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115
Hunt et al., eds., 2012, Vertebrate Coprolites. New Mexico Museum of Natural History and Science, Bulletin 57.
NEW COPROLITE ICHNOTAXA FROM THE BUCKLAND COLLECTION
AT THE OXFORD UNIVERSITY MUSEUM OF NATURAL HISTORY
ADRIAN P. HUNT1, SPENCER G. LUCAS2 AND JUSTIN A. SPIELMANN2
1 Flying Heritage Collection, 3407 109th St SW, Everett, WA 98204, e-mail: adrianhu@flyingheritage.com;
2 New Mexico Museum of Natural History and Science, 1801 Mountain Road NW, Albuquerque, NM 87104,
e-mail: spencer.lucas@state.nm.us; justin.spielmann1@state.nm.us
Abstract—The Buckland collection at the University of Oxford Museum of Natural History (UK) is the oldest
collection of coprolites in the world. It comprises Mesozoic and Cenozoic coprolites and other bromalites, the
majority of which are derived from the Lower Lias of southwestern England. The collection includes the most
comprehensive samples of two important British coprofaunas – the Lower Liassic of Lyme Regis and the Rhaetic
bone bed. We describe four new coprolite ichnotaxa from this collection: Ichthyosaurolites duffini ichnogen. et
ichnosp. nov., Strabelocoprus pollardi ichnogen. et ichnosp. nov. and Plektecoprus whitbyensis ichnogen. et
ichnosp. nov. from the Late Triassic and Early Liassic and Hyaenacoprus bucklandi ichnogen. et ichnosp. nov.
from the Late Pleistocene.
INTRODUCTION
William Buckland (1784-1856) was the first to study fossil feces
and coined the term “coprolite” (Buckland, 1822, 1824, 1829a-d, 1830,
1835, 1836; Duffin, 2006, 2009, 2012a-b; Hunt and Lucas, 2012a;
Pemberton, 2012). Buckland held academic positions at the University
of Oxford, first as Reader in Mineralogy and subsequently as Reader in
Geology (Duffin, 2006). He built up a collection of coprolites (the first
ever) through personal field work, purchases from the famous fossil
collector Mary Anning and fossil dealers and specimens donated from a
wide network of colleagues (Duffin, 2012a-b). His collection at the
University of Oxford Museum of Natural History is dominated by co-
prolite specimens from the Early Jurassic, but it also includes specimens
from the Late Triassic, Late Jurassic, Early Cretaceous and Late Pleis-
tocene and non-coprolite bromalites and infilled Recent shark intestines
(Figs. 1-5).
Duffin (1979; Swift and Duffin, 1999) first restudied coprolites
from the Rhaetic bone bed, from the Buckland and other collections, and
recognized four broad morphological types of coprolites. Hunt et al.
(2007) identified six morphotypes, including Liassocopros hawkinsi
and Saurocopros bucklandi.
Duffin (2010) identified seven morphotypes of coprolites in the
Buckland collection sample, and others, from the Lower Lias of the
coastal area of Dorset, England, including Falcatocoprus isp., Saurocoprus
bucklandi, Saurocoprus isp., Liassocoprus hawkinsi and three other
morphotypes. Other bromalites from the Buckland collection, notably
consumulites of ichthyosaurs, have been studied by several workers
(e.g., Pollard, 1968; Taylor, 1993). The purpose of this paper is to
describe four new ichnotaxa from the Buckland Collection of bromalites
at the University of Oxford Museum of Natural History, three from the
Triassic-Lias of Dorset and Somerset and one from the Pleistocene of
Yorkshire, all from the United Kingdom.
SYSTEMATIC PALEONTOLOGY
Ichthyosaurolites, ichnogen. nov.
Type ichnospecies: Ichthyosaurolites duffini Hunt et al., 2012.
Included ichnospecies: Known only from the type ichnospecies.
Etymology: From ichthyosaur, for the contents of the ichnofossil,
and the Greek lithos (rock).
Distribution: Lower Jurassic (Lower Liassic) of southwest En-
gland (Dorset).
Diagnosis: Bromalite that differs from other ichnogenera in con-
sisting of a wide, flattened rectangle with a rounded tip with abundant
phosphatic(?) groundmass and inclusions of multiple skeletal elements
of juvenile ichthyosaurs (vertebral diameters typically about 12-15 mm).
Discussion: This distinct morphology of this bromalite was first
recognized by Buckland (1836, pl. 15, fig. 18) and later by Duffin (2010,
pl. 77, fig. 1). Ichthyosaurolites represents a concentration of juvenile
bones, and thus it could potentially represent a gignolite (sensu Hunt and
Lucas, 2012a) or a bromalite. Putative ichthyosaur embryos are well
known (e. g., Böttcher, 1990, 1998; Deeming et al., 2001; Maxwell and
Caldwell, 2003) and Ichthyosaurolites specimens differ from these in
that: (1) the specimens are comprised principally of vertebrae in a disar-
ticulated mass in contrast to an articulated series; (2) no small non-
vertebral bones are preserved; and (3) the bones are contained in a dis-
tinct groundmass.
Ichthyosaurolites is not considered to represent a regurgitalite
because of the the large volume of groundmass relative to bone. The
discrete small volume of the specimens indicates that Ichthyosaurolites
does not represent an infilling of a portion of the gastrointestinal tract
and it is most parsimoniously considered a pelletized accumulation of
fecal material (incorporeal pelletite or coprolite of Hunt and Lucas, 2012a).
Thus, Ichthyosaurolites indicates active predation on juvenile ichthyo-
saurs. The size of the coprolites and the lack of spiral structure suggest
that the originator is a large marine reptile.
Vertebrate consumulites are most commonly preserved in aquatic
organisms, including ichthyosaurs (Fig. 2), because taphonomic factors
(e.g., water chemistry, deposition rates) in aqueous environments in-
crease the likelihood of the preservation of complete carcasses relative to
subaerial conditions. In addition, ichthyosaurs have long been considered
viviparous (Pearce, 1846). Thus, potentially a juvenile ichthyosaur skel-
eton within an adult one, as is known in about 50 instances (Wild, 1990),
could represent an embryo (e.g., Pearce, 1846; Seeley, 1880) or an act of
cannibalism (e. g., Quenstedt, 1858). There has been considerable discus-
sion of these hypotheses (e.g., Branca, 1908; Drevermann, 1926;
Liepmann, 1926) and it appears that both circumstances occur
(McGowan, 1991). Cannibalism by ichthyosaurs is a possibility for the
origin of Ichthyosaurolites, but the relative scarcity of these bromalites
(five specimens known from Lyme Regis) relative to the large number of
ichthyosaur specimens suggests that the predator was another taxon,
possibly a plesiosaur. O’Keefe et al. (2009) demonstrated ingestion of a
juvenile ichthyosaur (vertebral centra of 12-15 mm in diameter) by a
plesiosaur. They considered the ichthyosaur to be a voided embryo, but
the multiple specimens of Ichthyosaurolites suggest either that the size
of newborn ichthyosaurs has been overestimated or that spontaneous
abortions by ichthyosaurs were common.
116
FIGURE 1. Coprolites and a related specimen from the Buckland Collection at the Oxford University Museum of Natural History. A, GZ 103,
Coprolite from the Muschelkalk (Anisian-Ladinian), Lunéville, France. B, K 1272, Coprolite from Wiltshire, UK. The specimen label
indicates that this coprolite derives from the Cambridge Greensand (Aptian), however this unit does not cropout in that county (C. Duffin, pers.
commun., 2012). Thus, it is more likely that the specimen is from the Upper Greensand (Albian) or Lower Greensand (Aptian) if it is from
Wiltshire (C. Duffin, pers. commun., 2012). C, GZ 105, Liassocoprus isp. coprolite from the Keuper (Alaunschiefer)(Ladinian), near Gaildorf,
Germany. D, H 36, Coprolite from Rhaetic bonebed (Rhaetic), Aust Cliff, UK. E, J23883, Saurocopros bucklandi (Buckland, 1835, pl. 28, fig.
6) from the Lower Lias (Hettangian-Lower Pliensbachian) of Lyme Regis, UK. F, JZ 1701, Coprolite from Solnhofen Plattenhalk (Tithonian),
Solnhofen, Germany. G, GZ 104, Coprolite from Gaildorfer Keuper (Alaunschiefer)(Ladinian), near Gaildorf, Germany. H, R 4, Roman cement
infilled intestines of Recent dogfish (Squalus): Buckland, 1836, pl. 15, fig. 1. I, J23862-23865, Four coprolites on one block from the Lower
Lias (Hettangian-Lower Pliensbachian), Lyme Regis, UK.
117
FIGURE 2. Consumulites in ichthyosaur skeletons from the Buckland Collection at the Oxford University Museum of Natural History from the Lower Lias
(Hettangian-Lower Pliensbachian) of Lyme Regis, UK. A-B, OUM J12125, Consumulite in A, overview and B, close up (also see Pollard, 1968, pls. 72-
73). C, J OUM 12146, Consumulite in lateral view. D, OUM J10320, Consumulite in lateral view. E, OUM J13593, Consumulite in lateral view (also see
Buckland, 1836, pl. 14).
118
There are indications of Liassic ichthyosaur diet from the
Posidonienschiefer of Germany, for example, suggesting that prey selec-
tion changed through ontogeny (from fish to cephalopods) (Böttjer,
1989, 1998). There is also specific evidence that the Liassic
Temnodontosaurus trigodon preyed on Stenopterygius spp. (Böttjer,
1989, 1998), so it is also possible that Ichthyosaurolites represents the
coprolite of a large ichthyosaur such as Temnodontosaurus.
Ichthyosaurolites duffini, ichnosp. nov.
Holotype: OUM J 23905 (Fig. 3A-E; Buckland, 1836, pl. 15, fig.
18; Keller, 1977, fig. 6B; Duffin, 2010, pl. 77, fig. 1)
Etymology: For Christopher Duffin, to honor his contributions
to the study of coprolites and of the life and work of William Buckland.
Type locality: Lyme Regis, England.
Type horizon: Lower Liassic (Hettangian-Lower Pliensbachian).
Distribution: As for genus.
Referred specimens: OUM J 23922 (Figs. 3F-G), OUM J 23911
(Fig. 3M), OUM J23888a-b (Figs. 3J-L) and OUM J23771 (Figs. 3H-I),
all from the Lower Lias of Lyme Regis, England.
Diagnosis: As for genus.
Description: OUM J 23905 is 64 mm long and 39 mm wide with
a thickness of 24.4 mm. The only visible bones are many small ichthyo-
saur centra (six are visible on one surface), and they have widths ranging
from 14.1 to 15.4 mm (Figs. 3J-L). The matrix, which contains many
small bones, is eroded around the ichthyosaur elements, and they pro-
trude from the surface.
Discussion: There are four referred specimens (OUM J23888a-
b, OUM J 23922, OUM J 23911 and OUM J23771). OUM J23888a-b
consists of two parts that fit together, and they provide the best infor-
mation about the overall morphology of a complete coprolite (Figs. 3J-
L). OUM J23888a is a rectangular (99.3 mm by 84.2 mm) piece, and
OUM J23888b is narrower, subtriangular and has a rounded end. When
fitted together the coprolite is 143.2 mm long, with one side relatively
smooth and presumably approximating the original outer surface of the
coprolite, and the other very irregular (eroded?). The coprolite contains
several ichthyosaur vertebrae (10.6 to 14.6 mm in diameter) and other
angular bone fragments.
OUM J 23922 is a flattened ovoid, 73.3 mm long, 50.6 mm wide
and 29.4 mm thick (Figs. 3F-G). It contains several bone fragments and
vertebrae (diameters of 7.01, 10.8, 11.8 and 14.5 mm). This specimen
was mentioned by Keller (1977, p. 131, fig. 6b), who noted that it
“contains a sphenoid and a basioccipital bone of one individual. From the
size of these bones the body length of the swallowed and digested prey
can be estimated at 60-70 cm” (translation courtesy of L. H. Vallon).
OUM J 23911 (57.1 mm by 71.2 mm) is preserved on a sheet of matrix
(57.1 mm by 71.2 mm) and seems flattened and lacks the distinct margin
seen in the other specimens. This specimen contains angular bone frag-
ments and several vertebrae (diameters of 15.2, 15.6 and 13.6 mm).
OUM J23771 is a coprolite with one surface (51.6 by 50.7 mm ) ground
and polished and exhibiting bone fragments that are larger than in the
other specimens (Fig. 3H-I). The obverse side is irregular, with many
bone fragments. There are no obvious vertebrae in this specimen.
Strabelocoprus, ichnogen. nov.
Type ichnospecies: Strabelocoprus pollardi Hunt et al., 2012.
Included ichnospecies: Known only from the type ichnospecies.
Etymology: From the Greek strabelos (snail) in allusion to the
similarity in shape to a gastropod, and the Greek kopros (feces).
Distribution: Rhaetian Penarth Group (see discussion below)
and Lower Jurassic (Lower Liassic) of southwest England (Dorset and
Somerset).
Diagnosis: Heteropolar, microspiral coprolite that differs from
other ichnogenera in having a small number of coils (<3) in lateral view,
exhibiting very wide spirals in posterior view and in having a width that
exceeds half of its length.
Discussion: This ichnogenus is named for its gastropod-like mor-
phology. It is currently only known from the Upper Triassic and Lower
Lias of southwestern England. The spiral morphology suggests that the
coprolite was produced by a less derived fish such as a chondricthyan
that must have been of very large size. What clearly distinguishes
Strabelocoprus from other large microspiral copolites (e.g.,
Megaheteropolacoprus) is the large width to length ratio.
Strabelocoprus pollardi, ichnosp. nov.
Holotype: OUM J23743, coprolite (Fig. 4A-D).
Etymology: For John Pollard, to honor his contributions to
ichnology, including coprolite studies, in the UK.
Type locality: Watchet, Somerset, England.
Type horizon: Rhaetian Penarth Group (see discussion below).
Distribution: As for ichnogenus.
Referred specimens: OUM J23741, Lower Liassic (Hettangian-
Lower Pliensbachian) of Lyme Regis, England (Fig. 4E).
Diagnosis: As for ichnogenus.
Description: OUM J23743 is a complete phosphatic coprolite
with a length of 102.2 mm and a subcircular cross section with a maxi-
mum width of 62 mm and a lesser width of 58.3 mm (Figs. 4A-D). The
coprolite is heteropolar and microspiral with three coils and a relatively
elongate posterior spire (sensu Hunt et al., 2007; Hunt and Lucas, 2012b).
Fish scales are visible in several areas and are prominent on the lip of the
posterior spire. Adherent bilalves occur on one side of the coprolite,
Discussion: The holotype was collected in 1840, and its prov-
enance is listed as Lias. However, Duffin (pers. commun., 2012) has
raised legitimate issues with this putative stratigraphic derivation of the
holotype of Strabelocoprus pollardi. The stratigraphic sequence at
Watchet includes both the Lower Lias and the Rhetian Penarth Group.
The adherent bivalves on the holotype are Atreta intusstriata (Duffin,
pers. commun., 2012), which is most common in the Rhaetian Lilstock
Formation (upper Penarth Group: Swift, 1999), although it rarely occurs
in the Westbury Formation (lower Penarth Group) and also in the Lias
(Ivimey-Cook et al., 1999, p. 98, pl. 13, figs. 3-4). Atreta intusstriata in
the Lilstock Formation is “often attached to hard substrates” (Ivimey-
Cook et al., 1999, p. 98).
The referred specimen of Strabelocoprus pollardi (OUM J23741),
from the Lias of Lyme Regis, is flattened, but preserves the same overall
morphology and is of similar dimensions as the holotype, with a length
of 107.6 mm and a width of 73.4 mm (Fig. 4E). This specimen exhibits
some decay around the margins.
Hunt and Lucas (2010) noted that coprolites are commonly pre-
served in hydrodynamically-accumulated intraformational conglomer-
ates and bone beds in marine and nonmarine environments (e. g., Martill,
1999, fig. 6; Hunt and Lucas, 2010, fig. 2; Hunt et al., 2012, fig. 2F).
There is little actualistic information about the decay of feces, but the
excrement of the Recent lungfish Neoceratodus forsteri and Protopterus
annectans remains intact for several hours in an aqueous environment
(Jain, 1983). Thus, it is possible that the coprolites preserved in bone
beds could represent recently excreted feces. However, it seems more
parsimonious that coprolites preserved in bone beds represent previ-
ously-lithified feces (cf. Reif, 1971). The Atreta intusstriata attached to
one side of the holotype of Strabelocoprus pollardi indicate that this
specimen represents a lithified coprolite that lay on a sediment surface
before being incorporated in a sediment layer. Thus, OUM J23743 is a
taphonomically important specimen because it demonstrates that the
hypothetical circumstance of feces becoming lithified and subsequently
incorporated into a younger stratigraphic unit can in fact occur.
Plektecoprus, ichnogen. nov.
Type ichnospecies: Plektecoprus whitbyensis Hunt et al., 2012.
Included ichnospecies: Known only from the type ichnospecies.
Etymology: From the Greek plekte (rope) in reference to the
shape, and the Greek kopros (feces).
119
FIGURE 3. Ichthyosaurolites duffini ichnogen. et ichnosp. nov. from the Lower Lias (Hettangian-Lower Pliensbachian) of Lyme Regis, UK. A-E, OUM J
23905, Holotype coprolite in lateral (A-D) and terminal (E) views. F-G, OUM J 23922, Coprolite in lateral views. H-I, OUM J23771, Coprolite in lateral
views. J-L, OUM J23888a-b, Coprolite in lateral views. M, OUM J 23911, Coprolite in lateral view.
120
FIGURE 4. Strabelocopros pollardi ichnogen. et ichnosp. nov. and Plektecoprus whitbyensis ichnogen. et ichnosp. nov. (Hettangian-Lower Pliensbachian).
A-D, Strabelocoprus pollardi ichnogen. et ichnosp. nov., OUM J23743, Holotype in A-B, lateral, C, posterior and D, anterior views. E, OUM J23741,
Referred specimen, coprolite in matrix block. E-F, Plektecoprus whitbyensis ichnogen. et ichnosp. nov., OUM J29985, Holotype in two axial views.
Distribution: Lower Jurassic (Lower Liassic) of Yorkshire, En-
gland.
Diagnosis: Coprolite that differs from other ichnogenera in being
elongate, rounded in cross section with a loose spiral coil and having a
conical posterior end and a broad, rounded anterior end.
Discussion: This ichnogenus is currently only known from the
Lower Lias of Yorkshire. The producer of this coprolite could be a
marine reptile, given its size and lack of a tight spiral morphology.
Plektecoprus whitbyensis, ichnosp. nov.
Holotype: OUM J29985, coprolite (Fig. 4F-G).
Etymology: For the town of Whitby, the type locality.
Type locality: Whitby, Yorkshire, England.
Type horizon: Lower Liassic (Hettangian-Lower Pliensbachian).
Distribution: As for ichnogenus.
Referred specimens: None.
Diagnosis: As for ichnogenus.
Description: OUM J29985 is preserved in semi-relief on an ovoid
sheet of shale (Figs. 4F-G). The coprolite is complete, with a maximum
length of 69.3 mm with a loose spiral coil. The anterior end is conical, and
the posterior end is rounded.
Discussion: The elongate shape of this ichnospecies and its loose
coil may have made it susceptible to mechanic destruction, so Plektecoprus
may be rarely preserved.
121
FIGURE 5. Hyaenacoprus bucklandi ichnogen. et ichnosp. nov. from the Late Pleistocene of Kirkdale Cave, Yorkshire, UK. A-F,
OUM Q6168, Holotype partial coprolite in A-D, axial and E-F, polar views. G-J, Q6167, Partial coprolite in G-H, axial and I-J, polar
views.
122
Hyaenacoprus, ichnogen. nov.
Album graecum: Buckland, 1822, p. 186, pl. 24, fig. 6
Album graecum: Buckland, 1824, p. 20, pl. 10, fig. 6
Hyaino-coprus: Buckland, 1829a, p. 143
Hyaena-coprus: Buckland, 1830, p. 24
Type ichnospecies: Hyaenacoprus bucklandi ichnosp. nov.
Included ichnospecies: Known only from the type ichnospecies.
Etymology: Based on the term employed by Buckland (1830, p.
24), suggesting that these are hyaena coprolites.
Distribution: Upper Pliocene-Pleistocene of Europe, Asia and
Africa.
Diagnosis: Phosphatic coprolite that differs from other ichnogenera
in being composed of a series of rounded segments (pellets of Diedrich,
2012), some of which are sub-spherical and are white in color with many
small angular bone fragments.
Discussion: Buckland (1822, 1824) described the geology and
paleontology of Kirkdale Cave in Yorkshire, which he interpreted to be a
Late Pleistocene hyena den. He discovered “many small balls of the solid
calcareous excrement of an animal that had fed on bones, resembling the
substance known in the old Materia Medica by the name of album
graecum …. its external form is that of a sphere, irregularly compressed,
as in the feces of sheep, and varying from half an inch to an inch in
diameter; its colour is yellowish white, its fracture is usually earthy and
compact, resembling steatite, and some-times granular; when compact, it
is interspersed with minute cellular cavities: it was at first sight recognised
by the keeper of the Menagerie at Exeter Change, as resembling, both in
form and appearance, the faeces of the spotted or Cape Hyaena, which
he stated to be greedy of bones, beyond all other beasts under his care.
This information I owe to Dr. WOLLASTON, who has also made an
analysis of the substance under discussion, and finds it to be composed
of the ingredients that might be expected in faecal matter derived from
bones, viz. phosphate of lime, carbonate of lime, and a very small pro-
portion of the triple phosphate of ammonia and magnesia; it retains no
animal matter, and its originally earthy nature and affinity to bone, will
account for its perfect state of preservation” (Buckland, 1822, p. 186-
187). Subsequently, Buckland was able to conduct actualistic studies to
confirm his hypothesis: “I have had an opportunity of seeing a Cape
Hyaena at Oxford, in the travelling collection of Mr. Wombwell, the
keeper of which confirmed in every particular the evidence given to Dr.
Wollaston by the keeper at Exeter ‘Change……The keeper pursuing this
experiment to its final result [the feeding of bones to the hyena], pre-
sented me the next morning with a large quantity of album graecum,
disposed in balls, that agree entirely in size, shape, and substance with
those found in the den at Kirkdale” (Buckland, 1824, p. 38). Buckland
illustrated one specimen of album graecum (Buckland, 1822, pl. 24, fig.
6; 1824, pl. 10, fig. 6). This is a term that was used by apothecaries to
refer to dog feces that were especially rich in phosphate as a result of
feeding a bone-rich diet to dogs (Duffin, 2009). (Buckland (1829a, p.
143) later applied the term “Hyaino-coprus” to “the Album Graecum of
the fossil hyena” (Hyaena-coprus in Buckland, 1830, p. 24). We use the
latter name for the new ichnogenus to honor Buckland.
Hyena excrement is robust due to early diagenesis and cementa-
tion of the bone phosphate that commences in the intestines, and they
can survive hydrodynamic transport (Diedrich, 2012). Trampled latrinites
(sensu Hunt and Lucas, 2012a) occur in some European caves such as in
the Lindenthaler Hyänenhöhle in Germany (Liebe, 1876; Diedrich, 2012).
Hyaenacoprus bucklandi is widespread in Late Pleistocene caves
in the Old World. There is clearly an ichnofacies in North American caves
distinct from those in Europe, Asia and Africa. Old World caves are
dominated by hyena coprolites (e.g., Scott, 1987; Pesquero et al., 2011;
Diedrich, 2012), whereas those in North America are dominated by di-
verse herbivore coprolites (e. g., Mead and Agenbroad, 1989; Hunt and
Lucas, 2007; Mead and Swift, 2012).
Hyaenacoprus bucklandi, ichnosp. nov.
Holotype: OUM Q6168, partial coprolite (Figs. 5A-F).
Etymology: For the collector of the holotype, William Buckland,
to honor his contributions to the study of coprolites.
Type locality: Kirkdale Cave, Yorkshire, England.
Type horizon: Late Pleistocene cave fill.
Distribution: As for ichnogenus.
Referred specimens: OUM Q6167, partial coprolite (Figs. 5G-
J).
Diagnosis: As for ichnogenus.
Description: OUM Q6168 is a white coprolite fragment com-
posed of three principal, generally-rounded segments or pellets (sensu
Diedrich, 2012), the middle of which is sub-spherical on one side (Figs.
5A-F). These pellets are within the types e-f (irregular to round) of
Diedrich (2012). One side of the coprolite is generally flat and irregularly
pitted. The total length of the fragment is 55.4 mm, with a width of 57.4
mm. There are several small angular shards of bone visible in the copro-
lite.
Discussion: Referred specimen OUM Q6167 is a white ovoid
pellet with a nearly round cross section (Figs. 5G-J). It is 51.4 mm long
with widths of 31.4 and 32.6 mm. Each rounded end has a small broken
attachment for an adjoining pellet. This specimen corresponds to a type
d (long oval) pellet of Diedrich (2012).
We believe that Buckland was correct in hypothesizing that
Hyaenacoprus is the product of a hyena. However, in Africa today both
the lion and hyena produce feces of broadly similar morphology that are
tapering, segmented cylinders (Stuart and Stuart, 2000). Fresh hyena
feces are greenish in color, but they whiten when dry because of the high
bone content. Lion feces are usually dark in color, but they can also be
white if the diet is high in bone (Stuart and Stuart, 2000, p. 161 unnum-
bered fig. on lower left). Lion feces are usually of a larger size than those
of hyenas, typically over 4 cm in diameter, although this is not always
the case (Stuart and Stuart, 2000). Recent and fossil hyena feces/copro-
lites are usually concentrated at latrine/latrinite sites (Stuart and Stuart,
2000; Diedrich, 2012). Both hyenas (Crocuta crocuta spelaea) and lions
(Panthera leo spelaea, Panthera leo fossilis) occupied caves and other
sites in Pleistocene Europe and there is potential for confusion in identi-
fying their coprolites. Cave lions were larger than Recent subspecies and
so we hypothesize that their coprolites could be distinguished from
those of hyenas by larger size (greater than 4 cm in diameter) and isolated
occurrence. There is need for more study of Pleistocene coprolites in
Europe to test this hypothesis.
CONCLUSIONS
The Buckland Collection at the University of Oxford Museum of
Natural History is not only the oldest collection of coprolites but also
one of the most important. There is need for more description of this
collection, which includes the most comprehensive samples of two im-
portant British coprofaunas – the Lower Liassic of Lyme Regis and the
Rhaetic bone bed.
ACKNOWLEDGMENTS
We thank Paul Jeffery, Assistant Curator of the Geological Collec-
tions, Oxford University Museum of Natural History, for access to
collections. Chris Duffin and L. H. Vallon not only wrote insightful
reviews but also provided additional references and translations.
123
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