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JURASSIC DINOSAURS AND INSECTS: THE PALEOECOLOGICAL ROLE OF
TERMITES AS CARCASS FEEDERS
BROOKS B. BRITT
RODNEY D. SCHEETZ
ANNE DANGERFIELD
Earth Science Museum, Department of Geology, Brigham Young University, Provo, Utah 84602
USA <[email protected]>,<[email protected]>
ABSTRACT—Insect traces consisting of pits <1-20 mm wide are common on dinosaur bone
from the Upper Jurassic Morrison Formation in the western United States and are present on
bones from the Middle Jurassic of China. The pits occur on laminar bone in irregular clusters or
linear swaths. In some cases bone is consumed to a depth of more than a centimeter. Pitting is
most common on girdle and larger limb elements. Previously, the pits were interpreted as pupal
chambers of dermested beetles (Coleoptera). Pit morphology, size, patterns, deeply eroded
surfaces, and associated overlying burrows, however, match termite foraging traces on modern
bone. Thus termites are the responsible agent.
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The recognition that the pits are termite foraging traces is important because they: 1)
demonstrate termites originated at least by the Middle Jurassic and argue against the hypothesis
that termites could not have originated before the Early Cretaceous or Late Jurassic; 2) lend
support to identifications of Jurassic insect domichnia as termite mounds; 3) indicate termites
played a major, previously unrecognized, role in recycling nutrients from vertebrate carcasses;
and 4) show that in tropical and subtropical environments termites, not dermestids, are among
the last insects to infest carcasses.
Finally, termites consume exposed and buried bone of fresh to several-year-old carcasses.
Therefore, previous taphonomic interpretations involving supposed dermestids, which live only
on subaerially exposed, relatively fresh carcasses, must be reevaluated.
INTRODUCTION
MOST FOSSIL insect bone borings (Kubiak and Zakrzewska, 1974; Rogers, 1992; Paik, 2000)
and pits (Tobien, 1965; Martin and West, 1995; West and Martin, 1997; Hasiotis et al., 1999;
Laudet and Antoine, 2004) have been attributed to dermestid beetles. Short trails on modern
bones were attributed to lepidopteran larvae (Rivallain and Van Neer, 1983; Gautier, 1993) and
specifically, tineid moths (Behrensmeyer, 1978). Rivallain and Van Neer (1983) tentatively
associated shallow, flat-bottomed burrows on a 19th century turtle carapace to ants, although they
provided no explanation. Recently, pits and mandible marks on Neogene bone have been
attributed to termites (Kaiser, 2000; Kaiser and Katterwe, 2001; Fejfar and Kaiser, 2005). These
pits and marks compare well with those made by Australian termites (Watson and Abbey, 1986).
Shallow, circular pits, most with diameters of <10 mm, occur on dinosaurian bone from
the Upper Jurassic Morrison Formation. Vertebrate paleontologists have long been aware of
these pits but only recently have they been reported in the literature (Laws, et al., 1996; Hasiotis
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and Fiorillo, 1997; Hasiotis et al., 1999; Hasiotis, 2004). These authors identified the pits as
partial puparial chambers of dermestid beetles (Coleoptera, Dermestidae). Comparison of
feeding traces of the few modern insects known to attack bone, however, show the pits were
made by termites (Isoptera), not beetles. The pits are interpreted as termite feeding traces based
on morphology, linear clusters, and in-filled burrows.
Extant termites are notorious for their ability to eat wood and other plant materials
(Waller and La Fage, 1987a). They are also known to consume durable connective tissues, such
as ligaments and cartilage on carcasses (Coe, 1978). Less well known is their capacity to
consume bone (Derry, 1911; Behrensmeyer, 1978; Thorne and Kimsey, 1983; Tappen, 1994),
despite their ability to devour an entire human skeleton, leaving nothing but tooth crowns and
dwelling/transport chambers (Wylie, et al., 1987). Here, we present evidence that Jurassic
termites utilized dinosaur bone as a supplementary nutrient source.
ABBREVIATIONS
BYU, Brigham Young University, Provo, Utah; DNM, Dinosaur National Monument,
Jensen, Utah; GM, Gunma Museum, Gunma Prefecture, Japan; MOR, Museum of the Rockies,
Bozeman, Montana; MWC, Museum of Western Colorado, Grand Junction, Colorado; NAMAL,
North American Museum of Ancient Life, Lehi, Utah; SMM, Science Museum of Minnesota, St.
Paul, Minnesota; UMN, University of Minnesota, Minneapolis, Minnesota; UU, University of
Utah, Salt Lake City, Utah.
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METHODS
Pits are best observed with portable, directed, incandescent lamps, such as a mechanic’s
drop light, or fiber optic systems with the light at a low angle to the observed surface. Bones
with mottled coloring were whitened with ammonium chloride for photographic purposes.
LOCALITIES AND SPECIMENS
This study focuses on specimens from the Upper Jurassic Morrison Formation (Table 1)
but thousands of specimens from the overlying Lower Cretaceous Cedar Mountain Formation of
Utah, including BYU’s Dalton Wells and CEU’s Yellow Cat, Price River II, and Eolambia
quarries were also examined. Insect traces are common on the Lower Cretaceous bones but they
differ from those on Morrison Formation specimens and they will be described elsewhere. Pits
like those on Morrison bones are present on dinosaur bones from the Middle Jurassic Shishugon
Formation, Xinjiang Province, China (BBB, personal observation). We have not seen similar
pits on Late Cretaceous bones and we have not studied Triassic bones.
Bones from 20 dinosaur quarries (Table 1), ranging from isolated single skeletons to
large bone beds, all in the Brushy Basin Member of the Morrison Formation, were observed for
this study. Pitted bones occur in 70% (n=15) of the quarries. The SMM Poison Creek site bore
the highest percentage of pitted bones (49%), based on a random sample of 306 non-vertebral
bones of the over 400 bones collected. For logistical reasons, at no quarry were all bones
examined for pitting. For BYU’s Dry Mesa site, though, all theropod bones, n=779, were
observed and pits are present on 16%, which we extrapolate to be the average for the quarry
(Table 1). The most frequently pitted elements are femora, scapulocoracoids, and pubes with
pits on 17% to 55% of these bones. This concentration of pits on girdle elements and femora is
evident at all quarries where pitted bones are common.
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DESCRIPTION OF TRACE FOSSILS
Bones from the Morrison Formation exhibit two classes of pits. The rarest type, found on
a single skeleton in this study, has large, distinct mandible marks around bone pits. Similar starshaped Neogene traces are described by Kaiser (2000), Kaiser and Katterwe (2001), and Fejfar
and Kaiser (2005). By contrast, the trace fossils we describe here are common and consist of pits
that: 1) lack distinct mandible marks or have faint mandible marks on the rims and pit surfaces;
2) occur in clusters; 3) are present only on laminar bone; and 4) are not associated with
penetrating borings or surfical burrows.
The common pits are found occur primarily on one or two sides of a bone but it is not
uncommon for pits to occur on all non-articular surfaces. Bone from the Bone Cabin West
(NAMAL) quarries are usually pitted only on a couple surfaces while bones from the “Big Al”
(MOR) and Poison Creek (SMM) quarries, both in Wyoming, are commonly pitted on all sides.
Pits are roughly round, and occasionally elliptical (Fig. 1 & 2). Pit diameter ranges from
0.15-20 mm, with most being <10 mm. There is a continuous gradient of sizes, although one or
two sizes commonly dominate a pit cluster (Fig. 1.1). On a single bone pit sizes and spacing can
vary from fine, stipple-like concentrations to isolated pits (Fig. 2.8). Most pits are shallow, <2
mm (Fig. 1.1, 2.8-9), but they can be as deep as 5 mm (Fig. 2.3). Large pits can be smoothbottomed, negative hemispheres but most are composites of smaller, poorly-defined pits (Fig.
1.2, 2.1, 2.3, 2.7). Elliptical pits are generally elongate parallel to bone grain and are composites
of two or more subcircular pits (Fig. 1.1). To the naked eye most pits on dense bone appear to
have smooth margins (rims), but moderate magnification (6x) shows most pits have moderately
rough, irregular (scalloped) margins. Some pits exhibit faint scratch or “nibble” marks on the
surrounding rim. Pit details are seldom observed because they are often filled with matrix or
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covered with consolidant. Pits filled with matrix that contrasts with bone color are the most
obvious (Fig. 1.1).
Pits typically occur in clusters. The clusters, and patterns they form, are best seen on
bones that retain most of the original surface and are not deeply pitted. Loosely organized groups
of pits are referred to as patches or clusters. They are irregularly to roughly circular in outline,
with poorly defined edges (Fig. 1.5-1.8). Clusters can consist of small pits (Fig. 2.7, right side;
2.8, left center), a mixture of pit sizes (Fig. 1.4, right side; 1.5, 1.6), or large pits (Fig. 1.7, 1.8).
Less common than the generic clusters are lineations and rings. Rows ranging from
millimeters to meters long and one or more pits wide are here referred to as pit lineations, or
paths. These lineations can extend the full length of a long bone, as in the case of the anterior
face of a Camarasaurus femur, GM 101, but many are only several centimeters long (Fig. 2.1,
2.6, 2.7, 2.9). Lineations also occur at the sub centimeter level (top center of Fig. 2.8).
Pit lineations are occasionally associated with anatomical ridges. On an Allosaurus
fibula, BYU 5129, pits are present on various surfaces (Fig. 1.2) but occur in a line along the
caudomedial ridge (Fig. 1.3). Long rows of pits are found on the apex of the lateral ridges of the
cranial face of an Allosaurus chevron (Hanna, 2002, fig. 1.5) and rows of pits are evident on
sharp, low ridges of ligament scars on a Torvosaurus chevron (Fig. 2.9).
Furrows in the shape of a ring or nearly closed horseshoe (Fig. 3) occur on bone at most
quarries. Most rings are simple, with a single furrow (Fig. 3.1 and 3.3) but some are complex,
consisting of concentric rings resembling a flower (Fig. 3.2). Both ring morphs tend to occur in
clusters (Fig. 3.1, 3.2) and they are less common than simple pits. Rings have a table of
untouched bone (Fig. 3.3, solid white arrow; schematic cross section 3.5) or partially consumed
bone in the center (Fig 3.3 open white arrow and Fig. 3.6). Standard pits (Fig. 3.3, black arrow;
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Fig. 3.7) occur in clusters with rings and partial rings of the same diameter. Ring diameters
range from 2-15 mm. The rings are composed of closely-spaced small pits (Fig. 3.2).
In addition to pits and related traces on the bone itself, burrows are found in the matrix
surrounding some bones. The burrows are constructed in the matrix with one side of the burrow
in contact with the bone. One of the best examples of such burrows are found overlying the
pitted surface of a diplodocid sauropod scapula, MOR 057, from the MOR “Big Al” quarry in
Wyoming (Fig. 4). A number of disarticulated bones pertaining to the same individual are found
at the site and the bones are lightly to heavily pitted. On some specimens the burrows overlap, or
anastomose, essentially covering the bone with a mass of poorly defined burrows. The
undersides of the burrows are in contact with pits but the burrows do not penetrate the bone. The
burrows are broader than thick and the well delineated burrows shown in Figure 4 are 5 mm
thick and up to 40 mm wide. Areas beneath burrows are pitted while areas lacking visible
burrows are usually unpitted.
INSECTS THAT DAMAGE BONE
Pits on dinosaur bone from the Morrison, all of the same type described above, have been
previously reported by Laws, et al. (1996) Hasiotis and Fiorillo (1997), Hasiotis et al. (1999),
and Hasiotis (2004). Hasiotis et al. (1999) considered and ruled out a number of possible pitmakers, including soil and stomach acids, fungi, plants, and vertebrates. They determined that
insects, specifically dermestid beetles (Hasiotis, 2004) were the culprits. Other insects were
ruled out because they were thought incapable of excavating bone. Aside from dermestid
beetles, which can damage small delicate bones or less well ossified bone in the absence of flesh
(Sommer and Anderson, 1974), bone damaging insects include cloth moths and termites. Larvae
of the Christmas beetle, Anaplognathus have been found associated with burrowed bone
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(Haglund 1976), but the association is tenuous because termites were also present at the site.
Based on traces on modern bone we investigate tineid moths, dermestid beetles, and termites as
possible makers of the Jurassic pits.
Lepidoptera: moth traces.
The body fossil record of moths dates to the Early Jurassic (Davis and Robinson, 1999).
Most tineid moths (Lepidoptera, Tineidae) are detritivores, feeding primarily on fungi infected
debris (Davis and Robinson, 1999) but some are kerativores, feeding on the hair or feathers on
carcasses or debris in nests (Barbehenn and Kristensen, 2003). Tineids are found on carcasses in
the latter stages of decomposition – the dry and decay stages of Amendt et al. (2004) or stage 5
of Bornemissza (1957). The larvae feed on hair and other keratinous tissues and sometimes
excavate shallow trenches into bone just before pupation. The shallow excavations are less than
15 mm long and usually less than 4 mm wide and are usually found on horn cores (Rivallain and
Van Neer, 1983, and Gautier, 1992), where, we suggest, pupae were protected beneath the horn
sheath.
Coleoptera: dermestid traces.
The earliest body fossils of dermestid beetles are from the Oligocene (Carpenter, 1982).
Dermestids, which include carpet, hide, and lard beetles, are best known for their saprophagous
habits and osteologists are aware of their use in defleshing study skeletons (Sommer and
Anderson, 1974; Timm, 1982). The diet of dermestids, however, is quite varied. They feed on a
range of grains, feeds and animal products that contain protein, insects (eggs, larvae, and
carcasses), vertebrate carcasses (Hinton, 1945; Bornemissza, 1956) and even the flesh of live
birds (Samour and Naldo, 2003; Snyder, et al., 1984, and references therein) and they are known
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to kill small birds (Hinton, 1945). In the wild they commonly live in insect, bird, and rodent
nests and on vertebrate carcasses (Hinton, 1945). The larval stage is the most voracious phase of
the insect’s life but adults also feed on flesh (Schroeder et al., 2002). Following the last molt, the
mature larva immediately leaves the food (Kreyenberg, 1928), sometimes to a distance of 10
meters (Brimblecombe, 1938) and bores into “any substance at hand” (Illingworth, 1916),
including wood, mortar, cork, lead, and cloth (Kreyenberg, 1928; Brimblecombe, 1938; Hinton,
1945; Peacock, 1993; Schroeder et al., 2002; Panagiotakopulu, 2003) to avoid predation during
the pupal stage. Only members of the genus Dermestes burrow to pupate (Peacock, 1993).
Burrowing, however, is not required for pupation, e.g. in Dermestes lardarius, larvae that do not
burrow pupate within the last larval skin while those that burrow use the skin to cap the pupation
chamber (Kreyenberg, 1928; Hinton, 1945). Pupation burrows are essentially straight and range
from the length of the larva (Hinton, 1945) to 31 cm (Day, 1922) with a diameter slightly larger
than the larva (Kreyenberg, 1928) and the pupation chamber at the end of the tunnel. The borings
are round to oval in outline with sharp edges and steep walls (Kreyenberg, 1928; Roberts et al.,
2003). The pupation chambers are ovate in cross section (Kreyenberg, 1928, fig. 21).
With the exception of Schroeder et al. (2002), forensic studies report no dermestid
damage to bone (e.g., Smith, 1986; Iverson et al., 1996; Panagiotakopulu, 2003; Amendt et al.,
2004). Hinton (1945) reports damage to bone but does not elaborate and most of his references
simply note that shipments of bone were infested with dermestids, but they were probably
feeding on remnants of flesh as there is no mention of burrows or borings. Roberts et al. (2003)
noted larvae harvest marrow from larger bones but they can access the interior only if the bones
have been broken, drilled, or have natural entrances, such as large nutrient foramina (Stephen
Hinshaw, personal commun. to BBB, 2005) Schroeder et al. (2002) reported “feeding defects”
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on bone of a human corpse consisting of pits in the acetabulum and on the articular surface on
the proximal end of the humerus. The largest fossa is about 10 mm long, ovate, and may be a
pupal chamber. Most pupation, however, occurred in furniture, the upholstery of which was
riddled with holes. It is significant in this case that damaged bone was limited to articular
surfaces. It is possible that the hyaline cartilage was among the last soft tissues consumed and
the larvae ate through the cartilage into the relatively soft articular, grease-laden bone. The
notion that dermestids commonly bore into bones is likely based on a casual familiarity with
museum defleshing cages. In fact, it is rare for dermestids to damage bones in defleshing boxes,
unless the bones are poorly ossified (Stephen Hinshaw, personal commun. to BBB, 2005) and
the soft tissue supply exhausted (Roberts et al., 2003).
Isoptera: termite traces.
The oldest termite body fossils are from the Early Cretacous (Lacasa-Ruiz and MartinezDelclòs, 1986). Termites are renowned for their ability to recycle plant materials and infamous
for the damage done to wood structures. Although the complex architecture of termite galleries
has been widely studied and described (e.g., Lys and Leuthold, 1991), scant information on
surficial forage traces is available in the literature and even less is written about their ability to
consume bone. The following description of foraging traces on modern bone is based primarily
on Tappen (1994) and our observations of modern elephant bones (Fig. 2.4, 2.5) on which
termites were observed feeding in a tropical rainforest in Zaire (Martha Tappen, personal
communication to BBB, 2005). The primary traces left by termites foraging on bone are
abundant pits (Tappen, 1994) which usually overlap, giving the bone a pock-marked or eroded
(Wood, 1976) appearance. On thin bones, as in some human skull elements, termite marks may
be perforate (Wood, 1976). Linear groups of pits are present along termite foraging trails which
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give the trails their characteristic lobate margins (Fig. 2.2). Similar foraging trails are made by
termites on wood (Ebeling, 1975, fig. 72A). Pits vary in size from < 1 to at least 13 mm. All but
the smallest pits are composites of several nested or overlapping pits (Fig. 2.2, 2.4, 2.5).
Margins of most composite pits are ill-defined (Fig. 2.4, 2.5). Some modern bone with a scraped
appearance preserves what may be faint mandible traces (Thorne and Kimsey, 1983). Many pits,
however, lack definite mandible marks and preserve ledges representing layers of bone (Tappen,
1994, fig. 8). Some composite pits are smooth walled, save for bone texture which is a function
of vascularization (Fig. 2.4). Foraging galleries are covered with carton (Thorne and Kimsey,
1983) or earthen mounds where bones are in contact with the ground (Tappen, 1994). The
protective gallery walls are made of mineral grains, soil, or organic materials cemented with a
mouth secretions or feces, or simply feces (Lee and Wood, 1971).
Determination of trace maker.
A comparison of traces made in firm materials by moths, dermestids, and termites is
presented in Table 2. Tineid moths, which make simple, linear pupal furrows, are clearly not
responsible for the Morrison pits. Dermestid beetle traces share little in common with Morrison
pits (Table 2). Dermestid feeding traces consist of pits, but they are rare on bone, occurring only
on poorly ossified bones such as bones of juveniles or articular surfaces. Pupation borings are
narrow and elongate, usually straight, and do not overlap (Day, 1922) or are ovate and deep
creating a Swiss cheese effect (Kreyenberg, 1928). Both feeding and pupation burrows rarely
impact bone and they do not occur in patterns. Most invertebrate trace fossils in bone, whether
penetrating borings and burrows (Kubiak and Zakrzewska, 1974; Rogers, 1992; Paik, 2000), or
pits (Tobien, 1965; Martin and West, 1995; West and Martin, 1997; Laws, et al., 1996; Hasiotis
and Fiorillo, 1997; Hasiotis et al., 1999; Hasiotis, 2004; Laudet and Antoine, 2004), have been
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interpreted as dermestid pupation chambers. Probable dermestid pupal chambers like those
figured by Kreyenberg (1928) are found exclusively on Neogene bone (Laudet and Antione,
2004, and references therein).
The most detailed paper identifying Morrison pits as dermestid pupal chambers is
Hasiotis et al. (1999). That identification was made primarily because they considered beetles as
the only insects capable of boring bone, and dermestids were known to excavate pupal chambers.
They also argued the identification based on pit size, morphology, and evidence of an arid
climate for Morrison environs. They reported that pits occurred in limited size ranges; 0.5-1
mm, 2.5-3 mm, and 4-5 mm. Our observations, however, indicate the pit diameters cover the
spectrum between 0.15 mm to 20 mm. The pits were interpreted as the bottoms of pupation
chambers, assuming the balance of the pupal chamber was present in overlying dried tissues.
This caveat was necessary because complete dermestid pupation chambers are at least as deep as
the length of the larva. The pits on Morrison bone bear no resemblance to dermestid pupation
borings but they are vaguely similar to the dermestid feeding pits shown in Schroeder et al.
(2002). The difference is that the modern dermestid pits occur only on soft articular surfaces
while those in the Morrison occur only on dense laminar bone.
The characteristic pock-marked surfaces excavated by termites on modern bone consist
of pits of varying sizes, resulting in loss of much of the bone surface and, in extreme cases,
consumption of entire bones (Derry, 1911) or skeletons (Wylie, et al., 1987). Foraging trails are
marked by swaths of pits. Fine mandible marks are present on some modern bone (Thorne and
Kimsey, 1983; Watson and Abbey, 1986; personal observation). The ledged pits shown in
Hasiotis et al. (1999) match those figured for known termite pits (Tappen, 1994, fig. 8). Thus, the
pits so abundant on Morrison Formation bone are strikingly similar to modern termite foraging
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traces. The rarity of mandible marks on fossil bone likely relates to the termite taxon. Watson
and Abbey (1986) found that mandible marks on bone made by three termite genera varied in
size and density, and ranged from clear marks to partial “nibble” marks. Most modern termite
traces found on African elephant bone lack mandible marks.
Although pits are common on both modern and fossil termite impacted bone, rings have
only been reported on fossil bone. The rings and partial rings on Morrison bone are composed of
closely spaced pits added en echelon to form an arc. The arc may connect with the point of
origin to form a ring but often forms a nearly closed horseshoe-shape with a central table, or
plateau, of untouched or slightly consumed bone (Fig. 3.3, solid white arrow; Fig. 3.5). The
central table is gradually consumed leaving a lower table or dome (Fig. 3.3 open white arrow;
Fig 3.6) and the end result is a pit (Fig. 3.3, black arrow; Fig. 3.7). Rings in various stages of
development with an intimate association with standard pits, and the fact that mature rings are
pits, indicate the rings were also made by termites. The rings on fossil bone may reflect the
behavior of an extinct termite or simply a feeding pattern not yet observed in modern termites.
We speculate the rings are made by a single individual rotating about a fixed point and that the
unusual concentric patterns (Fig. 4.2) may be taxonomically diagnostic.
It is probable that the burrows that overlie pitted bones were made by termites. The
burrows are found only on pitted areas of the bone and the association between burrows and
pitting suggests the burrows represent foraging tubes. Some modern foraging tubes are actively
backfilled by termites (Lys and Leuthold, 1991) so finds of additional fossil burrows could be
thin sectioned and compared with termite backfill architecture (Lee and Wood, 1971). If the
burrows are termite foraging tubes, then termites harvested the bones in situ, that is, after burial.
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Excavation and preparation of bone from sites with pitting should be done with an eye for
burrows and carton.
TERMITES AND BONE
Osteophagy.
Some termites have at least a seasonal affinity for bone and have the capability to
consume an entire juvenile human skeleton (Wylie, et al., 1987). Behrensmeyer (1978) observed
termites on bones impacted by invertebrates and noted that termite-degraded bones were
common in some areas. In these cases, however, the termites were not been observed in the act.
The dearth of feeding observations is, in part, a function of bones being attacked long before
their discovery (e.g., Derry, 1911 – Egyptian and Nubian human remains; Light, 1929 – human
skull; and Wood, 1976 – Australian aboriginal remains). Furthermore, termites generally feed
while enclosed in galleries or underground (Ebeling, 1975). Fortunately, though, there are
several first-hand reports of termites feeding on bone. Thorne and Kimsey (1983) observed
termites consuming bones of vertebrate carcasses (boa, turtle, sloth, and agouti – a large rodent)
in a Panamanian tropical rainforest and Watson and Abbey (1986) induced termites to damage
bone in the laboratory. Termites are known to cause extensive damage to human skeletons in
rock shelters in Queensland, Australia (Wylie, et al., 1987) and Tappen (1994) observed termites
consuming elephant bones exposed for several years in a tropical rainforest in Zaire. Coe (1978)
saw the termite Odontotermes zambesiensis (Sjost.) consume intervertebral disks and hyaline
cartilage on two-year-old elephant carcasses. The same termites constructed foraging channels of
carton over bones, but there was no mention of bone consumption. Clearly, termites consume
bone, begging the question, “Why?”
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Bone and nutrition.
Termites feed on an array of materials, including live and dead plants, partially decayed
plants, dung, fungi, and carcasses (Waller and La Fage, 1987b). Most termite foods are
nutritionally depauperate (Waller and La Fage, 1987a) and it has been proposed that termites
seek out carcasses to supply phosphorous and nitrogen (Thorne, et al., 1983; Watson and Abby,
1986). Bone is a composite material comprised of about 65% (dry weight) hydroxylapatite, the
hard inorganic component, and about 35% collagen fibers, which forms a flexible matrix
(Francillon-Vieillot, et al., 1990). Hydroxylapatite, Ca5(PO4)3(OH), is potentially a source of
calcium and phosphorous. Nitrogen is a limiting factor in termite colonies (Traniello and
Leuthold, 2000, and references therein) and in bone this is found in collagen, a protein, but the
amino acids that make up collagen may, themselves, be the targeted compound. Because the
traces are found on dense laminar bone, not collagen-rich articular surfaces, phosphorous and/or
calcium are probably the nutrients sought by the termites. In any case, the pits and deeply
excavated surfaces on bone indicate bones were a source of food, i.e., the bones were not
casually probed or used as byways.
Foraging Techniques.
Termites are eusocial insects which divide nest/colony tasks among various casts
(soldiers, several grades of workers, queen, and king). Foraging routines vary according to
taxon, but in general the search for new food sources is usually led by soldiers, followed by
workers. The search is conducted above or below ground depending on whether the food is
disseminated or concentrated, respectively (Lee and Wood, 1971). Although the method by
which they find food is poorly understood because of their cryptic nature, it is known that at least
some subterranian termites utilize non-random geometric search patterns, while other termites
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utilize thermal shadows or chemical detection (Traniello and Leuthold, 2000). Once a new food
source is found, galleries consisting of a protective tube of carton or earth, or an excavated
tunnel, are constructed to connect the colony with the food source (Lys and Leuthold, 1991;
Traniello and Leuthold, 2000). Similar galleries are constructed on and in the food source,
including bones (Derry, 1911; Thorne, et al., 1983; Wylie, et al., 1987). Gallery coverings
protect termites from predatory invertebrates and help reduce water loss (Lee and Wood, 1971)
but in semiarid environments workers must descend to humid galleries to rehydrate (Ebeling,
1975). As harvesting continues, the entire food source can be covered with soil gallery
complexes or encased in saliva-cemented mud/sand or carton – a paper-like material like similar
to a wasp nest (Derry, 1911; Lys and Leuthold, 1991). By the time the food source, whether
wood or bone, is exhausted, only a thin covering of carton remains which mimics the original
shape of the now absent food source (Derry, 1911; Wylie, et al., 1987). Foragers, usually
workers, carry food particles into the nest’s interior for processing. A single colony can contain
over 6 million foragers with a range of 100 m, in the case of the Formosan subterranean termite,
Coptotermes formosanus, Shiraki (Su and Scheffrahn, 1988). A single Macrotermes bellicosus
colony has about 7 km of galleries within a 25 m radius, and in the savannas of Africa this
species has a density of about 33 colonies per hectare (Lys and Leuthold, 1991). With such large
numbers of individuals, termite colonies can have a major impact on food sources, including
bones.
Known osteophagous taxa.
Extant species known to consume bone in the wild are Nasutitermes carnarvonensis
(Hill) (Wylie, et al., 1987) and Nasutitermes nigriceps (Haldeman) (Thorne and Kimsey, 1983) a
neotropical wood feeder. Termite species suspected of damaging bone in archaeological sites
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were tested in the laboratory, showing that they could be induced to leave marks on bone. These
include unspecified species of Microcerotermes and Mastotermes as well as Coptotermes
ancinaciformis (Watson and Abbey, 1986). The termites observed in foraging trails on elephant
bone from Africa were not identified to genus. It is significant that bone consumption is not
limited to a single species of termites and includes genera from three families: Mastotermitidae
(Mastotermes), Termitidae (Nasutitermes and Microcerotermes), and Rhinotermitidae
(Coptotermes). Thus, osteophagy is relatively widespread both taxonomically, occurring in three
of seven total isopteran families, and geographically as well (Australia, Central America, and
Africa).
Geography and Environments.
Extant termites occur on all continents save for Antarctica. Most species are found in
tropical to semitropical environments, with the greatest diversity within 10º north or south of the
equator in tropical forests, but they also inhabit semiarid regions (Eggleton, 2000). Humidity is,
to a degree, a controlling factor, but termites can inhabit semiarid environments because
underground nests provide access to moisture (Ebeling, 1975). Modern pitted bones have been
reported from tropical rainforests to semiarid regions. Tappen (1994) found a large percentage
(59%) of bones in a tropical rainforest in Zaire were termite pitted, while in a nearby savanna
environment <1% were pitted, even though termites were abundant. Termite pits on bone have
also been reported from mboseli Park (Behrensmeyer, 1978) and Lake Baringo (Tappen, 1994),
Kenya, both of which are savanna environments. In Australia, termites consume bone in rock
shelters in the semiarid Carnarvon region of Queensland where the humidity was higher than the
unprotected environment (Wylie, et al., 1987). In a moist tropical island off the coast of Panama
termites infest carcasses in the dry season about 15 days postmortem, after most organ and
18
muscle tissues are lost to scavenging (Thorne and Kimsey, 1983). The termites then consume
bone under the protection of carton foraging tubes and make carton “nests” in the cranium and
pelvic areas. Termites abandon the carcasses with the first precipitation of the rainy season.
Phylogeny.
The Isoptera (termites) are closely related to Blattaria (cockroaches) and Mantodea
(mantids), which together comprises the Dictyoptera. The phylogenetic relationships within
Dictyoptera, however, remains to be resolved (Thorne, et al., 2000) although Blattaria
(specifically, the wood eating cockroach, Cryptocerus) and Isopera are traditionally considered
as sister groups (Nalepa and Bandi, 2000). The earliest termite body fossils are from the Early
Cretaceous (Upper Berriasian to Lower Valanginian) of Spain (Lacasa-Ruiz and MartinezDelclòs, 1986). Using molecular data and body fossils, it has been argued that termites likely
originated no earlier the Early Cretaceous (Nalepa and Bandi, 2000) or Late Jurassic (Thorne, et
al. 2000). If, however, Blattaria and Isoptera are sister taxa, then the origin must have been as
early as the late Carboniferous (Thorne, et al. 2000), an idea supported by the geographical
distribution of extant clades, suggesting all termite families originated prior to the breakup of
Pangaea (Eggleton, 2000). Domichnia trace fossils, interpreted as termite mounds, have been
reported from the Late Triassic (Norian) Chinle Formation (Hasiotis and Dubiel, 1995), the Early
Jurassic Clarens Formation of South Africa (Bordy, 2004) and the Late Jurassic Morrison
Formation (Hasiotis et al, 1997; Hasiotis, 2000; Hasiotis, 2002). Thorne et al. (2000) dismissed
such early trace fossils as coleopterid (beetle) galleries. The recognition that termites were
abundant in the Late Jurassic Morrison Formation based on feeding traces, however, suggests the
galleries were correctly identified as termite mounds.
19
COMPARISON OF DERMESTIDS AND TERMITES
The environments and feeding conditions pertaining to dermestids and termites are
compared in Table 3. With the exception of the temperate zones, termites impact bone under a
wider variety of conditions than dermestids. Furthermore, because they are colonial, termites
can impact a carcass more severely and rapidly than dermestids, which require time for eggs to
hatch (days) and probably time for a second generation hatch (weeks) before full infestation.
Because termites can control temperature and humidity within the nest and to a lesser degree in
foraging tubes, they function effectively in deserts and tropical rain forests. Dermestids, on the
other hand, exert no control over their environment and the growth of larvae is adversely
impacted by conditions of low humidity and temperature (Kreyenberg, 1928; Hinton, 1945;
Ebeling, 1975).
Studies of extant termite carcass consumption reveals: 1) bone consumption by isopterans
is known from China at 31º N and Egypt at 30º N to the Brooloo Range, Queensland, Australia
at 26º S; 2) termites infest carcasses in both semiarid and humid environments; 3) all bone
surfaces may be affected, but the underside-substrate contact seems to be preferred (Tappen,
1993); 4) the heaviest infestations are in the pelvis and cranium; 5) bone consumption is
deliberate and provides critical nutrients to the colony; 6) bone is utilized as long as nutrients are
present, which may be years after death, long after dermestids and most other saprophagous
insects abandon carcasses; and 7) exposed carcass infestation is limited to the dry season.
CONCLUSIONS
We concur with Roberts et al. (2003) that care should be taken when assigning trace
fossils to a maker. In this study, traces on fossil bone primarily from the Upper Jurassic
20
Morrison Formation are compared to equivalent modern traces on the same substrate – bone.
The traces compare in general ways, such as pit size and morphology, which reflects the
excavation method and size of the organism. Additionally, the organization of pits into clusters
and lineations reflects harvesting behavior by large numbers of individuals along broad areas and
trails (galleries), respectively. Consequently we have a high degree of confidence that the pits
were made by termites. We itemize our findings as follows:
1. Invertebrate trace fossils on bones, consisting of pits of various sizes, some in linear or
circular clusters, occur on bones in the majority of quarries in the Brushy Basin Member of the
Morrison Formation. When pits are present they occur on <1% to 49% of the bones.
2. The morphology, size, and distribution patterns of the trace fossils on bone match foraging
traces made by extant termites on bone. Therefore, termites, not dermestid beetles, likely created
the fossil traces.
3. Rejection of the dermestid hypothesis means there is currently no trace or body fossil evidence
of dermestids prior to the Oligocene.
4. The recognition of Middle and Late Jurassic termite traces indicates Isoptera originated even
earlier, supporting the hypothesis that Blattoidea and Isoptera are sister taxa and that termites
originated in the late Paleozoic.
21
5. Termites were abundant in Morrison Formation environments, at least in Utah, Colorado, and
Wyoming and played a major role in recycling carcass nutrients. They may have also played a
major role in vegetal recycling and soil modification.
6. The termite foraging traces in the Late Jurassic Morrison Formation support the interpretation
of large, mound-shaped domichnia in same formation as termite mounds.
7. Extant termites are most abundant in tropical to subtropical latitudes and termite traces on
bone are more common in tropical rain forests than in semiarid areas. Thus termite traces do not
necessarily indicate a semiarid climate. In tropical climates, termite consumption of bone occurs
only during the dry season.
8. The standard model in which dermestid beetles are presumed be the last scavengers of dry,
and even weathered carcasses is incorrect. Dermestids inhabit a carcass only as long as soft
tissues are present, whereas termites consume bare bone days to years following death. The
taphonomic implications of this timing are significant.
9. Bone buried at the time of termite utilization could be burrowed on all sides and may preserve
foraging tubes, whereas bones of exposed carcasses are more likely to be burrowed on the
undersurface, (i.e., the bone-substrate contact).
Additional studies of bone consumption by modern termites are needed. With the data at
hand, however, paleotaphonomic studies can show whether termites harvested bone shortly after
22
death or much later by determining the cross cutting relationships of weathering cracks and
pitting. If the termites were primarily wood-feeding, termite burrows will be found in fossil logs
in the formation, whereas if they were primarily litter-feeding, foraging tubes will be in the
uppermost layers of paleosols. In the absence of pre-Cretaceous termites body fossils, and the
rarity of Mesozoic and older insect body fossils, termite foraging traces on bone can be used to
better bracket the timing of the origin of termites.
The termite interpretation of these traces fossils can be tested. If extensive foraging
trails, carton, or back-filled foraging tubes are found associated with pitted bones, it would
corroborate the hypothesis. Such foraging tubes will be difficult to recognize during fossil
preparation unless the preparator is conscious of the subtle differences in the matrix near the
bone. If careful searches do not produce such structures, the hypothesis may be rejected. Data
presented above, however, indicate that the termite hypothesis is currently the most parsimonious
option.
ACKNOWLEDGEMENTS
We are most grateful to M. Tappen for providing unpublished information on modern termite
pitted bone. S. VanderVoss’s keen eye and careful preparation of MOR specimens was
invaluable. B. Erickson graciously provided access the SMM collections and D. Waddington’s
help with forensic literature was invaluable. C. Miles generously donated critical specimens. N.
Anderson, P. Valora, M. Pickard, and T. Tomlin assisted with data collection and D. Turley
helped with imaging. We especially thank R. Beal, A. Herrmann, J. Kingsolver, and S. Hinshaw
for dermestid beetle information. BBB was funded by grants from BYU’s College of Physical
23
and Mathematical Sciences and AD was funded by a BYU ORCA grant and the BYU
undergraduate mentoring program.
REFERENCES
AMENDT, J., KRETTEK, R., AND ZEHNER, R. 2004. Forensic entomology.
Naturwissenschaften, Vol. 91: 51-65.
BARBEHENN, R .V., AND KRISTENSEN, N. P. 2003. Digestive and excretory systems.
Handbook of Zoology, Volume IV, Arthropoda: Insecta. Kristensen, N. P., (ed.)
Lepidoptera, Moths and Butterflies, Volume 2: Morphology, Physiology, and
Development, Pt. 36, p. 165-187.
BEHRENSMEYER, A.K. 1978. Taphonomic and ecologic information from bone weathering.
Paleobiology, Vol. 4, p. 150-162.
BORDY, E. M., BUMBY, A. J., CATNEANU, OCTAVIAN, ERIKSSON, PATRICK G. 2004.
Advanced Early Jurassic Termite (Insecta: Isoptera) Nests: Evidence from the Clarens
Formation in the Tuli Basin. Southern Africa Palaios, Vol. 19, p. 68-78.
BORNEMISSZA, G. F. 1957. An analysis of arthropod sucession in carrion and the effect of its
decomposition on the soil fauna. Australian Journal of Zoology, Vol. 5, p. 1-12.
BRIMBLECOMBE, A. R. 1938. Hide beetle damage to Oregon pine. Journal of the Australian
Institute of Agricultural Science, Vol. 4, No. 1, p. 49-50.
CARPENTER, F. M. 1982. Treatise on invertebrate paleontology. Superclass Hexapoda. Pt. R,
Arthropoda 4, Geological Society of America and University of Kansas Press, 655 p.
COE, M. 1978. The decomposition of elephant carcasses in the Tsavo (East) National Park,
Kenya. Journal of Arid Environments, Vol. 1, p. 71-86.
24
CROWSON, R. A. 1981. The Biology of the Coleoptera. Academic Press, London, 802 p.
DAVIS, D. R., AND ROBINSON, G. S. 1999. The Tineoidea and Gracillarioidea, p. 91-117.
In N.P. Kristensen (ed.), Handbook of Zoology: A natural history of the animal kingdom,
Vol. IV, Arthropoda: Insecta, Pt. 35. Lepidoptera, Moths and Butterflies Volume 1:
Evolution, Systematics, and Biogeography. Walter de Gruyter, New York.
DAY, F. H. 1922. Dermestes lardarius L., feeding on wood. Entomological Monograph
Magazine, Vol. 58, p. 209.
DERRY, D. E. 1911. Damage done to skulls and bones by termites: Nature, Vol. 86, p. 245–
246.
EBELING, W. 1975. Urban Entomology. University of California, Division of Agricultural
Sciences, Los Angles, 695 p.
EGGLETON, P. 2000. Global patterns of diversity, p. 25-51. In T. Abe, D. E. Bignell, and M.
Higashi (eds.), Termites: Evolution, Sociality, Symbiosies, Ecology. Kluwer Academic
Publishers, Dordrecht, Netherlands.
FRANCILLON-VIEILLOT, H., DE BUFFRÉNIL, V., CASTANET, J. GÉRAUDIE,
MEUNIER, F.J., SIRE, J.Y., ZYLBERBERG, L., AND DE RICQLÈS, A. 1990.
Microstructure and mineralization of vertebrate skeletal tissues, p. 471–548. In J. G.
Carter, (eds.), Skeletal biomineralization: patterns, processes and evolutionary trends,
Vol. 1. Van Nostrand Reinhold, New York.
FEJFAR, O., AND KAISER, T. M. 2005. Insect bone-modification and paleoecology of
Oligocene mammal-bearing sites in the Doupov Mountains, northwestern Bohemia.
Palaeontologia Electronica, Vol. 8, Issue1:8A:11p, 1 MB; http//palaeoelectronica.org/;aloe/2005_1fejfar8/Issue1_0505.htm
25
GAUTIER, A. 1993. Trace fossils in archaeozoology. Journal of Archaeological Science, Vol.
20, p. 511-523.
HAGLUND, L. 1976. The Broadbeach Aboriginal Burial Ground. University of Queensland
Press, Brisbane.
HASIOTIS, S. T. 2000. The invertebrate invasion and evolution of Mesozoic soil ecosystems:
The ichnofossil record of ecological innovations, p. 141-169. In R. Gastaldo and W.
DiMichele (eds.), Phanerozoic Terrestrial Ecosystems: Paleontological Society Short
Course, Vol. 6.
HASIOTIS, S. T. 2002. Complex ichnofossils of solitary and social soil organisms:
understanding their evolution and roles in terrestrial paleoecosystems. Palaeogeography,
Palaeoclimatology, Palaeoecology, Vol. 192, p. 259-230.
HASIOTIS, S. T. 2004. Reconnaissance of Upper Jurassic Morrison Formation ichnofossils,
Rocky Mountain Region, USA: paleoenvironmental, stratigraphic, and paleoclimatic
significance of terrestrial and freshwater ichnocoenoses. Sedimentary Geology, Vol. 167,
p. 177-268.
HASIOTIS, S. T., AND DUBIEL, R. R. 1995. Termite (Insecta: Isoptera) nest ichnofossils
from the Upper Triassic Chinle Formation, Petrified Forest National Park, Arizona.
Ichnos, Vol. 4, p 119-130.
HASIOTIS, S. T., AND FIORILLO, A .R. 1997. Dermestid beetle boring in sauropod and
theropod dinosaur bones, Dinosaur National Monument, Utah – keys to the taphonomy of
a bone bed. Combined Rocky Mountain/Southcentral Geological Society of America
Meeting, Abstracts with Program, El Paso, Texas, Vol. 29, No. 2, p. 13.
26
HASIOTIS, S. T., FIORILLO, A. R., AND HANNA, R. R. 1999. Preliminary report on borings
in Jurassic dinosaur bones: Evidence for invertebrate-vertebrate interactions, p. 193-200.
In D. Gillette (ed.), Vertebrate Paleontology in Utah, Utah Geological Survey
Miscellaneous Publication 99-1.
HASIOTIS, S. T., PETERSON, F., DEMKO, T. M., AND TURNER, C. 1997. Giant termite
(Insecta: Isoptera) nests from the base of the Upper Jurassic Morrison Formation,
northwestern New Mexico. Geological Society of America Abstracts with Program, Vol.
28, p. A461.
HINTON, H. E. 1945. A monograph of the beetles associated with stored products, Vol. 1,
British Museum (Natural History), London, 443 pp.
ILLINGWORTH, J. F. 1916. Notes on the life-history of Dermestes cadaverinus Fab.
Proceedings of the Hawaii Entomological Society, Vol. 3, No. 3, p. 255-257.
IVERSON, K., OCONNOR, B. M., OCHOA, R., AND HECKMANN, R. 1996. Lardoglyphus
zacheri (Acari: Lardoglyphidae), a pest of museum dermestid colonies, with observations
on its natural ecology and distribution. Annals of the Entomological Society of America,
Vol. 89, No. 4, p. 544-549.
KAISER, T. M. 2000. Proposed fossil insect modification to fossil mammalian bone from PlioPleistocene hominid-bearing deposits of Laetoli (northern Tanzania). Annals of the
Entomological Society of America, Vol. 93, No. 4, p. 693-700.
KAISER, T. M., AND KATTERWE, H. 2001. The application of 3-D-microproofilometry as a
tool in the surface dagnosis of fossil and sub-fossil vertebrate hard tissue. An Example
from the Pliocene Upper Laetolil beds, Tanzania. International Journal of
Osteoarchaeology, Vol. 11, p.350-356.
27
KREYENBERG, J. 1928. Experimentell-biologische untersuchungen über Dermestes lardarius
L. und Dermestes vulpinus F. Ein Beitrag zur frage nach der Inkonstanz der
häutungszahlen bei Coleopteren. Eitschrift fuer Angewandte Entomologie, Vol. 14, No.
1, p 140-188.
KUBIAK, H., AND ZARKRZEWSKA, G. 1974. Fossil mammals in Upper Paleolithic site
with dwellings of mammoth bones. Folia Quaternaria, Vol. 44, p. 77-79.
LACASA-RUIZ A, MARTINEZ-DELCLÒS X. 1986. Meiatermes: nuevo género fosil de
insecto Isoptero (Hodotermitidae) de las calizas Neocomienses del Montsec (Provincia de
Lérida, Espana). Publ. Inst. d'Estidis Ilerdencs Diput. Prov. Lléida. p. 5-65.
LAUDET, F., AND ANTOINE, P. O. 2004. Des chambres de pupation de Dermestidae
(Insecta : Coleoptera) sur un os de mammifère tertiaire (phosphorites du Quercy):
implications taphonomiques et paléoenvironnementales. Geobios, Vol. 37, Issue 3, p.
376-381.
LAWS, G. R., HASIOTIS, S. T., FIORILLO, A. R., CHURE, D., BREITHAUPT, B. H., AND
HORNER, J. 1996. The demise of a Jurassic dinosaur after death – three cheers for the
dermistid (sic) beetle. Geological Society of America National meeting, Abstracts with
Program, Vol. 28, No. 7, p. 299.
LEE, K. E., AND WOOD, T. G. 1971. Termites and Soil. Academic Press, London, 251p.
LIGHT, S.F. 1929. Present status of our knowledge of the termites of China. Lignan Science
Journal, Vol. 7, p. 581-600.
LYS J. A., AND LEUTHOLD., R. H. 1991. Morphology of the gallery system around the nest
and gallery development under experimental conditions in the termite Macrotermes
bellicosus (Smeathman). Insectes Sociaux, Vol. 38, p. 63-76.
28
MARTIN, L. D., AND WEST, D. L. 1995. The recognition and use of dermestid (Insecta,
Coleoptera) pupation chambers in paleoecology. Palaeogeography, Palaeoclimatology,
Palaeoecology, Vol. 113, p. 303-310.
NALEPA, C. A., AND BANDI, C. 2000. Characterizing the ancestors: paedomorphosis and
termite evolution, p. 53-75. In T. Abe, D. E Bignell, and M. Higashi (eds.), Termites,
Evolution, Sociality, Symbiosis, Ecology. Kluwer Academic Publishers, Netherlands.
PAIK, I. S. 2000. Bone chip-filled burrows associated with bored dinosaur bone in floodplain
paleosols of the Cretaceous Hasandong Formation, Korea. Palaeogeography,
Palaeoclimatology, Palaeoecology, Vol. 157, p. 213-225.
PANAGIOTAKOPULU, E. 2003. Insect remains from the collections in the Egyptian Museum
of Turin. Archaeometry, Vol. 45, No. 2, p. 355-362.
PEACOCK, E. R. 1993. Adults and Larvae of Hide, Larder and Carpet Beetles and their
relatives (Coleoptera: Dermestidae) and of Derodontid Beetles (Coleoptera:
Derodontidae). Handbooks for Identification of British Insect, Vol. 5, London, Royal
Entomological Society of London, 144 p.
RIVALLAIN, J., AND VAN NEER, W. 1983. Les fouille de Koyom (Sud du Tchad). Etude du
matériel archéologique et faunique. L’Anthropologie, Vol. 87, p. 221-239.
ROBERTS, E., ROGERS, R., AND FOREMAN, B.Z. 2003. An experimental approach to
identifying and interpreting dermestid (Insecta, Coleoptera) bone modification. Journal
of Vertebrate Paleontology, Abstracts with programs, Vol. 23, No. 3, p. 89A.
ROGERS, R. R., 1992. Non-marine borings in dinosaur bones from the Upper Cretaceous Two
Medicine Formation, northwestern Montana. Journal of Vertebrate Paleontology Vol. 12,
No. 4., p. 528-531.
29
SAMOUR, J. H., AND NALDO, J. L. 2003. Infestation of Dermestes carnivorus in a saker
falcon (Falco cherrug). The Veterinary Record Vol. 153, p. 658-659.
SCHROEDER, H., KLOTZBACH, H., OESTERHELWEG, L. L, AND PÜSCHEL, K. 2002.
Larder beetles (Coleoptera, Dermestidae) as an accelerating factor for decomposition of a
human corpse. Forensic Science International, Vol. 127, p. 231-236.
SMITH, K. G. V. 1986. A manual of forensic entomology. Cornell University Press, Ithica,
New York, 205 p.
SNYDER, N. F. R., OGDEN, J. C., BITTNER, J. D., AND GRAU, G. A. 1984. Larval
dermestid beetles feeding on nestling snail kites, wood storks, and great blue herons. The
Condor Vol. 86, p. 170-174.
SOMMER, H. G., AND ANDERSON, S. 1974. Cleaning skeletons with dermestid beetles: two
refinements in the method. Curator, Vol. 17, p. 290-298.
SU, N.-Y., AND SCHEFFRAHN, R .H. 1988. Foraging population and territory of the
Formosan subterranean termite (Isoptera:Rhinotermitidae) in an urban environment.
Sociobiology, Vol. 14, p. 353–359.
TAPPEN, M. 1994. Bone weathering in the tropical rain forest. Journal of Archaeological
Science, Vol. 21, p. 667-673.
THORNE, B. L., AND KIMSEY, R .B. 1983. Attraction of neotropical Nasutitermes termites
to carrion. Biotropica, Vol. 15, No. 4, p. 295-296.
THORNE, B. L., GRIMALDI, D. A., AND KRISHNA, K. 2000. Early fossil history of the
termites, p. 77-93. In T. Abe, D. E. Bignell, and M. Higashi (eds.), Termites: Evolution,
Sociality, Symbiosies, Ecology. Kluwer Academic Publishers, Dordrecht, Netherlands.
TIMM, R. M. 1982. Dermestids. Field Museum of Natural History Bulletin, Vol. 53, p. 14-18.
30
TOBIEN, H. 1965. Insekten-frassspuren an Tertiären und Pleistozänen Säugertierknochen.
Senckenbergiana Lethaia, Vol. 46a, p. 441-451.
TRANIELLO, J .F. A., AND LEUTHOLD, R .H. 2000. Behavior and ecology of foraging in
termites, p. 141-168. In T. Abe, D. E. Bignell, and M. Higashi (eds.), Termites:
Evolution, Sociality, Symbiosies, Ecology. Kluwer Academic Publishers, Dordrecht,
Netherlands.
WALLER, D. A., AND LA FAGE, J. P. 1987a. Nutritional ecology of termites, p. 487-532. In
F. Slansky Jr., and J.G. Rodriguez (eds.), Nutritional Ecology of Insects, Mites, and
Spiders. Wiley & Sons, New York.
WALLER, D .A., AND LA FAGE, J. P. 1987b. Food quality and foraging response by the
subterranean termite Copotermes formosanus Shiraki (Isoptera: Rhinotermitidae).
Bulletin of Entomological Research, Vol. 77, p. 417-424.
WATSON, J. A. L., AND ABBY, H. M. 1986. The effects of termites (Isoptera) on bone: some
archaeological implications. Sociobiology, Vol. 11, p. 245-254.
WEST, D. L., AND MARTIN, L. D. 1997. In Proceedings of the 1993 bone modification
conference, Hot Springs, South Dakota, edited by L.A. Hannus, L. Rossum, and R.P.
Winham, p. 88-97. Occasional Publication No. 1, Archaeology Laboratory. Augustana
College, Sioux Falls, South Dakota.
WYLIE, F. R., WALSH, G. L., AND YULE, R. A. 1987. Insect damage to aboriginal relics at
burial and rock-art sites near Carnavon in central Queensland. Journal of the Australian
Entomological Society, Vol. 26, p. 335-345.
WOOD, W. B. 1976. The skeletal material from the Brooloo Range and Rocky Hole Creek
burial sites. Archaeology and Physical Anthropology in Oceania, Vol. 11, No. 3, p. 175-
31
185.
32
FIGURE CAPTIONS
FIGURE 1—Pits on dinosaur bone from the Morrison Formation. 1. Fine, shallow pits on
Allosaurus right ischium, MOR 693. 2. Pits on Allosaurus left fibula, BYU 5129. 3. Detail of
caudomedial surface, BYU 5129. 4. Detail of distomedial surface, BYU 5129. 5. Pits on
superior surface of Torvosaurus right metatarsal, BYU 5277. 6. Detail of BYU 5277. 7. Deep
pits on Allosaurus left scapula, BYU 13652. 8. Detail of BYU 13652. Unlabelled scales are 1cm.
FIGURE 2—Pits on fossil dinosaur and modern elephant bone. Fossils are from the Morrison
Formation; elephant bone is from the Ituri Rain Forest, Zaire. 1. Pit trail, Torvosaurus chevron,
BYU 4676a. 2. Linear pit trail made by termites (From Tappen, 1994, courtesy Academic
Press). 3. Deep pitting with loss of all surficial bone, sauropod bone fragment, BYU 17946. 4.
Deep pitting, loss of all surficial bone, underside of elephant mandible, UMN ZA-91-1. 5.
Shallow pits bordered by intact surface, elephant scapula, UMN ZA-91-2. 6. Pit trails and
shallow pit clusters, Torvosaurus scapula, BYU 4900. 7. Detail of trail and cluster on BYU
4900. 8. Pits of varying sizes and fine pit trails, Torvosaurus chevron, BYU 4976c. 9. Linear
pit trails following ligament attachment ridges, Torvosaurus chevron, BYU 4976a. Unlabelled
scales are 1cm.
FIGURE 3—Ring pits on sauropod bones from the Morrison Formation and schema of ring
development. 1. Ring furrows, diplodocid rib, SMM XXXX. 2. Complex rings, matrix-filled,
sauropod rib, SMM XXXX. 3. Matrix-filled concentric rings, diplodocid pubis. 4. Cross section
of laminar bone prior to ring development. 5. Early stage of ring development with central table.
33
5. Late stage of ring development with partially consumed table. 6. Pit formed by destruction of
central table. Scales = 1cm
FIGURE 4—In-filled burrows. 1. Gray burrows over pits on diplodocid scapula, MOR 957,
Morrison Formation. Most burrows covering pits were removed. 2. Line drawing of MOR 957,
burrows are white.
34
TABLES
TABLE 1—Quarries in the Brushy Basin Member of the Morrison Formation examined for
pitted bones. See abbreviations section for explanations for institutions. Quarry location data
are on file with each institution. [Double Column]
Pit
Institution
BYU
Quarry
Accumulation
Pitted?
Abundance
Matrix
Dry Mesa
bone bed
1
common, 16%
sandstone
Kalico Gulch
bone bed
1
common
sandstone
Cactus Park
bone bed
1
rare
mudstone
Hinkle
1 individual
0
none
sandstone
Agate Basin
1 individual
0
none
siltstone
Jensen-Jensen
bone bed
0
none
sandstone
Jones
1 individual
0
none
mudstone
Easter/CAD
2 individuals
0
none
limestone
Dominguez-
abundant,
DNM
Carnegie
bone bed
1
40%
sandstone
MOR
Big Al (693)
1 individual
1
common, 13%
sandstone
BHTL
1 individual
1
abundant
sandstone
714
bone bed
1
rare
?
790
bone bed
1
rare
?
592
bone bed
1
rare
?
957
bone bed
1
rare
?
Mygatt-Moore
bone bed
0
absent
mudstone
MWC
35
NAMAL
Bone Cabin
many
West
individuals
1
common
sandstone
Meilyn
3 individuals
1
common
sandstone
Stego 99
bone bed
0
none
sandstone
abundant,
SMM
Poison Creek
bone bed
1
49%
siltstone
bone bed
1
rare
mudstone
ClevelandUU
Lloyd
Quarries with
pitted bones
14
Quarries
examined
20
% of quarries
with pits
70
36
TABLE 2—Comparison of traces made by moths, dermestid beetles, and termites with pits on
Morrison Formation bones. [Format to Double Column]
Character
Tineid Moth
Dermestid Beetle
Termite
Morrison Pits
Pit diameter
undocumented
3.5-5.2 mm
<1-13mm
0.15-20mm
Pit sizes on
minor variation
minor variation
variety of sizes
variety of sizes
furrows, ellipses
round to elliptical
round to
round to elliptical.
elliptical,
primarily round
single bone
Shape, plan
view
primarily round
Shape, x-
U-shaped
section
narrow, deep shaft to U-
U-shaped
U-shaped
shaped
Pit depth
undocumented
few to 15 mm
few mm
shallow, max 5mm
Pit patterns
none
none to single row
random, to wide
random, to wide
lineations
lineations
lineations, rings
none
none
0 to cm scale
0 to cm scale
no
no
yes
yes
yes
yes
no
no
no
no
yes
yes
Depth of
surface loss
Composite
pits?
burrows in
bone
Associated
large
burrows in
substrate
37
TABLE 3—Comparison of dermestid and termite habits. [Format to Single Column]
Character
Dermestids
Termites
Amount of flesh
moderate to high
minimal to none
Age of carcass
days-months
days to years
Carcass exposure
subaerial
buried to subaerial
Feeding surface
sheltered upper surface
underside, exposed surfaces
Temperature range
tropical to temperate
tropical to subtropical
Humidity range
moderate
low to high
Full infestation
3 days to weeks
immediate (colony invades)
38
Running Title: Jurassic dinosaurs and termites
Keywords: Termites, Isoptera, Dermestidae, Dinosauria, Late Jurassic, paleoecology,
taphonomy
TABLE 1—Quarries in the Brushy Basin Member of the Morrison Formation
examined for pitted bones. See abbreviations section for institutional names.
Institution
BYU
DNM
MOR
MWC
NAMAL
SMM
UUVP
Quarry
Accumulation Pitted?
Dry Mesa
bone bed
1
Kalico Gulch
bone bed
1
Cactus Park
bone bed
1
Hinkle
1 individual
0
Agate Basin
1 individual
0
Jensen-Jensen
bone bed
0
Dominguez-Jones 1 individual
0
Easter/CAD
2 individuals
0
Carnegie
bone bed
1
Big Al (693)
1 individual
1
BHTL
1 individual
1
714
bone bed
1
790
bone bed
1
592
bone bed
1
957
bone bed
1
Mygatt-Moore
bone bed
0
Bone Cabin West many individuals
1
Meilyn
3 individuals
1
Stego 99
bone bed
0
Poison Creek
bone bed
1
Cleveland-Lloyd
bone bed
1
Quarries with pitted bones
14
Quarries examined
20
% of quarries with pits
70
Pit Abundance
common, 16%
common
rare
none
none
none
none
none
abundant, 40%
common, 13%
abundant
rare
rare
rare
rare
absent
common
common
none
abundant, 49%
rare
Matrix
sandstone
sandstone
mudstone
sandstone
siltstone
sandstone
mudstone
limestone
sandstone
sandstone
sandstone
?
?
?
?
mudstone
sandstone
sandstone
sandstone
sandstone
mudstone
Deposit
fluvial channel
fluvial channel
overbank
fluvial channel
overbank
fluvial
overbank
lacustrine
fluvial channel
fluvial channel
fluvial channel
?
?
?
?
fluvial-lacustrine
flood plain
fluvial
fluvial
fluvial channel
fluvial-lacustrine