88Lucas, S.G. and Spielmann, J.A., eds., 2007, The Global Triassic. New Mexico Museum of Natural History and Science Bulletin 41.

A REVIEW OF VERTEBRATE COPROLITES OF THE TRIASSICWITH DESCRIPTIONS OF NEW MESOZOIC ICHNOTAXA

ADRIAN P. HUNT, SPENCER G. LUCAS, JUSTIN A. SPIELMANN AND ALLAN J LERNER

New Mexico Museum of Natural History, 1801 Mountain Road NW, Albuquerque, NM 87104-1375

Abstract—Coprolites are the least studied and most under-sampled vertebrate trace fossils. They are verycommon in some Triassic localities. We recognize six new coprolite ichnotaxa: Alococopros triassicus, A. indicus,Saurocopros bucklandi, Liassococopros hawkinsi, Malericopros matleyi and Falcatocopros oxfordensis. Thedistribution of coprolite ichnotaxa is: Permian - Hyronocopros amphipolar and Heteropolacopros texaniensis;Early Triassic - Hyronocopros amphipolar and Alococopros triassicus; Middle Triassic - Alococopros triassicusand ?Liassocopros sp.; Late Triassic - Heteropolacopros texaniensis, Alococopros triassicus, Dicynodontocoprosmaximus, Malericopros matleyi, Liassocopros hawkinsi and Saurocopros bucklandi; Early Jurassic- Liassocoproshawkinsi and Saurocoporos bucklandi. We recognize the Liassocopros and Heteropolacopros coprolite ichnofacies.

INTRODUCTION

Coprolites are the least studied and most under-sampled verte-brate trace fossils. When we started extensively collecting Triassic verte-brate fossil assemblages in the early 1980s, we were struck by the preva-lence of vertebrate coprolites at many localities but their virtual absencein museum collections. Other paleontologists noted that they didn’tcollect these coprolites or that they subsequently disposed of themrather than accession them. A similar lack of attention (or respect) befellhuman coprolites in archeological sites (Bryant and Dean, 2006). Wehave strived to sample vertebrate coprolites as assiduously as otherfossils, and thus the New Mexico Museum of Natural History andScience now has the largest collection of Triassic vertebrate coprolites(Appendix).

There is an acme for vertebrate coprolites in Permian-Triassicredbeds (Hunt and Lucas, 2005b) with a worldwide distribution of Tri-assic assemblages (Fig. 1). Buckland (1829, p. 227), the founder of thestudy of coprolites (which we term paleoscatology), first described themin detail from the Rhaetian Westbury Formation of Great Britain(Buckland and Conybeare, 1822, p. 302, pl. 37 had earlier noted thembut not recognized them as coprolites): “some similar substances whichhave long been known to exist at Westbury, Aust Passage, and Watchet,on the banks of the Severn, and which now also prove to be faecal ballsof digested bone: they mostly occur in a thin bed of sandy micaeous lias,so full of bones and teeth and spines of reptiles and fishes, as to form abony breccia known to geologists by the name of bone-bed, and occupy-ing the lowest place at the bottom of the lias.” Subsequently, much laterin the twentieth century, there were several published studies of Triassiccoprolites (e.g., Rusconi, 1947, 1949, Ochev, 1974; Jain, 1983). Re-cently, there have been more detailed studies of the ichnotaxonomy,ichnofacies and biostratigraphy of Triassic vertebrate coprolites (e.g.,Lucas et al., 1985, Hunt et al., 1993, 1994, 1998; Northwood, 2005).However, these studies were very preliminary. The purpose of thispaper is to provide a first review of Triassic coprolites and provide astimulus for future work. In the course of this work, we recognized theneed to describe several new coprolite ichnotaxa from the Triassic, Juras-sic and Cretaceous.

Institutional abbreviations: BCM, Bristol City Museum andArt Gallery, Bristol; BMMNH, Natural History Museum (formerlyBritish Museum of Natural History), London; GSI, Geological Surveyof India, Calcutta; ISI, Indian Statistical Institute, Calcutta; MNA, Mu-seum of Northern Arizona in Flagstaff; NMMNH, New Mexico Mu-seum of Natural History and Science, Albuquerque; NMW, NationalMuseum of Wales, Cardiff; UCMP, University of California Museum ofPaleontology, Berkeley; UMMP, University of Michigan Museum ofPaleontology, Ann Arbor; YPM PU, Princeton collection at the YalePeabody Museum, New Haven.

SYSTEMATIC ICHNOLOGY

Introduction

Currently, there are only two named ichnogenera of Triassic co-prolites. Hunt et al. (1998) named Heteropolacopros texaniensis for aheteropolar-coiled coprolite and Dicynodontocopros maximus for largecoprolites presumed to have been produced by dicynodonts (Fig. 2).

In the course of our review of Triassic coprolites, we have notedthe need to formalize a number of distinct ichnotaxa. These include aform that is currently only known from the Jurassic but that we expectto be present in the Triassic, and an ichnogenus that has two species, oneof which is Cretaceous in age.

Alococopros igen. nov.

Type species: Alococopros triassicus isp. nov.Included species: A. triassicus and A. indicus.Etymology: From the Greek alocos for “furrowed” and kopros

for “dung.”Distribution: Early Triassic–Late Cretaceous of Australia, India

and North America.Diagnosis: Differs from other copolite ichnogenera in often being

arcuate in lateral view and sub-rounded in cross-section with regularlyspaced, thin, longitudinal grooves.

Discussion: Specimens here ascribed to this distinctive ichnogenuswere first described from the Upper Triassic of West Texas (Case, 1922,figs. 33C-D). It may be possible to distinguish between thinner, straighterforms (e.g., Northwood, 2005, fig. 2F) and broader, more arcuate forms(e.g., Case, 1922, figs. 33C-D).

Alococopros triassicus isp. nov.

Holotype: UMMP 7253 (partim), coprolite (Fig. 3A).Type locality: Crosby County, Texas.Type horizon: Tecovas Formation.Etymology: Named for the Triassic Period, which yields all known

specimens of this species.Distribution: Early-Late Triassic of Australia, India and North

America.Referred specimens: UMMP 7253 (partim), coprolite (Fig. 3B).Diagnosis: Differs from A. indicus in being less than one-fourth

as long (typically 2 cm in length).Discussion: This ichnospecies is currently only known from the

Triassic. Northwood (2005) discussed the origin of these types of co-prolites at length. Longitudinal intestinal rugae occur in both amphibiansand reptiles, but Northwood (2005) argued that Alococopros triassicus(her “longitudinally striated coprolites”) represent archosauromorphs,because: (1) this ichnotaxon first occurs in the Early Triassic; (2) some

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extant reptiles have longitudinal rugae; and (3) they resemble extantcrocodile feces (Young, 1964). This is a reasonable hypothesis, since thisichnospecies is restricted to the Triassic, as are basal archosauromorphs.

Alococopros indicus isp. nov.

Holotype: BMNH, unnumbered Matley Collection, two sectionsof the same coprolite (Fig. 3C-D).

Type locality: North of Kadubana, India.Type horizon: Lameta Formation.Etymology: Named for the country of India from which the type

specimens originate.Distribution: Upper Cretaceous of India.Referred specimens: Unnumbered coprolites, Matley Collec-

tion (Matley, 1939b, pls. 74, 75, figs. 1-4).Diagnosis: Ichnospecies that differs from A. triassicus in being

more than four times as long (typically 10 cm in length).Discussion: This ichnospecies is currently only known from the

Lameta Formation of India. This ichnospecies is considerably larger thanA. triassicus.

Saurocopros igen. nov.

Type species: Saurocopros bucklandi isp. nov.Included species: Known only from the type ichnospecies.Etymology: From the Greek sauros for “reptile” and kopros for

“dung” to honor Buckland’s (1829, p. 227, caption for plate 28) use ofthe term “sauro-coprolites” for specimens of this ichnotaxon.

Distribution: Late Triassic-Late Cretaceous of Europe and NorthAmerica.

Diagnosis: Microspiral heteropolar coprolite that differs from

Malericopros in being tapered below the spiral demarcation and fromHeteropolacopros in having a small number of wide spirals (typically 3)at the anterior end (compare Fig. 2A-I and Fig. 4).

Discussion: We name this ichnogenus with the full knowledgethat this coprolite does not pertain to a reptile. Rather, we name it tohonor William Buckland, who used the term “Sauro-coprolites” to referto coprolites of this morphology from the lower Lias of Lyme Regis (e.g.,Buckland, 1829, p. 227, pl. 28, figs. 6, 7, 9). These coprolites are abun-dant in the Lower Jurassic of England (e.g., Buckland, 1829; Hawkins,1834, 1840). Hunt and Lucas (2005c) described large heteropolar copro-lites from the Lower Permian of Texas. These specimens may pertain toSaurocopros.

Saurocopros bucklandi isp. nov.

Holotype: BMMNH R. 2102 (Fig. 4B: Hawkins, 1840, pl. 29).Type locality: Lyme Regis, England.Type horizon: Lower “Lias.”Etymology: Named for the Rev. William Buckland, who first

described specimens of this ichnogenus.Distribution: As for the ichnogenus.Referred specimens: BMNH R. 41285 (Fig. 4A), BMNH R.

1402 (Fig. 4C-D) and other coprolites from the Lower Lias of LymeRegis, England (Fig. 4E-G).

Diagnosis: As for the ichnogenus.Discussion: For obvious reasons, it is only appropriate to name

coprolites after scholars who have made contributions to paleoscatologyand who would presumably consider the attribution an honor. Such is thecase with William Buckland, who not only coined the term “coprolite”but who also founded and pursued the field of paleoscatology.

FIGURE 1. Distribution of principal Triassic coprolite-rich areas on Triassic Pangea. Locations are: 1, Queensland, Australia (Early Triassic); 2, Pranhita-Godavari basin, India (Middle-Late Triassic); 3, Mendoza region, Argentina (Middle Triassic); 4, Chinle and Moenkopi basins, United States (Early-LateTriassic); 5, Newark Supergroup basins, United States and Canada (Late Triassic); 6, United Kingdom and Germany (Middle-Late Triassic); 7, Kazakhstanand Russia (Middle Triassic). Base map after Wing and Sues (1992).

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Liassocopros igen. nov.

Type species: Liassocopros hawkinsi isp. nov.Included species: Known only from the type species.Etymology: From the Liassic of England, which has yielded the

first and most numerous specimens of this ichnogenus.

Distribution: Late Triassic-Late Cretaceous of Europe, India andNorth America.

Diagnosis: Coprolite that differs from most ichnotaxa in beingheteropolar and that differs from Heteropolacopros and Saurocopros inbeing macrospiral (see definition below) in morphology.

Discussion: Neumayer (1904) introduced the terms “heteropo-lar” and “amphipolar” to describe the coiling of spiral coprolites, andthese terms have been widely accepted (e.g., Williams, 1972; Duffin,1979; Jain, 1983; Hunt et al., 1994, 1998). To provide a framework fordescription, we refer to the tightly coiled end of a heteropolar coproliteas anterior and the line of separation at the posterior end of the tightly-coiled segment as the spiral demarcation. The anterior portion (typically30-40%) of heteropolar coprolites is tightly coiled, and the posteriorfraction consists of one long coil with a wide lip (sensu Jain, 1983).Amphipolar coprolites (sensu Neumayer, 1904) exhibit an even distri-bution of coils (e.g., Hyronocopros: Hunt et al., 2005d, fig. 3).

Jain (1983) utilized the term “amphipolar” to refer to coprolitesthat have multiple spirals that extend for more than 50% of the length ofthe coprolite (e.g., Jain, 1983, fig. 2B) but that do not extend the wholelength, so they are not truly amphipolar (sensu Neumayer, 1904). Thesecoprolites are often reminiscent of trochospiral gastropods in overallmorphology (e.g., Fig. 5A) and are, technically as well as etymologically,heteropolar in form. We introduce here the terms “microspiral” for themore typical heteropolar coprolites such as Heteropolacopros in whichthe markedly spiral portion constitutes less than 50% of the overalllength, and “macrospiral” for the forms described by Jain (1983) inwhich the tightly spiral portion of the coprolite constitutes 50% or moreof its length (Fig. 6). As with microspiral heteropolar coprolites, thelargest diameter of the macrospiral Liassocopros is at the posterior endof the tightly coiled fraction of the coprolite. Liassocopros is broaderrelative to its height than are other heteropolar coprolites.

These coprolites are abundant in the Lower Jurassic of England(e.g., Buckland, 1829; Hawkins, 1834). The first coprolite describedfrom North America derives from the Upper Cretaceous of New Jerseyand probably also represents this ichnotaxon (DeKay, 1830, pl. 3, fig. 6).

Liassocopros hawkinsi isp. nov.

Holotype: BMNH R. 2107, coprolite (Fig. 5D-E).Type locality: Lyme Regis, England.Type horizon: Lower Lias.Etymology: Named for Thomas Hawkins, who described speci-

mens attributed here to this ichnogenus in 1834.Distribution: As for the ichnogenus.Referred specimens: Coprolites from the Lower Lias of Lyme

Regis, England (Fig. 5A-C: Buckland, 1829, pl. 28, figs. 4, 7, pl. 29, fig.1).

Diagnosis: As for genus.Discussion: Hawkins (1834, pls. 27-28; 1840, pls. 29-30) illus-

trated a number of coprolites from the Lower Lias of Lyme Regis that, in1840, were still listed as “in the Author’s Collections, not yet transferredto the British Museum” (Hawkins, 1840, unnumbered page – list ofplates). These specimens, which were collected by Mary Anning, weresubsequently “transferred,” and several are illustrated herein, includingthe holotype of Liassocopros hawkinsi (compare Fig. 5D and Hawkins,1834, pl. 28; 1840, pl. 30) and a referred specimen of Saurocoprosbucklandi (compare Fig. 4A and Hawkins, 1834, pl. 27; 1840, pl. 29 –note that the image is reversed in Hawkins’ plates and that the specimenhas lost part of its posterior extremity during the last 167 years!).

It is possible that two forms may be distinguishable within thisichnospecies. One form is trochospiral with an acute anterior tip (Fig.5A) and the other has much more rounded anterior and posterior extremi-ties (Fig. 5D).

Malericopros igen. nov.

Type species: Malericopros matleyi isp. nov.

FIGURE 2. A-B, Heteropolacopros texaniensis, UMMP 7253 (partim),holotype, in lateral views from the Tecovas Formation, Crosby County,West Texas, USA. C-D, Heteropolacopros texaniensis, ISI P.58, in lateralviews from the Maleri Formation, India. E-F, Heteropolacopros texaniensis,ISI P.51, in lateral views from the Maleri Formation, India. G-I,Heteropolacopros texaniensis, UMMP 7253 (partim), topotypes of H.texaniensis in lateral views from the Tecovas Formation, Crosby County,West Texas, USA. J-K, Dicynodontocopros maximus , UMMP 7255,holotype in lateral views from the Tecovas Formation, Crosby County,West Texas, USA. Note that Hunt et al. (1998, p. 228, 229) incorrectlylisted the number of the holotype of Dicynodontocopros maximus as UMMP7253 and UMMP 7285). A-B, after Hunt et al. (1998, fig. 2K-L); C-F, afterJain (1983, pl. 82, figs. 5-6, 10-11).

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Included species: Known only from the type species.Etymology: From the Maleri Formation, which yielded the holo-

type of the ichnogenus, and the Greek kopros for “dung.”Distribution: Late Triassic of India.Diagnosis: Microspiral heteropolar coprolite that differs from

Saurocopros and Heteropolacopros in that the maximum diameter isposterior to the spiral demarcation.

Discussion: In all other heteropolar coprolites (Heteropolacopros,

Saurocopros, Liassocopros) the maximum diameter is near the posteriorend of the tightly-coiled portion of the coprolite, and the portion of thecoprolite posterior to the spiral demarcation tapers in lateral view.

Malericopros matleyi isp. nov.

Holotype: GSI K.42/419 (Fig. 5F; Matley, 1939a, pl. 33, fig. 1;Jain, 1983, pl. 82, fig. 9).

Type locality: Near Maleri, India.Type horizon: ?Lower Maleri Formation (upper Carnian).Etymology: Named for C. A. Matley, who first described Indian

Triassic and Cretaceous coprolites in detail.Distribution: As for the ichnogenus.Referred specimens: ISI P.71 (Jain, 1983, pl. 81, fig. 10).Diagnosis: As for the ichnogenus.Discussion: Currently, this ichnospecies is only known from the

Upper Cretaceous of India.

Falcatocopros igen. nov.

Type species: Falcatocopros oxfordensis isp. nov.Included species: Known only from the type species.Etymology: From the Latin falcatus for “crescent,” referring to

the thin, curved shape of this coprolite, and the Greek kopros, for “dung.”Distribution: Early-Late Jurassic of England.Diagnosis: Differs from other coprolite ichnogenera in being long,

narrow and arcuate in lateral view, rounded to sub-rounded in crosssection with a width that gradually decreases from one end to the other.

Discussion: This ichnogenus is currently only documented fromthe Jurassic, but it may be present in Rhaetian ichnofaunas.

Falcatocopros oxfordensis isp. nov.

Holotype: BMNH R. 2094, coprolite (Fig. 5H).Type locality: Near Peterborough, England.Type horizon: Oxford Clay.Etymology: Named for the Oxford Clay, which yielded the holo-

type.Distribution: As for the ichnogenus.Referred specimens: BMMNH R 2110, coprolite, Lower Lias,

Lyme Regis, England (Fig. 5G; Hawkins, 1834, pl. 35; 1840, pl. 30).Diagnosis: As for the ichnogenus.Discussion: This highly distinctive ichnospecies is uncommon,

probably, at least in part, the result of a taphonomic artifact related to itsslender morphology.

TRIASSIC VERTEBRATE BIOCHRONOLOGY

Lucas and co-workers (Lucas and Hunt, 1993; Lucas, 1997, 1998,1999; Lucas and Hancox, 2001; Lucas and Huber, 2003) have developeda global biochronological scheme for Triassic tetrapods. This schemeinvolves the definition of eight land-vertebrate faunachrons (lvfs) toencompass Triassic time. Subsequently, Lucas and others (Lucas, 1997,1998; Hunt, 2001; Hunt et al., 2005a; Lucas et al., 2007) further refinedthis biochronology. In the following review of Triassic coprolites, weutilize this biochronology wherever possible.

TRIASSIC VERTEBRATE COPROLITE RECORD

Early Triassic

Northwood (1997, 2005) published the most thorough study of aTriassic coprolite ichnofauna, describing specimens from the ArcadiaFormation in Queensland, northeastern Australia. Northwood (2005)recognized three main forms of coprolites (although obviously did notutilize the ichnotaxa erected herein): (1) amphipolar coprolites assign-able to Hyronocopros amphipola (Hunt et al., 2005c); (2) longitudinally-striated coprolites representing Alococopros triassicus; and (3) indeter-minate coprolites. Hyronocopros amphipola and Alococopros triassicus

FIGURE 3. A-B, Alococopros triassicus igen. et isp. nov., UMMP 7253(partim), in lateral views, from the Tecovas Formation, Crosby County,West Texas, USA. C-E, Alococopros indicus igen. et isp. nov., BMNHunnumbered (Matley collection), in lateral views from the Lameta Formation,India.

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constitute less than 25% of the sample. She also noted that some of thebroken coprolites may represent a heteropolar form based on the largenumber of whorls in cross section. The Acadia coprolites commonlycontain inclusions (over 50%)that include two kinds of cyanobacteria,macrofloral specimens and rare invertebrate specimens (e.g., conchostracanvalves, impressions of an insect wing, an insect head segment), scales,teeth, tooth plates and bones of actinopterygian and dipnoan fish andfragmentary amphibians (Northwood, 2005). Dipnoan remains were rela-tively more common in Alococopros specimens (Northwood, 2005).

Benz (1980) reported coprolites from the Moqui Member of theMoenkopi Formation in northern Arizona. However, Benz (1980) in-cluded the lower portion of the superjacent Holbrook Member (MiddleTriassic) within the Moqui and there none of the coprolites that shedescribed are actually from the Lower Triassic.

Middle Triassic

Ochev (1974) described coprolites from four Middle Triassic lo-calities, one in Kazakhstan and three in Russia: (1) Mollo-Khara-Bala-Kantemir (Kazakhstan); (2) Karagachka; (3) Donguz I; and (4) BukobayV. Ochev (1974) discriminated three types of coprolites that he com-pared with those described from the Upper Triassic of West Texas byCase (1922). The most easily identified are longitudinally striated formsassignable to Alococopros triassicus (Ochev, 1974, fig. 1e-f). These co-prolites are described as being quite common.

Ochev (1974, fig. 1d) compares spirally-coiled specimens to those

illustrated by Case (1922, fig. 33A-B) assigned to Heteropolacoprostexaniensis by Hunt et al. (1998). However, the one specimen that isillustrated appears to be amphipolar in form, rather than heteropolar asdescribed (Ochev, 1974, fig. 1d). These coprolites are noted as less com-mon. The third form of coprolite is described as large (2-10 cm long),with a smooth surface and containing possible plant impressions. Theonly illustration of this type of coprolite is a cross section (Ochev, 1974,fig. 1c). Ochev (1974) compares this large form with specimens thatCase collected and briefly described, but did not illustrate, some of whichrepresent Dicynodontocopros maximus (Hunt et al., 1998, fig. 2A-B).Ochev (1974) listed the occurrence of his three types of coprolites as: (1)Mollo-Khara-Bala-Kantemir – all three forms; (2) Karagachka -Alococopros triassicus and small specimens of the large morphotype;and (3) Donguz I and Bukobay – spiral and large forms.

Rusconi (1947, figs. 1-4; 1949, figs. 2-6) described Triassic andPermian coprolites from the Mendoza area in Argentina. The large Trias-sic sample is dominated by spiral forms (e.g., Rusconi, 1949, figs. 2-4)but also includes small, cylindrical forms (Rusconi, 1949, fig. 5) andlarge, wide amorphous forms (Rusconi, 1949, fig. 6). The large forms areup to 120 mm in length and 58 mm in width. They are comparable in sizeto Dicynodontocopros but differ in having more rounded terminationsand a more regular width. It is possible that these differences aretaphonomic in origin.

The spiral coprolites appear to be dominantly heteropolar (e.g.,Rusconi, 1949, fig. 4, first two coprolites in first row) although a fewmay be amphipolar (e.g., Rusconi, 1949, fig. 2, bottom left). They arerelatively short and wide compared to the holotype of Heteropolacopros(Hunt et al., 1998, fig. 2K-L). The heteropolar coprolites are apparentlymainly macrospiral. Some spiral forms include ganoid scales, possiblyreferable to the holostean fish Pholidophorus. We tentatively assignsome of these coprolites to Liassocopros (e.g., Rusconi, 1949, fig, 2,center right) based on their macrospiral structure and width:length ratios.

Benz (1980) reported coprolites from the Holbrook Member ofthe Moenkopi Formation at Radar Mesa in northern Arizona. Benz(1980, pl. 7) illustrated some indeterminate coprolites and noted thatcoprolites were locally abundant. Many contain temnospondyl bones,including intercentra (Morales, 1987). Coprolites are present at otherMoenkopi localities, but they have not been described. We, for example,have observed coprolites at several localities near the town of Holbrook.There is an unstudied coprolite collection at the MNA.

Fraas (1891) reported that spiral coprolites are common in theGerman Muschelkalk, and he attributed them to sharks. The Muschelkalkranges in age from Anisian to Ladinian.

In India, the Yerrapilli Formation (early Middle Triassic) yieldsspherical, ovoid and elliptical coprolites (Chatterjee, 1967; Jain, 1983).These specimens are covered by desiccation cracks and differ in mor-phology from those from the Late Triassic of India (Jain, 1983).

Late Triassic

The majority of Triassic vertebrate coprolites in museum collec-tions and mentioned or described in the literature are from the LateTriassic. Vertebrate coprolites are common and locally abundant in strataof the Upper Triassic Chinle Group of Lucas (1993) in western NorthAmerica (Hunt and Lucas, 1989, 1993a, b; Murry, 1989; Murry andLong, 1989; Heckert et al., 2005; Hunt et al., 1998, 2005c).

The Newark Supergroup of eastern North America ranges in agefrom Middle Triassic-Early Jurassic. There has been more study of thevertebrate trace fossils of this stratigraphic unit, almost exclusively tracks,than any other over the last 150 years (Hitchcock, 1858; Lull, 1953;Olsen, 1988; Olsen et al., 1998). However, the coprolites of the Newarkhave been virtually ignored. The few references to coprolites in pub-lished works suggest that they are most common in the Carnian andJurassic portions of the Newark (Olsen, 1988; Olsen et al., 1989, 2003,2005a,b; Olsen and Flynn, 1989; Olsen and Huber, 1998; Olsen andRainforth, 2002; Gilfillian and Olsen, 2000).

FIGURE 4. A-G, Saurocopros bucklandi igen. et isp. nov. from the Lower“Lias” of southwestern England. A, BMNH R 41285 from Charmouth inlateral view. B, BMNH R 2102, holotype of Saurocopros bucklandi igen. etisp. nov. from Lyme Regis. C-D, BMNH R 1402, one specimen in lateralview from Lyme Regis. Note the abundant inclusions and that one side isabraded. E-G, Specimens from the Buckland collection, presumably at theUniversity of Oxford, three specimens in lateral view from Lyme Regis. E-G, after Buckland, 1829, pl. 28, figs.6, 7, 9, specimens rotated 180° fromoriginal publication, are to the same scale and G, is 10.7 cm long. A-D,Specimens collected by Mary Anning.

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CarnianVertebrate coprolites are common and locally abundant in the

upper Carnian strata of the Upper Triassic Chinle Group of Lucas (1993)in western North America. The oldest coprolites are from the Otischalkianof West Texas. Elder (1978, 1987) described coprolites from the Colo-rado City Formation near Midland, noting that they are particularlyabundant at Otis Chalk quarries 1 and 2. Elder (1978) explicitly discrimi-nated the same three morphologies as Case (1922), notably heterospiralforms representing Heteropolacopros texaniensis (Elder, 1978, pl. 14,fig. 1a), longitudinally-striated forms assignable here to Alococoprostriassicus (Elder, 1978, pl. 14, fig. 1b) and a third variable and indetermi-nate form (Elder, 1978, pl. 14, figs. 1c-d). The indeterminate forms, atleast as illustrated, do not represent Dicynodontocopros. The NMMNHcollection includes indeterminate coprolites from the Popo Agie Forma-tion of Wyoming (Hunt et al., 1998).

Outcrops of younger Carnian (Adamanian) Chinle strata are muchmore widespread. Lipman and McLees (1940) described a new speciesof bacteria, Thiobacillus coproliticus, from a coprolite from Arizona, butdid not describe the coprolite that yielded it. Case (1922) recognizedthree coprolite forms from the Tecovas Formation of West Texas thatinclude the holotype and referred specimens of Heteropolacoprostexaniensis (Case, 1922, fig. 33A-B; Hunt et al., 1998, fig. 2C-L). Case(1922, fig. 33C-D) also described specimens now referable to Alococoprostriassicus. Coprolites of the third category described by Case (1922, p.83) are large (5-18 cm long), smooth surfaced and lack vertebrate inclu-sions. One of these specimens in the UMMP collection is the holotypeof Dicynodontocopros maximus (Hunt et al., 1998, fig. 2A-B). Otherspecimens in the collection are smaller and lack a distinct morphology.

Hunt et al. (1998) described Dicynodontocopros maximus fromthe Bluewater Creek Formation at the Placerias quarry near St. Johns,

Arizona. Coprolites are locally abundant in the Placerias quarry (e.g.,Camp and Welles, 1956; Kaye and Padian, 1994). Hunt et al. (1998) alsonoted that Heteropolacopros texaniensis occurs in the Blue Mesa Mem-ber of northeastern Arizona at Petrified Forest National Park (Hunt andSantucci, 1994). Coprolites, some of which contain fish scales, teeth andplant debris, are common in the Blue Mesa Member at the “DyingGrounds” locality in Petrified Forest National Park (e.g., Murry andLong, 1989; Heckert, 2001, 2004). Undescribed coprolites occur in theBlue Mesa and Painted Desert members of the Petrified Forest Forma-tion at Petrified Forest National Park. Wahl et al. (1998) described evi-dence of coprophagy in the Blue Mesa Member of Petrified ForestNational Park.

Ash (1978a, b) described a large number of coprolites from alacustrine mudstone unit in the Bluewater Creek Formation in westernNew Mexico that he subsequently donated to the NMMNH. Ash (1978a)recognized three main forms: cylindrical, cigar-shaped with tapered ends(rare), and spiral. The spiral coprolites are microspiral and heteropolar;some clearly represent Heteropolacopros texaniensis (e.g., Ash, 1978a,fig. 2h) and at least one specimen represents Alococopros triassicus(Ash, 1978a, fig. 2g). Weber and Lawler (1978) analyzed the lipid con-tent of a sample of these coprolites. Other localities in the BluewaterCreek Formation yield abundant coprolites (Heckert and Lucas, 2003).

Other Adamanian coprolites in New Mexico are known from theLos Esteros Member of the Santa Rosa Formation, Garita Creek Forma-tion, lower Petrified Forest Formation and Salitral Formation (Hunt andLucas, 1988, 1990, 1993; Hunt et al., 1989). Parrish (1999) reportedabundant coprolites from the Monitor Butte Formation in southern Utah.

There are a few references to coprolites in the Carnian portion ofthe Newark Supergroup. Olsen (1988) noted abundant coprolites in theCumnock Formation. The Lockatong Formation yields coprolites fromseveral localities (Olsen et al., 1989; Olsen and Flynn, 1989; Olsen andRainforth, 2002; Jenkins in Häntzschel et al., 1968; YPM PU speci-mens). Olsen and Huber (1998, table 1) noted coprolites in the PekinFormation in North Carolina.

Burmeister et al. (2006, fig. 6) described coprolites from the Isalo“Group” (Isalo II beds) of Madagascar. These coprolites are 10-60 mmin length and nonspiral. About 5% of the coprolites contain fish bonesand scales.

Carnian/NorianOldham (1859, pl. 15, figs. 11-12) first described coprolites from

the Maleri Formation of India. The Maleri Formation is known to spanthe Carnian/Norian boundary and to contain both late Carnian and early-middle Norian faunas (Bandyopadhyay and Sengupta, 2006). Most fos-sils appear to derive from the upper Carnian portion of the MaleriFormation, but we are not certain of the exact age of any of the Malericoprolites described by Oldham or many subsequent workers.

King (1881, p. 271-272) noted that, in the Maleri, the “common-est remains are coprolites which lie about the fields in large numbers, ofall sizes and shapes, from the short cylindrical forms with tapering endsand spiral foldings up to large flat rudely discoid coils.” Aiyengar (1937,p. 104) mentioned that “coprolites are abundant about a mile W.S.W. ofMaleri” and he later reported (in Matley, 1939a, p. 531) that thesecoprolites are found in red clays in association with Ceratodus and twolarge reptile vertebrae “which have been described by F. von Huene as anew species of reptile” and so these coprolites thus presumably derivefrom the lower Maleri (Huene, 1940). Aiyengar (in Matley, 1939a) alsonotes that another locality about a mile southwest of Maleri yielded largereptile bones from a calcareous sandstone and lacked coprolites. Matley(1939a) described coprolites first described by Oldham (1859) and onethat is inferred to have been collected by Aiyengar from the lower Maleri.Matley (1939a, pl. 33) described these coprolites as fusiform and spiralin structure and varying in length from 55 to about 80 mm long. Thesecoprolites include the holotype of Malericopros matleyi (Matley, 1939a,pl. 33, figs.1a-b), possible specimens of Heteropolacopros texaniensis

FIGURE 5. A-E, Liassocopros hawkinsi igen. et isp. nov. from the Lower“Lias” of Lyme Regis, England. A-C, Three specimens in the Bucklandcollection, presumably at the University of Oxford, in lateral view. D-E,BMNH R 2107, holotype of Liassocopros hawkinsi igen. et isp. nov., inlateral views. F, Malericopros matleyi igen. et isp. nov., GSI K. 42/419,holotype in lateral view, from the ?lower Maleri Formation, near Maleri,India. G-H, Falcatocopros oxfordensis igen. et isp. nov. from the Jurassicof England. G, BMNH R 2110, from the Lower “Lias” of Lyme Regis, inlateral view. H, BMNH R 2094 (Leeds collection), holotype of Falcatocoprosoxfordensis igen. et isp. nov., from the Oxford Clay at Peterborough, inlateral view. A-C, after Buckland (1841, v. 2, pl. 15, p. 27-29); F, after Jain(1983, pl. 82, fig. 9).

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(Matley, 1939a, pl. 33, figs. 4), probable specimens of Liassocoproshawkinsi (Matley, 1939a, pl. 33, figs. 5a), a possible specimen ofSaurocopros bucklandi (Matley, 1939a, pl. 33, fig. 8) and apparentlyamphipolar forms (Matley, 1939a, pl. 33, fig. 3).

Sohn and Chatterjee (1979) described ostracodes from coprolitesfrom the lower Maleri Formation. These coprolites are described as adistinct type from near Achlapur village. They are large, with lengthsfrom 7 to 10 cm and widths from 50 to 80 mm. Sohn and Chatterjee (pl.1, fig. 4-5) only illustrated a fragment of the end of one coprolite. Thesecoprolites were found near some rhynchosaur bones, so they are clearlyfrom the lower (upper Carnian) portion of the Maleri.

Jain (1983) described a sample of coprolites from the lower MaleriFormation that are heteropolar, amphipolar and non-spiral. Some ofthese specimens pertain to Heteropolacopros texaniensis (Jain, 1983, pl.82, figs. 1-6, 10-11), Malericopros matleyi (Jain, 1983, pl. 82, fig. 9) andLiassocopros hawkinsi (Jain, 1983, pl. 81, figs. 5, 10). Other fragmen-tary spiral coprolites are heteropolar (e.g., Jain, 1983, pl. 81, figs. 6, 8,11-14) and possibly amphipolar (e.g., Jain, 1983, pl. 81, fig. 16).

Buckland (1841, p. 13) noted that “Professor Jaeger has recentlydiscovered many Coprolites [sic] in the alum slate of Gaildorf [sic] inWirtemberg [sic]; a formation which he considers to be in the lowerregion of that part of the new red sandstone formation which in Germanyis called Keuper.” The classic “Keuper,” like the Maleri, is of both lateCarnian and Norian age. Fraas (1891) reported common spiral coprolitesfrom the “Keuper,” which he attributed to sharks. Major Europeanmuseum collections do not include “Keuper” coprolites (e.g., NaturalHistory Museum, London and Museum für Naturkunde, Stuttgart).

DeBlieux et al. (2006, figs. 9A-C) illustrated numerous coprolitesfrom the Petrified Forest Formation of Zion National Park in southernUtah. These specimens could be of either Carnian or Norian age.

NorianThe Bull Canyon Formation of east-central New Mexico yields

large coprofaunas. Lucas et al. (1985) described three morphologies ofcoprolites: (1) longitudinally furrowed specimens that representAlococopros triassicus (Lucas et al., 1985, fig. 7M-R); (2) small, rod-liketo oval morphology (> 90% of sample) (Lucas et al., 1985, fig. 7A-L);and (3) large, irregularly shaped forms with numerous inclusions (fishscales, bone fragments) (Lucas et al., 1985, fig. 7S-U). This sample(NMMNH locality 110) is from the younger Lucianoan sub-lvf of theRevueltian. The NMMNH also contains a large sample from the olderBarrancan time interval (NMMNH locality 1) and numerous isolatedspecimens from various Barrancan loaclities.

Coprolites are present at other Revueltian Chinle localities inNew Mexico, including the upper Petrified Forest Formation in the SanYsidro area (Hunt and Lucas, 1990) and Chama Basin (Hunt and Lucas,1993), Trujillo Formation (Hunt, 1991) and Correo Sandstone Memberof Petrified Forest Formation at Mesa Gigante and the Hagan Basin(Hunt and Lucas, 1993b).

In Arizona, Revueltian coprolites occur in the Painted DesertMember of the Petrified Forest Formation at Petrified Forest NationalPark (Hunt and Santucci, 1994). Coprolites are also common in the OwlRock Formation at Ward Terrace (Kirby, 1989).

Late Norian/RhaetianCoprolites are locally common in Apachean strata of the Chinle

Group, notably in New Mexico and Utah. Hunt et al. (1993) noted thatcoprolites were common in the Bell Springs Formation in northeasternUtah. Coprolites occur on the main track bed at the Shay Canyon tracksite(Rock Point Formation) in southeastern Utah (Lockley, 1986; Lockleyand Hunt, 1995, fig. 3.8).

In New Mexico, coprolites are locally abundant in the RedondaFormation of east-central New Mexico. The largest concentration is atthe Gregory quarry (NMMNH locality 485) in Apache Canyon. Thislarge sample lacks Heteropolacopros texaniensis and Alococoprostriassicus.

One of the most interesting occurrences of coprolites in the Trias-sic occurs at the Coelophysis quarry in north-central New Mexico. Co-prolites occur associated with skeletons of Coelophysis (Rinehart et al.,2005a,b). These coprolites occur in the vicinity of the cloaca in morethan one skeleton and include bones assignable to Coelophysis, whichindicates cannibalism in this early dinosaur (Rinehart et al., 2005a, bcontra Nesbitt et al., 2006).

RhaetianBuckland (1829) first recognized coprolites from the Rhaetian

Westbury Formation (Penarth Group) of England (Swift and Duffin,1999). Coprolites are common in the bone beds of the Westbury Forma-tion (Buckland, 1829; Duffin, 1979; Storrs, 1994; Martill, 1999; Swiftand Duffin, 1999). Duffin (1979; Swift and Duffin, 1999) recognizedfour broad morphological types of coprolites. However, two of thesecategories included both amphipolar and heteropolar forms, which weregard as fundamentally distinct morphologies representative of differ-ent ichnotaxa (e.g., Hunt et al., 1998, 2005c). Therefore, we recognize sixcategories:

1. Large (up to 80 mm), usually brown, often tapered with well-defined amphipolar structure. Undigested vertebrate remains include fishscales (often tangential or normal to spiral folds) and crustacean remains(Tropifer laevis, possible isopods). Discrete food “boli” are discernablein thin section. Swift and Duffin (1999) interpreted these specimens torepresent sharks, possibly myriacanthid holocephalans and palaeoniscidchondrostreans (coprolites with vertebrate inclusions) or dipnoans (co-prolites with arthropod inclusions).

2. Large (up to 80 mm), usually brown, often tapered with well-defined heteropolar structure. Swift and Duffin (1999, fig. 32A) de-scribed coprolites of this general form as amphipolar, but the specimenthat they illustrate is clearly heteropolar in morphology. Undigestedvertebrate remains include fish scales (often tangential or normal to spiralfolds) and crustacean remains (Tropifer laevis, possible isopods). Dis-crete food “boli” are discernable in thin section. Swift and Duffin (1999)interpreted these specimens to represent sharks, possibly myriacanthidholocephalans and palaeoniscid chondrostreans (coprolites with verte-brate inclusions) or dipnoans (coprolites with arthropod inclusions).

3. Elongate (~30 mm long) with amphipolar coiling and no visiblevertebrate and invertebrate inclusions (Swift and Duffin, 1999, fig. 32B).Swift and Duffin (1999) attributed these coprolites to Ceratodus ormyriacanthid holocephalans.

4. Elongate (~30 mm long) with heteropolar coiling and no visiblevertebrate and invertebrate inclusions. Swift and Duffin (1999) attrib-uted these coprolites to Ceratodus or myriacanthid holocephalans.

5. Small (maximum 30 mm long), capsule-shaped, lacking spiralform or inclusions. They are often black and shiny due to polishing andabrasion during post-fossilization transport. These coprolites are usu-ally homogeneous in thin section with disseminated pyrite (Swift andDuffin, 1999, fig. 32C).

FIGURE 6. Principal morphotypes of amphipolar (A), microspiralheteropolar (B-D) and macrospiral heteropolar (E) coprolites. A,Hyronocopros. B, Heteropolacopros. C, Malericopros. D, Saurocopros. E,Liassocopros. Not to scale.

956. Small (up to 30 mm long) flattened, shiny forms. They include

undigested scales and teeth, but no internal spiraling (Swift and Duffin,1999, fig. 32D). Swift and Duffin (1999) attributed these coprolitespossibly to small reptiles. Duffin (1979) notes that types 5 and 6 are byfar the most common.

Some Type 2 (notation above, not that of Duffin or Swift andDuffin) coprolites pertain to Liassocopros hawkinsi (BCM 4891; Duffin,1979, pl. 21, fig. 1; Swift and Duffin, 1999, fig. 32A). Some specimens ofType 4 may represent Saurocopros bucklandi (NMW G2066; Duffin,1979, pl. 21, fig. 3; Swift and Duffin, 1999, fig. 32B).

Coprolites also occur in other Rhaetic bone beds in western Eu-rope although none have been described.

TRIASSIC COPROLITE BIOSTRATIGRAPHY

Coprolites are potentially of biochronological utility (e.g., Hunt,1992; Hunt et al., 1998, 2005a). Trace fossils generally represent higherlevel taxonomic groups of body fossils. Thus, track ichnogenera arecommonly only equivalent to the “family” level of body fossils (Lucas,2007). Coprolites probably represent, in most cases, even higher leveltaxonomic levels (“order” or above). However, the stratigraphic distribu-tion of coprolites obviously mirrors the stratigraphic ranges of the ani-mals that produced them. Also, given also that some localities/strati-graphic units produce numerous coprolites and no body fossils, there isa potential to utilize coprolites in biochronology. We can presently rec-ognize the following ranges for ichnotaxa that are present in the Triassic:

Permian: Hyronocopros amphipolar, Heteropolacoprostexaniensis

Early Triassic: Hyronocopros amphipolar, Alococopros triassicus.Middle Triassic: Alococopros triassicus, ?Liassocopros isp.Late Triassic: Heteropolacopros texaniensis, Alococopros

triassicus.Dicynodontocopros maximus, Malericopros matleyi, Liassocopros

hawkinsi, Saurocopros bucklandi.Early Jurassic: Liassocopros hawkinsi, Saurocopros bucklandi.Alococopros triassicus is a good index fossil for the Triassic be-

cause it is easily identifiable, widespread and relatively common. Thecharacteristic Early Permian Hyronocopros amphipolar also appears tobe restricted to the Early Triassic. Even though certain Middle Triassicspecimens could represent this ichnotaxon, it is certainly absent in theLate Triassic.

Dicynodontocopros maximus and Malericopros matleyi are bothrestricted to the Late Triassic (upper Carnian), but their distribution islimited. The ubiquitous Early Jurassic Liassocopros hawkinsi andSaurocopros bucklandi have their first appearance in the Late Triassic.Heteropolacopros texaniensis is not currently known from strata youngerthan Carnian.

There appears to be a change in coprofaunas near the end of theNorian. Apachean (upper Norian/Rhaetian) and Rhaetian assemblageslack the long-ranging Alococopros triassicus and/or include good examplesof the characteristic Jurassic Liassocopros hawkinsi and Saurocoprosbucklandi. This change is apparent in both the nonmarine Chinle Groupand the mixed marine and nonmarine Westbury Formation.

TRIASSIC COPROLITES AND ICHNOFACIES

Hunt et al. (1994, 1998) recognized “coprofacies” in the UpperTriassic of western North America. Hunt and Lucas (2007) noted that

these should be referred to as ichnocoenoses because of their relativelylimited distribution in space and time.

Hunt et al. (1994, 1998) distinguished three ichnocoenoses: (1)Dicynodontocopros ichnocoenosis, in which coprolites occur in gray toblack mudstones that formed in alternating wet and dry conditions, in-cluding periods of standing water, and are associated with aquatic verte-brate microfossils; (2) Heteropolacopros ichnocoenosis, which occurs influvial redbeds; and (3) ovoid, structureless coprolite ichnocoenosis,which occurs in highly carbonaceous strata that formed in ponds. Thereis a third obvious Triassic ichnocoenosis that yields significant speci-mens of spiral coprolites, notably Liassocopros hawkinsi, and occurs inshallow marine strata, with the exemplar being the Westbury Formation.

Are any of these ichnocoenoses pervasive enough, spatially andtemporally, to be considered to be ichnofacies? Arguably, at least tworepresent widely distributed ichnofacies. The Liassocopros ichnofaciesis characterized by a prevalence of spiraled coprolites that occur inshallow marine strata. This ichnofacies is represented at least in thePennsylvanian (Zangerl and Richardson, 1963), Early Permian (Will-iams, 1972) and Early Jurassic (Buckland, 1829) as well as the UpperTriassic.

The Heteropolacopros ichnofacies is characterized by the pres-ence of microspiral heteropolar coprolites that occur in fluvial redbeds.This ichnofacies occurs at least from the Early Permian (Hunt et al.,2005b, c) until the Late Triassic.

It seems reasonable that a Dicynodontocopros ichnofacies, whichcontains large herbivore coprolites, might characterize swampy environ-ments and that an ichnofacies, which we could term the Alococoprosichnofacies, should characterize ponds. However, we do not have thedata to support these hypotheses.

PROSPECTUS FOR FUTURE WORK

In the last few years we have made a concerted effort to describeand document Permo-Triassic coprolites (e.g., Hunt et al., 1994, 1998,2005a, b, c; Hunt and Lucas, 2005a, b, c). This work is based on theextensive samples that we have collected and the very limited collectionsin other museums. We have four basic purposes in these works:

1. To raise awareness of the general abundance of the vertebratecoprolite fossil record and its potential importance.

2. To demonstrate that distinct morphologies can be discrimi-nated, described and of utility.

3. To illustrate that vertebrate coprolites have importance inbiochronology.

4. To suggest that coprolites have utility in ichnofacies studies.Despite these lofty goals, we realize that vertebrate coprolites

have been grossly undersampled and that paleoscatology is in a proteanstage. We hope that other workers will be inspired to collect and describemore vertebrate coprolites and to further this still nascent sub-disciplineof paleontology.

ACKNOWLEDGMENTS

We thank Gregg Gunnell, Sandra Chapman, Angela Milner, MichaelMorales, Joseph Gregory, John Ostrom, Walter Joyce and Kevin Padianfor access to specimens in their care and Jerry Harris and Larry Rinehartfor helpful reviews.

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APPENDIX

List of Triassic coprolite specimens in the Geoscience Collection of the New Mexico Museum of Natural History and Science, Albuquerque.

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