Taphonomic Investigations of Owl Pellets

Author(s): Peter Dodson and Diane Wexlar

Source: Paleobiology , Summer, 1979, Vol. 5, No. 3 (Summer, 1979), pp. 275-284
Published by: Cambridge University Press

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Paleobiology, 5(3), 1979, pp. 275-284

Taphonomic investigations of owl pellets

Peter Dodson and Diane Wexlar

Abstract.-Owls are important consumers of small vertebrates, and because they regurgitate pellets rich
in bone, they may be important potential contributors of the concentrated remains of small vertebrates
to the fossil record. Owls of three sizes, the large great horned owl (Bubo virginianus), the medium-sized
barn owl (Tyto alba), and the small screech owl (Otus asio), were fed a common diet of mice. The bony
contents of the pellets were analyzed to determine the amount of bone loss by digestion, bone complete-
ness, and sites of bone breakage. For all three species, only about half the number of bones ingested
were recovered in the pellets. Mandibles and femora were most abundant, and pelves and scapulae were
the least abundant. Screech owls broke 80% of the cranial and limb elements, barn owls only 30%.
Skulls fared poorly in great horned and screech owl pellets, while barn owls returned 80% of the skulls
intact, with only the caudal portion of the cranium damaged; barn owls also returned articulated strings
of vertebrae and complete paws. These results provide a baseline for the recognition of owls as agents
of accumulation of small bones in the fossil record and suggest that the actions of ancient predators may
be revealed by species-specific patterns of bone destruction of an assemblage of fossil prey species.

Peter Dodson. Laboratories of Anatomy, School of Veterinary Medicine, University of Pennsylvania,

Philadelphia, PA 19104, and Research Associate, Academy of Natural Sciences, 19th and Parkway,
Philadelphia, PA 19103

Diane Wexlar. Laboratories of Anatomy, School of Veterinary Medicine, University of Pennsylvania,

PA 19104

Accepted: May 9, 1979

Owls in Taphonomy conceptions or unwise generalizations persist;

for instance, owls are alleged to ingest prey
A fundamental and widely appreciated fact
items whole, to digest no bone, and to regur-
of avian biology is the observation that owls
gitate bone largely undamaged (Mayhew 1977).
regurgitate the bones of their prey. Pellets of a
All of these views are challenged in this paper.
group of owls averaged 46% bone, whereas a
The purpose of this study is to examine from a
group of hawks yielded only 6.5% bone by
paleontological perspective the pellets produced
weight (Duke et al. 1975). The striking differ-
by several species of live owls with view to pos-
ence between hawks and owls evidently stems
ing three questions: 1. Are there characteristics
from the difference in levels of stomach acidity,
by which a suite of small bones can be confi-
the basal pH of hawks averaging 1.6, that of
dently recognized as representing an owl pellet?
owls 2.35. Owl pellets have been used by mam-
2. Do different species of predator impose on
malogists to survey the extant small mammal
the remains of the prey a characteristic taphon-
fauna of a region (e.g. Long and Kerfoot 1963;
omic signature by which the identity of the
Dexter 1978). The paleoecological potential of
predator may be revealed? and 3. What role
owl pellets has been commented upon (Davis
does size of the predator play in the character
1959; Mellett 1974), and the action of owls has
of the suite of bones produced?
been inferred by paleontologists for specific de-
posits (e.g. Guilday et al. 1977; Gnidovec 1978).
Procedure
However, despite the use of owl pellets by bi-
ologists and the recognition of the potential sig- To study the effects of differential predation,
nificance of owl pellets by paleontologists, there captive owls of three different sizes were fed a
has not yet been a systematic study of the os- common diet of mice: great horned (Bubo vir-
teological characteristics of the bones of small ginianus), average male weight 1300 g (Craig-
animals that have served as prey for owls. Mis- head and Craighead 1969); barn (Tyto alba),

? 1979 The Paleontological Society. All rights reserved. 0094-8373/79/0503-0005/$ 1.00

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276 PETER DODSON & DIANE WEXLAR

TABLE 1. Proportional representation of skeletal elements. The number of specimens (n) is the minimum number of
specimens based on major elements (mandible and femur for great horned owl; skull or barn owl; and femur for screech
owl). Percentage present (%) expresses the proportion of specimens present relative to the potential number (i.e. n x 2 for
limbs, n x 18 for unguals, n x 54 for vertebrae). Percentage complete is the proportion of elements complete relative to
the total number actually present.

Great Horned Owl n = 37 Barn Owl n = 17 Screech Owl n = 12

Com-
Present % Complete % Present % Complete % Present % plete %

Skull 35 94.6% 1 2.9% 17 100% 14 82.4% 8 66.7% 1 12.5%

Mandible 73 98.7% 28 38.4% 32 94.1% 29 90.6% 16 66.7% 5 31.3%
Scapula 45 60.8% 1 2.2% 30 88.2% 12 40.0% 8 33.3% 0 0.0%
Humerus 61 82.4% 38 62.3% 28 82.4% 25 89.3% 16 66.7% 2 12.5%
Radius 51 68.9% 28 54.9% 29 85.3% 24 82.8% 18 75.0% 6 33.3%
Ulna 59 79.7% 29 49.1% 27 79.4% 17 63.0% 20 83.3% 5 25.0%
Pelvis 60 81.0% 2 3.3% 26 76.5% 8 30.8% 19 79.2% 0 0.0%
Femur 73 98.7% 51 69.9% 24 70.6% 24 100.0% 23 95.8% 8 34.8%
Tibia 78 105.4% 4 5.1% 23 67.7% 17 73.9% 19 79.2% 2 10.5%
Total 536 85.2% 182 34.0% 236 81.7% 170 72.0% 147 72.1% 29 19.7%

The following small elements, mostly intact, were recorded only as present
Clavicle 40 54.1% 19 55.9% 4 16.7%
Astragulus 63 85.1% 25 73.5% 19 79.2%
Calcaneum 60 81.1% 23 67.7% 16 66.7%
Unguals 64 9.6% 88 22.1% 61 28.2%
Vertebrae 1,048 52.5% 622 67.8% 283 43.7%
Grand Total 1,811 51.5% 1,013 62.7% 530 46.5%

average male weight 285 g; and screech (Otus
asio), average male weight 167 g. Pellets used
in the study came from two different groups,
one from the Academy of Natural Sciences of
Philadelphia (ANSP), and the other from the
Philadelphia Zoo. Thus the findings reported
are not the idiosyncrasies of single birds, but for
each species a minimum of three individuals
and two institutions are involved. Ten pellets
were analyzed for each of the two larger species,
and 20 were used for the screech owl.
In order to obtain complete extraction of
bone, the pellets were prepared as follows. Each
pellet was soaked in shallow water for an hour
to initiate mechanical breakdown. As many
bones as possible were carefully removed from
the softened pellet. Commercial hair remover
(Nair, Carter Products, New York; active in-
gredients sodium and calcium thioglycolate)
was added (approximately 20 cc per pellet) and
allowed to stand for 30 min. Then the pellet
was transferred to a beaker and diluted with
tweezers under a magnifying lens. The yield of
bone is similar whether mechanical processing
alone or the chemical method is used, but the
latter is both much quicker and permits a higher
level of confidence that complete extraction of
bone has been accomplished. Using the chemi-
cal method, articulated strings of caudal verte-
brae and complete paws have been removed, so
the use of the hair remover does not appear to
have an adverse effect on either ligaments or
bones in these conditions.
For each pellet, bones were sorted by ana-
tomical element and then subdivided into cat-
egories representing intact condition and var-
ious characteristic non-intact states. Ribs,
metapodials and cranial fragments, the latter
particularly abundant, were not tabulated ow-
ing to their small size and fragile nature. Results
were pooled for all pellets of each species (Table
1).
Observations

200 cc of water. The addition of 5 cc of ethanol
and 5 cc of ether helps breakdown the emolients
in the hair remover. The residue was then de-
canted into a vacuum filtration unit, changing
filter paper frequently to avoid clogging. Bones
were easily picked off the filter paper with
At the Academy of Natural Sciences three
great horned owls over an 8 day interval con-
sumed an average of 5.6 mice per day per bird;
during the same interval two barn owls ate 3.0
mice per day while two screech owls ate but 1.8
mice per day per bird. It is reported that the

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 TAPHONOMY OF OWL PELLETS 2 7 7

BO .
1.0 *
GH.

0 .4-

--50 \ . 7/ \**
.4 SO \ / \ :1 \.

.4 I

.2

Sk M S H R U P F T Ci A C Un V
FIGURE 1. Proportional representation of bones in
number potentially present based on minimum
representation of mandibles and femora, and po
element; ordinate: proportion preserved. Abbreviations; SK: skull; M: mandible; S: scapula; H: humerus; R: radius; U:
ulna; P: pelvis; F: femur; T: tibia-fibula; Cl: clavicle; A: astragulus; C: calcaneum; Un: unguals; V: vertebrae. BO: barn
owl-solid line (-); GH: great horned owl-dotted line ( ); SO: screech owl-dashed line (--).

mean and modal values of pellets cast per meal
are close to 1.0 for great horned owls (Duke et
al. 1975), but observations at the Academy of
Natural Sciences indicate that certain individ-
uals are more likely than others to cast either
two pellets or none. ANSP barn owls cast an
average of 1.5 pellets per meal, while the
screech owls cast fewer than one pellet per meal
despite daily feeding (one screech owl cast a sin-
gle pellet on three days, none on three other
days; the other one cast three pellets once, two
pellets once, one pellet twice and no pellets
twice).
Table 1 summarizes the contents of the pel-
lets. Ten great horned owl pellets contained an
average of 201 identifiable bones or bone frag-
ments (exclusive of ribs and cranial fragments
as indicated above) and together represented a
minimum of 37 mice (based on mandibular and
femoral count). Barn owl pellets averaged 107
identifiable bone fragments each, with 10 pellets
representing a minimum of 17 mice (based on
mandibles). Screech owl pellets averaged 59
identifiable bone fragments and represented a
minimum of 12 mice (based on femora). Several
screech owl pellets were barren or essentially
so, consisting only of compacted hair.
A mouse can be expected to yield 54 verte-
brae, 18 unguals, one skull, and a pair each of
mandibles, limb elements and limb girdles. For
all three species, the actual yield for all elements
combined was about half the potential yield,
though slightly greater for the barn owl (62.7%)
than for the other species (Table 1). Mandibles
and femora were the best represented elements
(Figure 1), though the former were not well rep-
resented for screech owls and the latter for barn
owls. The fragile scapula is under-represented
in the pellets of great horned and screech owls
but not in the barn owl pellets.
The three species differ greatly in the amount
of mechanical damage to the bones of their prey
(Table 2; Figure 2). For barn owls, 72.0% of
the bones were recovered intact (maximum
100% for femur; minimum 30.8% for pelvis);
while for the screech owl a mere 19.7% of bone
escaped breakage (maximum 34.8% for femur;
minimum 0% for scapula and pelvis). Despite
the great differences among species in absolute
amount of damage, there is remarkable similar-
ity in the pattern of relative susceptibility of
various skeletal elements to damage. For all
three owls the scapula and pelvis are most sus-
ceptible to damage, and the femur (and man-
dible and humerus to a lesser extent) is the least
susceptible. Only the humerus of the screech

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278 PETER DODSON & DIANE WEXLAR

1.0

BO

.8

.60

2
so

GH/

Sk M S H R U P F T
FIGURE 2. Occurrence of entire bones in owl pellets. Proportion of intact, undamaged specimens of each element relative
to the number actually represented-data from Table 1. Note the very much higher proportion of undamaged bone for
barn owls than for the other two species. Note also the relatively good preservation of mandible, humerus and femur,
poor preservation of scapula and pelvis. Abbreviations as in Figure 1.

owl pellets fails to conform to the pattern of
similarity of all three species, being consider-
ably more prone to breakage, both relatively
and absolutely, than for the other owls.
Characteristic sites of bone damage can be
identified in the mouse bones of owl pellets
(Table 2). The skull is a site of intense destruc-
tive activity. No skulls passed undamaged (Fig-
ure 3). All skulls tabulated as "complete" in
Table 1 actually lacked the back half of the cra-
nium (parietals, temporals and occipital). Only
a single skull each appeared in the great horned
and screech owl samples of ten pellets each. The
skull in the great horned owl pellet lacked both
nasals as well as right premaxilla and maxilla;
that in the screech owl sample lacked even fron-
tals, consisting only of paired premaxillae and
maxillae. Maxillae, premaxillae and petrosals
are abundant, well preserved skull elements;
other skull parts are highly fragmented. Quite
in contrast are the skulls from the barn owl pel-
lets; each pellet contained at least one skull, two
contained two skulls and one contained three.
One skull had the mandibles in articulation,
with both temporals and the left parietal pres-
ent; only the occipital and right parietal were
missing. At least one nasal was present in 7 of
14 skulls, and only one skull lacked a premax-
illa. In addition, parietals, temporals and occip-
itals, highly fragmented in the other owls, are
often preserved separate but intact.
In great horned owls and screech owls there
is a tendency for either the front end or the back
end of the mandible to break off, or infrequently
both (Figure 3). In great horned owls, loss of
the back of the mandible is far more frequent
(Table 2).
The scapula is a delicate bone very suscepti-
ble to breakage (Figure 4). Very common is ero-
sion of the dorsal border to a greater or lesser
extent, imparting a "feathery" appearance.
Often only the robust ventral (articular) portion
remains; broken scapular spines are among the
identifiable small parts recovered. The robust
humerus is twice as likely to be injured at the
proximal end than at the distal end or at both

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 TAPHONOMY OF OWL PELLETS 279

TABLE 2. Sites of bone breakage. N refers to the numb

The numbers may differ slightly from n in Table 1, in
numbers. "-" designates missing; e.g. "mandible-front" means "front end of mandible missing"; "Femur-prox" means
"proximal end of femur missing." il = ilium; is = ischium; pu = pubis; fib = fibula.

Great Horned Owl Barn Owl Screech Owl

N % N % N %

Mandible Intact 28 38.4% 29 90.6% 5 27.8%

-front 2 2.7% 1 3.1% 5 27.8%
-back 39 53.4% 2 6.3% 6 33.3%
front, back 4 5.5% 0 0 2 11.1%
Total 73 32 18

Scapula Intact 1 2.2% 12 40.0% 0 0

feathery 19 42.2% 10 33.3% 2 25.0%
distal 25 55.6% 8 26.7% 6 75.0%
Total 45 30 8

Humerus Intact 38 62.3% 25 89.3% 2 12.5%

-prox 11 18.0% 2 7.1% 6 37.5%
distal 5 8.2% 1 3.6% 4 25.0%
shaft only 7 11.5% 0 0 4 25.0%
Total 61 28 16

Radius Intact 28 63.6% 24 82.8% 6 33.3%

-prox 2 4.5% 1 3.4% 0 0
distal 13 29.5% 4 13.8% 9 50.0%
shaft only 1 2.3% 0 0 3 16.7%
Total 44 29 18

Ulna Intact 29 51.8% 17 65.4% 5 25.0%

-prox 5 8.9% 1 3.8% 1 5.0%
-distal 16 28.6% 8 30.8% 8 40.0%
shaft only 6 10.7% 0 0 6 30.0%
Total 56 26 20

Pelvis Intact 2 6.5% 8 47.1% 0 0

-tip 4 12.9% 5 29.4% 1 12.5%
back 10 32.3% 2 11.8% 0 0
-tip, back 9 29.0% 0 0 6 75.0%
i1, is 3 9.7% 0 0 0 0
il, pu 2 6.5% 0 0 0 0
is, pu 1 3.2% 2 11.8% 1 12.5%
Total 31 17 8

Femur Intact 51 71.8% 24 100% 8 47.0%

-prox 6 8.5% 0 0 2 11.8%
distal 13 18.3% 0 0 5 29.4%
shaft only 1 1.4% 0 0 2 11.8%
Total 71 24 17

Tiba-Fibula IntacL 4 5. 1% 17 73.9% 2 10.5%

-fib 24 30.8% 0 0 1 5.3%
-prox 4 5.1% 1 4.4% 1 5.3%
distal 1 1.3% 4 17.4% 2 10.5%
prox, fib 18 23.1% 0 0 2 10.5%
distal, fib 14 18.0% 1 4.4% 3 15.8%
shaft only 13 16.7% 0 0 8 42.1%
Total 78 23 19

ends (Table 2; Figure 4). Surprisingly, the del- ulna, injury to the distal end is very much more
icate radius is little more susceptible to injury common than to the proximal end (Figure 4).
than is the humerus and is consistently less bro- The pelvis is another site of heavy destruc-
ken than the ulna. For both the radius and the tion, and 50% or more of pelves are reduced to

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280 PETER DODSON & DIANE WEXLAR
FIGU
(occi
artic
plus
and
to th
angl
skull fragments: premaxilla, petrosal, parietal.
separate bones (ilium, ischium, pubis or pieces
thereof) (Figure 5). The most common sites of
injury to intact pelves are the cranial tip of the
ilium and the caudal border of the ischium (Fig-
ure 5). The robust femur sustains breakage at
its distal end (Figure 5). The tibia-fibula is a
compound bone, the delicate fibula being free
proximally and fused distally (Figure 5). The
fibula is likely to be present in less than 20% of
cases, except in barn owl pellets. Though for
great horned owl pellets loss of the proximal
tibia is somewhat more common than loss of the
distal tibia, the reverse is true for barn owl pel-
lets; for screech owl pellets the tendency is for
loss of both ends (Table 2).
Discussion
Owls are both geographically widespread and
of considerable antiquity; Bubo dates from the
Lower Oligocene, and Tyto and Otus appeared
in the Miocene (Brodkorb 1971). Representa-
tives of each of the three genera used in this
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study were found worldwide. Tyto alba, the
barn owl, is cosmopolitan (Presst and Wagstaffe
1973); Bubo virginianus, the great horned owl,
ranges through North and South America (Fog-
den 1973), while Otus asio, the eastern screech
owl, is confined to North America east of the
Rocky Mountains (Hekstra 1973). The potential
of these and other owls for contribution of the
remains of small vertebrates to the fossil record
may thus be considerable.
Accumulations of small vertebrates have
been important sources of information to pale-
ontologists (e.g. Estes 1964; Estes and Berber-
ian 1970; Wolff 1973), and their modes of for-
mation have been the subject of several studies
(e.g. Dodson 1973; Mellett 1974; Korth in
press), though the problem remains unresolved.
This study is an attempt to establish criteria by
which owl-related accumulations of small bones
can be recognized in the fossil record. Different
species of predators appear to impose distinctive
characteristics upon the remains of their prey
such that it may be possible for the paleonto-
 TAPHONOMY
FIGURE 4. Representative types of damage to the forelimb
resulting from ingestion by owls. A. Scapula, from left to
right: intact, "feathery," dorsal portion missing, distal end
only. B. Humerus: intact, proximal missing, distal end, dis-
tal missing, shaft only. C. Radius: intact, distal missing,
shaft only; Ulna: intact, distal missing, proximal missing.
logist to identify the agent of accumulation of
a given suite of bones. This approach is nascent
in taphonomy. Einarsen (1956) described the
effects of a variety of raptorial birds and carniv-
orous mammals on carcasses of their prey. Brain,
in an important series of papers (1967, 1969,
1976) has distinguished the effects of aboriginal
man and domestic dogs on carcasses from the ef-
fects of leopards on the same animals. The char-
acteristics of bone concentration by African por-
cupines (Hystrix) have been documented by
Hendey and Singer (1965) and Brain (in press).
Field studies of bone accumulation by hyaenas
and bat-eared foxes are currently under way in
Africa (A. K. Behrensmeyer, personal communi-
cation).
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OF OWL PELLETS 281
em
FIGURE 5. Representative types of damage to the hind-
limb resulting from ingestion by owls. A. Pelvis: upper left,
intact; lower left, damage to back; upper middle, damage
to front and back; lower middle, front missing; upper right,
ilium; middle right, ischium; lower right, pubis. B. Femur:
intact, damage (digestive erosion?) to proximal end, distal
missing, proximal only. C. Tibia-fibula, upper row: intact,
fibula missing, proximal end damaged, proximal end plus
fibula missing; lower row: distal end missing, distal end plus
fibula missing, shaft only, shaft only less fibula.
African porcupines (which are not predators
but which collect bone in prodigious quantities
and gnaw it) collect bones of species in propor-
tion to their actual abundance (Brain, in press).
Owls, by contrast, may be extremely selective
in their choice of food species. Pellets of barn
owls from northeastern Ohio (Dexter 1978) re-
vealed 14 of 28 species of small mammals living
in the region, and 97% consisted of the remains
of but three species (77% Microtus, 17% Bla-
282 PETER DODSON & DIANE WEXLAR
rina, 3% Peromyscus). A deposit that exhibits
a narrow range of size and taxonomic compo-
sition of prey items may be indicative of an ex-
tremely selective predator. Barn owls take 95%
of their prey in the mouse size range; screech
owls take 83% of their prey in this same size
range (Guilday et al. 1977) although insects are
also a significant food source. For great horned
owls, mice form but 33% of the diet, while rab-
bit- and squirrel-sized items constitute 57% of
the diet.
Owls are moderately to extremely destructive
to carcasses of small prey, though large prey
may be fastiduously flensed with minimal skel-
etal disarticulation or destruction (Einarsen
1956). Indeed, a careful study of pellet contents
in relation to food ingested over a period of time
for three species of owl (Asio otus, Strix aluco,
Tyto alba) showed that between 8 and 21% of
rodents ingested failed to show up in the pellets,
complete digestion of bone having taken place
(Raczynski and Ruprecht 1974). Young owls
have a greater tendency to digest bone than do
mature ones, evidently incorporating bone salts
from their prey into their owl skeletons. Bone
digestion by owls appears to be a strangely se-
lective process, delicate ribs often being re-
turned intact, for example. Digestive erosion on
the mandible of rodents, exposing the spongiosa
and tooth roots has been described (Raczynski
and Ruprecht 1974).
Due to their strong tendency to digest bone
(Duke et al. 1975), falconiforms (falcons,
hawks, eagles, etc., the diurnal predators of
Mayhew 1977) would seem to have a poor po-
tential for contribution to the fossil record.
Mayhew (1977), however, in analyses of pellets
of European kestrel (Falco tinnunculus) and
buzzard (Buteo buteo), has drawn attention to
the appearance of teeth and bones regurgitated
by falconiforms. Breakage is considerable
(though unquantified) and digestive erosion may
cause rounding of broken bone surfaces and of
tooth cusps, exposure of mandibular alveoli, re-
duction of enamel to a soft chalky consistency,
and erosion of teeth on non-occlusal surfaces.
He recognized these characteristics in more than
25% of isolated small mammal teeth in a sample
from a Middle Pleistocene deposit at West Run-
ton, England. Bone breakage alone is not evi-
dence of hawk predation. Digestive erosion of
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the mandible has been documented for owls
(Raczynski and Ruprecht 1974), and Korth (in
press) has produced abrasion by simulating
stream processes in a tumbling barrel. Used
with caution, however, it appears that the con-
dition of small mammal teeth can potentially
serve as reliable indicators of falconiform pre-
dation.
It might be surmised that the amount of bone
breakage of the prey should be inversely pro-
portional to the size of the owl. A mouse is a
large item to a small predator such as a screech
owl, but a small item to a large predator such
as a great horned owl, which can eat rabbits
also. Indeed, screech owls are the most destruc-
tive (Table 2). However, barn owls exhibit the
very important peculiarity of consistently re-
gurgitating a high proportion of undamaged,
often partially articulated bone (including intact
manus and pes, and articulated strings of ver-
tebrae). More than 80% of skulls were returned
intact by barn owls (though missing the occiput
and the back of the cranium), in contrast to the
3 to 12% of skulls returned by the other two
species. The results corroborate the finding of
Raczynski and Ruprecht (1974) that the barn
owl "digests its prey to a lesser extent than do
the other species."
Feeding Observations
The condition of regurgitated bone reflects in
part, at least, the mode of prehension and inges-
tion. Prey are seized in the talons and are swiftly
dispatched with a nip to the neck. Several rap-
tors were observed in feeding at the Academy
of Natural Sciences. A great horned owl took
a mouse transversely in its beak, transferred it
from mid-body forward in a series of 4 or 5 bites
(during which the sounds of breaking bones
(ribs, forelimbs, skull) could be heard), then
swallowed the mouse head first. The great
horned owl paid no attention to the brain, and
no damage to the hind portion of the body ap-
parently occurred during ingestion. A barn owl
held the mouse in its talon, carefully severed
the head from the neck (opening the cranium in
the process?) and swallowed it whole. It then
opened the thoracic cavity and fastidiously re-
moved through the enlarged thoracic inlet the
thoracic and abdominal viscera, which it neatly
 TAPHONOMY OF OWL PELLETS 283
consumed; finally it swallowed the hollowed out
carcass, neck first. Observed destruction of
bone during ingestion was thus minimal. Un-
fortunately the screech owls to which we had
access would not eat with observers present,
even in the quiet evening hours. A kestrel (Falco
sparverius) may constitute a suitable feeding
analogue. This small falcon, holding the mouse
in its talon, consumed its prey mouthful by
mouthful, commencing at the snout and system-
atically working its way caudally; each small
bite resulted in some bone breakage. The dam-
age resulting from this process is similar to that
generally discovered in screech owl pellets, in
which only 20% of bone escapes breakage.
Bones generally reside in the stomachs of owls
for 8 to 16 hr (average 10 to 13 hr) before being
cast (Duke et al. 1976; Duke and Rhoades
1977).
Depending upon their level of hunger, kes-
trels may cease feeding at any time, abandoning
a variably-sized caudal half of their carcass.
This is also observed for screech owls, which
may or may not consume the tail. This habit
may account for the depression of the relative
frequency of preservation of vertebrae in
screech owl pellets as compared to those of the
other two species. Raczynski and Ruprecht
(1974) called attention to the possibility that,
during times of rodent flushes when food is su-
perabundant, owls may selectively feed only on
rodent heads, resulting in unusual concentra-
tion of cranial remains.
Conclusions
The data presented in this report form a base-
line for characterizing accumulations of bone
resulting from food processing activities of sev-
eral species of owls. Important general charac-
teristics of owl pellet accumulations include 1.)
abundance and high quality of bone; 2.) possible
extreme inequitability of species distribution; 3.)
good representation of all skeletal parts includ-
ing both mandibles and vertebrae that contrast
strongly in their transport behavior (Dodson
1973); 4.) high representation of mandibles (pos-
sibly with damage to the articular region) and
complete femora relative to other skeletal parts;
5.) highly fragmented skulls with isolated max-
illae and premaxillae well represented, or intact
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skulls with occiput and cranial vault removed;
6.) heavily damaged scapulae and pelveses; 7.)
breakage of the proximal humeri, distal radii,
ulnae and femora; and 8.) heavy damage to
either end of the tibiae.
The three species of owl in this study, the
great horned owl (Bubo virginianus), barn owl
(Tyto alba) and screech owl (Otus asio) can be
clearly distinguished in terms of the destructive
effects on the bones of their prey: screech owls
cause breakage of 80% of the cranial and ap-
pendicular elements of their prey, compared to
65% for great horned owls and only 30% for
barn owls. Barn owls are particularly distinc-
tive in the low degree of breakage and high de-
gree of articulation of the bone produced. In
summary, the answers to the questions posed in
this study appear to be affirmative: that is, the
osteological characteristics of owl pellets are dis-
tinctive; the three species of owl can be distin-
guished in terms of their effect on their prey;
and the size of the predator is important in de-
terming the condition of the bone of the prey.
Owls date from the Eocene and are very
widespread geographically and ecologically in
the modern world. Thus through much of the
Cenozoic, they may have been important con-
tributors to the fossil record. This study is only
a starting point. Future studies of these species
should concentrate on the pellets of free-living
individuals. There are 130 species of owls in the
world; it is not known what degree of generality
these results may have. Many species of mam-
malian predators would be profitable subjects
for study. We believe that the approach out-
lined in this paper will prove to be a fruitful one
for extracting new levels of ecological insights
from the fossil record. We hope that this study
will prove a stimulus to further research.
Acknowledgments
We are very grateful to the workers of the
Animal Unit of the Academy of Natural Sci-
ences of Philadelphia, particularly Jerry Frei-
lich, Roger Clark and Marjan van der Schraaf.
They supplied us with quantities of pellets, kept
records of food consumption and pellet produc-
tion, freely offered us their own observations,
cheerfully answered our questions and gener-
ously premitted us to intrude and observe. Bob
284 PETER DODSON & DIANE WEXLAR
Callahan of the Philadelphia Zoo kindly sup-
plied us with pellets from the zoo. James Mellett
provided us with samples of wild carnivore scat
for comparative purposes. Doug Purvis of the
Laboratories of Anatomy, School of Veterinary
Medicine, provided us with technical assistance
and support at the initiation of our study. David
Weishampel, John Guilday, John Barry and
William Korth each provided us with a valuable
lead into the literature. Doug Purvis and David
McDevitt most generously donated their time
and services in the exacting task of preparing
photographs. Thanks also to two anonymous
reviewers for critical reading of the manuscript.
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