwise before being
placed on willow racks to be dried in the shade. This
process was undertaken only during cool weather in
order to avoid spoilage. Like small fish, larger fish were
also baked in ashes when fresh (Fowler, 1992: 63).
Although Fowler (1992) does not discuss whether or
not the bones of larger fish were consumed, presumably
they were not, given that small fish were preferred
because one need not worry about the bones (Fowler,
1992: 63). This assumption is supported by archaeological evidence from Hidden Cave, Nevada, suggesting
that the bones of small tui chubs were ingested, while
the bones of larger fish were not (Smith, 1985: 172178).
Lindstroms data suggest that if processing time is
not taken into account, small spawning fish, including
tui chubs, harvested with a basketry scoop rank top
among the 58 food types considered in her diet breadth
model (Lindstrom, 1996). When Lindstrom took processing time into account, assuming the same type of
processing for small fish as for large fish, mass capture
of spawning tui chubs fell in rank to number 20 out of
58. However, as discussed above, ethnographic and
archaeological data suggest that small tui chubs and
large tui chubs were not necessarily processed in a
similar fashion, and that processing time for small fish
was probably considerably less than for larger ones.
Specifically, small fish could be dried whole on the
ground, with no need to build drying racks or to split
the fish. Additionally, small fish could be eaten whole,
with no need to remove the bones or the viscera, which
would not only reduce processing time, but would
increase the nutritional value of the fish with respect
to vitamins, minerals and calories (e.g. Raymond &
Sobel, 1990; Lindstrom, 1996).
It is also worth noting that there is relatively little
time and effort invested in manufacturing a basketry
tray or scoop, which probably served other functions
in addition to fishing. In contrast, construction of a gill
net is a very labour intensive endeavour, and the
smaller the mesh, the greater the effort involved in
manufacture and repair (Lindstrom, 1996). Additionally, there is ample anecdotal evidence to suggest that
smaller tui chubs are easier to mass harvest than larger
ones. Thus, when equipment manufacture and repair

979

Figure 9. Feature 68, shown here after one-quarter of the pit was
excavated.

time are taken into consideration, plus the fact that

small tui chubs occur in high density aggregations in
near-shore settings more frequently than large ones, it
appears that small tui chubs may well have provided
higher return rates than larger ones, particularly outside of the spawning season.
In sum, then, the predictions of optimal foraging
theory, as well as ethnographic and archaeological
evidence, suggest that people did make a distinction
between small tui chubs and large ones, and that there
is a relationship between fish size and the manner in
which the fish were captured, processed and eaten.
Thus, an hypothesis was proposed that tui chub fishing
at the Harney Dune site was deliberately size-selective
and involved the use of gill nets. In order to test this
hypothesis, it was necessary to reconstruct the size
range of the tui chubs represented in the assemblage.
The reconstructed size distribution of the archaeological specimens could then be compared with hypothetical gear selectivity curves and lacustrine tui chub total
population curves, and evaluated in the context of
other archaeological data from the site. Three features
were selected for this analysis: Feature 68, Feature 89,
and Feature 66.
Feature descriptions
Feature 68 is a shallow bowl-shaped pit 160 cm in
diameter that occurs as a remnant in a scoured and
eroded blowout (Figure 9). Excavation of one-half of
the feature revealed the presence of fire-cracked rocks,
charcoal, burned and carbonized sticks identified as
chenopod, small amounts of chenopod seeds, mammal
bones and debitage and thousands of tui chub bones.
Most of the tui chub bones were burned and many
of them were completely carbonized. Two charcoal
samples returned calibrated calendar ages of
151060 (Beta 65166) and 161070 (Beta 64974)
(Greenspan & Raymond, 1996).
Feature 89 is an irregularly shaped erosional
remnant of midden deposit covering approximately

980

R. L. Greenspan

15 m2. A test unit was excavated into the midden and

resulted in the recovery of fire-cracked rock, debitage,
two Desert series projectile points, one mano and
thousands of fish bones. Two charcoal samples returned calibrated calendar ages of 1850100 (Beta
75246) and 189060 (Beta 75247) (Greenspan &
Raymond, 1996).
Like Feature 89, Feature 66 is an erosional remnant
of midden deposit, covering an irregularly shaped area
approximately 8 m2. A test unit was excavated into an
eroding edge of the feature, resulting in the recovery of
fire-cracked rock, debitage, blade flakes and a blade
core, four Desert series projectile points, three bone
gorges and large quantities of fish bones. Three charcoal samples were submitted for radiocarbon dating;
two returned modern dates (Beta 65164 and Beta
75244) and the third returned a calibrated date of
179070 (Beta 75245) (Greenspan & Raymond,
1996).
The fill from the features was removed from the field
in bulk samples, and was later screened and sorted by
students in the laboratory. Bulk samples from Feature
66 and Feature 68 were screened through nested sieves
of 3 mm and 1 mm mesh, and the analytical sample
selected for this study included specimens recovered in
both the 3 mm and the 1 mm fractions. Feature 89 was
screened only through 1 mm mesh.
Methodology
In order to estimate fish length, measurements were
made on bones from two modern comparative collections of tui chubs with a total of 44 specimens.
Regression equations were calculated based on the
modern reference collections; 14 different measurements were taken on a total of five different skeletal
elements. Measurements on the hyomandibular and
the basioccipital were selected for the analysis because
those particular portions of those two elements (Figure
10) were the most abundant and best preserved in the
archaeological assemblage, and the measurements
are highly correlated with fish length. The regression
equations and other relevant statistics are presented
in Table 1. The archaeological specimens were then
measured and fish length was predicted from the
regression equations.
Results
Estimation of the size ranges of the tui chubs recovered
from Features 66, 68 and 89 indicates that all of the
specimens included in the analysis were adult fish. The
estimated total lengths ranged from 130342 mm. In
comparison, the tui chubs in Davis Eagle Lake study
reached 130 mm some time between their second and
third summers, with an average maximum size for
adult fish of 314 mm. The largest tui chub measured by
Davis was 374 mm (Davis, 1986).
There are distinct differences between the three features at the Harney Dune site with respect to size range

(a)

(b)

Figure 10. Tui chub basioccipital (a) and hyomandibular (b), with
measuring points indicated.
Table 1. Output from regression of logN bone length and logN fish
length (TL)

Regression formula
R squared
Standard error of y estimate
Exponent of standard error
of y estimate

Hyomandibular

Basioccipital

1047 (x)+3140
0985
0045

1113 (x)+3911
0964
0069

111

117

Table 2. Estimated size (TL) in mm of fish remains from Harney Dune

site features
Feature

Minimum

Maximum

Mean

SD

68
89
66

86
93
17

130
161
240

228
321
342

184
227
277

207
240
267

(Table 2; Figure 11). The specimens from Feature 68

are the smallest, while those from Feature 66 are the
largest. The specimens from Feature 89 are intermediate in size between those from Features 68 and 66.
Table 2 shows the size range, mean and standard
deviation of the reconstructed fish lengths from each
feature. Histograms of the estimated lengths of the fish
from each feature are shown in Figure 11. Features 68
and 89 both have unimodal, roughly normal distributions, as indicted by the normal probability plots in
Figure 12. Feature 66, in contrast, does not show any
particularly strong patterning, which might be a function of the relatively small number of specimens
(N=17) available from that feature.

Cultural or Natural Deposits?

None of the three features has the population structure
of a living lacustrine tui chub population, which is
multimodal and contains many small individuals (e.g.
Figure 3). However, the population structure alone is
not enough to rule out a natural fish kill. Certainly

Interpretation of Archaeological Fish Remains

981

22
18

Frequency

N = 86

(a)

13
2
9
1
4
0
0
120

140

160
180
Total length (mm)

200

220
1

30
2

Frequency

25
20

3
100

N = 93

15

150

200

250

200

300

400

300

350

3
(b)

10
2
5
0
120

1
160

200
240
280
Total length (mm)

320

360
0

5
1

Frequency

4
2

N = 17
3

3
100
1
2
(c)
0
220

240

260

320
280
300
Total length (mm)

340

360
1

Figure 11. Size frequency distributions of reconstructed length (TL)

of tui chubs recovered from (a) Feature 68, (b) Feature 89 and (c)
Feature 66.

some fish die-offs kill all the fish in a given area, or

non-selectively kill a sample of them. But it is also
possible for a lethal agent to act selectively on certain
age or size classes. For example, very young fish may
be more susceptible to changes in water temperature
or chemistry; similarly, they may be more vulnerable
to infectious disease than mature individuals. On the
other hand, adult fish that have recently spawned may
be highly vulnerable to disease or stressful conditions
due to their depleted energy stores (Logan & Markle,
1993: 237). Thus, a total population curve is not

2
200

250
Length

Figure 12. Normal probability plots of the size frequencies of tui

chubs from (a) Feature 68, (b) Feature 89 and (c) Feature 66.

982

R. L. Greenspan

necessarily an appropriate proxy for a catastrophic

death curve in the case of natural fish kills.
Furthermore, it is necessary to consider the taphonomy of natural fish kills before relying solely on
population structure as an indicator of cultural selectivity in an assemblage. Numerous researchers have
noted that small and immature specimens tend to be
significantly underrepresented in vertebrate fossil assemblages due to inherently poor preservation in very
young individuals (e.g. Caughley, 1966: 909; Klein,
1982a: 56). In the case of fish kills, an additional factor
to be considered is selective scavenging of the carcasses, particularly by birds (Logan & Markle, 1993:
237; A. Munhall, pers. comm.). Specifically, birds that
are feeding on a fish kill may carry away small fish,
while larger fish might be passed over or dismembered
at the site of the fish die-off. This would result in an
increased likelihood that remains of the larger fish
would be incorporated into archaeological deposits,
while the smaller ones might go largely unrepresented.
Thus, although the population structure of the fish
remains from the Harney Dune site shows a pattern
consistent with deliberate, size-selective fishing, there
are a number of non-cultural factors that could result
in similar patterns. It is therefore particularly important to rely on as many different lines of evidence as
possible in interpreting the archaeological fish assemblage. At the Harney Dune site there other lines of
evidence that strongly indicate that the fish remains
are, in fact, of cultural origin. First, the fish bones are
clearly associated with the cultural deposit. In contrast,
no fish bones were reported from a trench cut into an
off-site portion of Harney Lake Dune as part of a
geomorphic study (Dugas, 1992: 4147). Secondly, a
large population of the assemblage is burned, and the
sites setting along the shore of an alkaline lake, in an
area where there is very little fuel to sustain a natural
brush fire, strongly suggests the burning is cultural.
And finally, the presence of artefacts that are generally interpreted to represent fishing gear is additional
evidence of deliberate fishing at the Harney Dune site.

Evidence of Fishing Technology

Assuming, then, that the tui chub bones were culturally
deposited, does the size distribution of the fish tell us
anything about how the fish were caught? The size
distribution curves from Features 68 and 89 (Figure
11) are very much like what would be expected from
gill-netted populations. However, depending upon the
time of year and the location of the fishing effort, fish
caught by other means could exhibit curves of similar
shape. Tui chubs are schooling fish, and frequently
form schools that are segregated by size (Douglas
Markle, pers. comm.). Thus fishing methods that target
an individual school of fish, such as dip nets or
basketry scoops, may be sampling a sub-population
that is fairly restricted in size. In such a case, the
overall shape of the distribution curve could be similar

to that observed for gill nets. Although the shape of the

distribution curve may be the same, the limited ethnographic evidence available suggests that the size range
of fish obtained by basketry scoops and gill nets would
be quite different (Fowler, 1992: 6162). This is supported by Lindstroms (1996) experimental data and by
anecdotal reports from fisheries biologists.
In the case of the Harney Dune site, the presence
of notched stone net weight features suggest that
weighted nets were used, although that certainly does
not rule out the use of other methods of catching fish
as well. The population size structure of the fish
remains from Features 68 and 89 is more consistent
with the use of gill nets than bag or seine nets to catch
tui chubs. Theoretically, a seined population should
have a more abrupt cut-off on the small end of the
graph (Figure 1). Fish too large to escape through the
mesh should be retained in the net regardless of how
much larger they are than the minimum size caught.
In practice, however, this may not always be the
case. Davis reported that in Eagle Lake he caught tui
chubs with gill nets in June and early July, but efforts
to catch them with beach seines during the same time
period were completely unsuccessful. In mid-July large
numbers of young of the year were obtained by seining;
no larger fish were caught in the seine nets, although
they were caught in gill nets during the same time
period (Davis, 1986). Thus, it may be that beach seines
are effective for catching small tui chubs, but possibly
less so for catching large ones, such as those represented in the Harney Dune site features. Again, this is
consistent with the ethnographic fishing practices of
the Cattail-Eater Northern Paiute, who used gill nets,
traps and hook and line for obtaining large tui chubs,
and used basketry scoops and dip nets (which were
used in the fashion of a seine) for catching smaller
tui chubs (Fowler, 1992: 6163).
Feature 66 differs from the other two features in
several respects. The sample size is small (N=17),
which may reflect a smaller catch, or it may reflect a
recovery or sampling problem. The size distribution is
not normal, or even obviously unimodal (Figures 11,
12). Also, the fish represented in Feature 66 are larger
than those from the other two features, overlapping
with the larger specimens from Feature 89, but showing virtually no overlap with Feature 68 (Figure 13).
Three bipointed bone gorges, believed to have been
used as composite fishhooks, were recovered from the
Harney Dune site. All three were found in Feature 66,
suggesting the possibility that these fish were caught
with baited hooks.

Discussion
The hypothesis that deliberate and size-selective fishing
for tui chubs occurred at the Harney Dune site is
supported by the present study. The fish recovered
from the Harney Dune site are relatively large for tui
chubs. The reconstructed total lengths range from 130

Interpretation of Archaeological Fish Remains

40

Frequency

30

Feature 68
Feature 89

20

Feature 66

10

0
120

160

200
240
280
Total length (mm)

320

360

Figure 13. Comparison of size frequencies of tui chubs from all three
features.

to 342 mm total length (TL), with a mean of 212 mm;

more than half the specimens represent individuals
larger than 210 mm.
In comparison, Butlers specimens from site
26CH1062 at Stillwater Marsh range from 52 to
143 mm standard length (SL), with a mean of 89 mm
(Butler, 1996: 704). Raymond & Sobel (1990: 8)
measured samples of dried chub found in prehistoric
cache pits from several cave sites in Nevada. Specimens
from Stick Cave ranged from 75 to 120 mm fork length
(FL); those from Humboldt Cave ranged from 80 to
220 mm (FL), and those from Lovelock Cave ranged
from 43 to 130 mm (FL).
It should be pointed out here that the sizes of the fish
in these assemblages are not all directly comparable. I
used total length (TL) in the Harney Dune site study,
which is the greatest length that can be measured and is
defined as the distance from the tip of the snout to the
end of the caudal fin (tail). Butler used standard length
(SL), which is the distance from the tip of the snout to
the end of the vertebral column. Raymond & Sobel
used fork length (FL), which is the distance from the
tip of the snout to the deepest point in the fork of
the caudal fin. This means that the differences in size
between the Harney Dune site specimens and the specimens from the various Nevada assemblages is not as
great as the raw numbers would suggest. Nevertheless,
even taking the different measurements into account,
the Harney Dune site specimens are relatively large.
Taken together, the cultural context and burned
condition of the bones, the quantities of fish remains in
some features, the relatively large size and narrow size
range of the fish from each feature and the presence of
net weights and bone gorges are fairly compelling
evidence that tui chubs were an important, deliberately
fished resource for the inhabitants of the Harney Dune
site. The unimodal, approximately normal size distributions of the reconstructed fish lengths from Features
68 and 89 suggest deliberate fishing with size selective
gear. This pattern is consistent with gill net fishing. The

983

distributions from Features 68 and 89 have different

ranges and statistically different means (t=
1278,
P<0001), which suggest that the fish were caught with
two different nets. This inference is further supported by
the radiocarbon dates, which indicate the features may
differ in age by up to a few hundred years. The data
from Feature 66 are inconclusive, although the large
size of the fish, the relatively small number of individuals represented and the presence in the feature of bone
gorges may be indicative of hook and line fishing.
Whether an individual feature represents a single
fishing event or multiple events remains unknown. It
might be possible to determine season of death for
some of the specimens through the analysis of incremental growth rings; multiple events occurring during
different times of the year might be detectable through
such analysis. Multiple events occurring within a single
season, or during the same season in different years,
are not likely to be recognized in the archaeological
record. With some fishing gear, including scoops and
seine and dip nets, one could get different size profiles
with the same gear, depending upon fishing location
and time of year. Such an effect would be less apparent
with gill nets, because of their greater size selectivity.
The use of gear selectivity models and mortality
profiles has aided in testing the hypothesis that the tui
chubs represented at the Harney Dune site were caught
with gill nets. The successful application of such models
is dependent upon a careful consideration of the behaviour and population structure of the species under consideration, including schooling behaviour and seasonal
movements. It is also important to consider the population structure, potential selectivity and taphonomic
factors affecting natural fish kills. However, the usefulness of such models is limited unless the assemblages
are evaluated in light of their archaeological and environmental contexts. When these factors are all taken
into account, gear selectivity models and mortality
profiles can provide a useful framework within which to
evaluate and interpret archaeological fish remains.

Acknowledgements
This research was supported by the U.S. Fish and
Wildlife Service. I thank A. W. Raymond for inviting
me to participate in the Harney Dune site project, and
for his input and support during every phase of the
research. This paper has benefitted from discussions
with K. A. Cruz-Uribe, D. Markle, A. Munhall
and G. R. Smith. Assistance with various aspects of
this project was provided by P. J. Bartlein, L. Davis,
J. Ricks, W. L. Minckley, D. W. Steinman and
Heritage Research Associates, Inc. L. Davis generously
allowed me to include his unpublished data on Eagle
Lake tui chubs. Figures 8 and 10 were drafted by
K. McCornack. K. A. Cruz-Uribe helped prepare
Figures 1, 2, 5 and 13. The photographs for Figures 6,
7 and 9 were provided by A. W. Raymond. Drafts of

984

R. L. Greenspan

this paper were read and commented on by K. A.

Cruz-Uribe, R. Minor, A. W. Raymond, and two
anonymous reviewers. I take full responsibility for any
errors or omissions.

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