380 20.1. Number of Tracksites Through Time. Note the Late Jurassic

Home , Parabrontopodus

0 10 20 30 40 50 60 Maastrichtian Campanian Santonian Coniacian Turonian-Coniacian Cenomanian 100.5 My Albian Aptian Barremian Hauterivian-Barremian Hauterivian Valanginian Berriasian Tithonian-Berriasian ~145.0 My Tithonian Kimmeridgian-?Tithonian Kimmeridgian Oxfordian-Kimmeridgian Oxfordian ~163.5 My Callovian Bathonian Bajocian-Bathonian Bajocian Aalenian ~174.1 My Toarcian Sinemurian-Pliensbachian Sinemurian Hettangian-Pliensbachian Hettangian-Sinemurian Hettangian 0 10 20 30 40 50 60

20.1. Number of tracksites through time. Note the Late Jurassic and the Early Cretaceous peaks related to the high number of tracksites from the Reuchenette Formation (Late Jurassic, Switzerland) and the Glen Rose Formation (Early Cretaceous, United States). No record for the Pliensbachian, Toarcian (Early Jurassic), the Aalenian, Callovian (Middle Jurassic), and for the Santonian and Campanian stages (Late Cretaceous) is known. Number of tracksites is on the y-axis, the Jurassic and Cretaceous stages are on the x-axis.

380

DinoTracks.indb 380 1/22/16 11:23 AM A Review of the Dinosaur Track Record from Jurassic and Cretaceous Shallow Marine 20 Carbonate Depositional Environments

Simone D’Orazi Porchetti, Massimo Bernardi, Andrea Cinquegranelli, Vanda Faria dos Santos, Daniel Marty, Fabio Massimo Petti, Paulo Sá Caetano, and Alexander Wagensommer

An extensive literature on dinosaur ichnology et al. 2003, 2010), and more recently also from the south- is available today, with hundreds of papers describing di- ern hemisphere in South America (Moreno and Pino, 2002; nosaur tracks in different depositional settings. In recent Moreno and Benton, 2005; Pazos et al., 2012) and Madagascar years, it has become common practice in paleontology to (Wagensommer, Latiano, and Nicosia, 2010; Wagensommer gather data in databases to ease organization, managing, et al., 2012). and analysis of large amounts of information. A review of the Lockley, Hunt, and Meyer (1994) first noticed the recur- occurrences of dinosaur tracks in shallow marine carbonate rent, statistically significant association of sauropod tracks, depositional environments is presented here, based on 131 pa- especially wide-gauge tracks, together with less-abundant pers published between 1917 and 2013 and describing a total theropod tracks, in low latitude carbonate-dominated sys- of 212 tracksites. The raw data set, which adopts the tracksite tems, and introduced the term Brontopodus ichnofacies as the basic unit, reveals an abundance of saurischian foot- for this association. The occurrence of dinosaur tracks in prints and a relative paucity of ornithischian tracks in these these environments also led to some important paleoenvi- depositional environments. Theropods are dominant during ronmental and paleogeographical reinterpretations, as in most of the Jurassic and the Cretaceous period, whereas sau- the case of the Tethyan, peri-adriatic carbonate platforms ropods are always well represented and dominant at least in (Bosellini, 2002; Dalla Vecchia, 2003, 2005; Nicosia et al., the Late Jurassic. A complementary database for siliciclastic 2007; Zarcone et al., 2010), and the Jura carbonate platform coastal and inland depositional settings is required before (e.g., Meyer, 1990, 1993; Meyer and Hauser, 1994; Marty and testing for trends and patterns to investigate the relationships Meyer, 2013), which prior to the discovery of dinosaur tracks between dinosaur tracks and facies/environment in general. were thought to be devoid of terrestrial biota. However, it is This contribution intends to promote new large-scale evolu- still rather poorly understood whether dinosaurs used these tionary studies based on track data. environments only occasionally as migration corridors, if they were present seasonally for feeding or even breeding, Introduction or if at least some of them continuously inhabited these environments (see also, Lopez-Martínez, Moratalla, and Sanz, Intrinsically, the nature of vertebrate tracks deals with in 2000; Marty, 2008; Myers and Fiorillo, 2009; Diedrich, 2011; situ trace fossils that can be unambiguously linked to the Fricke, Hencecroth, and Hoerner, 2011; Marty and Meyer, environment in which they were left (Lockley 1991, 1998). 2013; Rowe et al., 2013). Particular depositional environments are interpreted as hav- This contribution reviews the Jurassic and Cretaceous ing been preferred by specific groups of dinosaurs and other dinosaur track record of shallow marine carbonate deposi- vertebrates, thus giving rise to the tetrapod ichnofacies con- tional systems, briefly “carbonate platforms,” and presents cept (Lockley et al., 1994; Lockley, Hunt, and Meyer, 1994; a database (http://dx.doi.org/10.6084/m9.figshare.1348706) Hunt and Lucas, 2007; Lockley, 2007). that was built for this purpose. This database should serve Dinosaur tracks are often recorded in coastal, shallow as a reference for future (statistical) analyses based on this marine environments of Europe and North America (e.g., coastal dinosaur track record, and it may also be completed Farlow, Pittman, and Hawthorne, 1989; Farlow, 1992; Dalla with complementary data from other (siliciclastic) environ- Vecchia, 1994, 1998; Lockley et al., 1994; Meyer and Pittman, ments and time periods. For the time being, Triassic dino- 1994; Avanzini et al., 1997, 2006; Kvale et al., 2001; Marty saur track occurrences, including those usually labeled as

381

DinoTracks.indb 381 1/22/16 11:23 AM “dinosauroid” or “dinosauromorph” tracks (e.g., Marsicano, warm (sub)tropical climate, and low input of sediment from Domnanovich, and Mancuso, 2007; Brusatte, Niedz´wiedzki, the continent. Several tracksites are from peripheral conti- and Butler, 2011; Kubo and Kubo, 2013; Niedz´wiedzki, Bru- nental (e.g., Texas, United States) or intracontinental (low- satte, and Butler, 2013), are not included as they are often land coastal, e.g., Wyoming, United States) settings, whereas not easy to distinguish from other nondinosaurian archo- others are from intraoceanic (e.g., Southern Italy, Croatia) saur footprints or are hard to refer to one dinosaur group or and epeiric (Jura Mountains, Switzerland) carbonate plat- another (i.e., Saurischia, Ornithischia). However, recently forms located far away from emergent mainland. Accord- more and more Triassic dinosauroid and dinosaur ichnites ingly, the database includes tracksites from a range of differ- are discovered in shallow marine carbonate deposits of Cen- ent marginal marine environmental settings, and at a single tral Europe (e.g., Gand and Demathieu, 2005) and notably tracksite, several subenvironments of proximal to distal peri- the Alps (e.g., Avanzini, 2002; Furrer and Lozza, 2008; Meyer tidal facies (e.g., sabkha, algal marsh, mudflat, tidal channel) et al., 2009, 2010, 2013; Bernardi et al., 2013), and once these may also be observed in close lateral and/or vertical vicinity. tracks are better understood, they will certainly become an However, a detailed description of the spatial distribution of important addition to the database. shallow water carbonate depositional environments during the Jurassic and the Cretaceous is outside the scope of this Materials and Methods chapter, and the reader is referred to the sedimentological literature on this topic (e.g., Hardie and Shinn, 1986; Tucker et The database was built on data gathered from the literature, al., 1990; Demicco and Hardie, 1994; Wright and Burchette, and to the best of our knowledge, all available reports of di- 1996; Schlager, 2002; Kiessling, Flügel, and Golonka, 2003). nosaur tracksites from carbonate-dominated shallow marine In track-rich regions, numerous tracksites may occur at deposits of Jurassic and Cretaceous age have been included. short distances from one another and even belong to the In total, 131 publications published between 1917 and 2013 are same stratigraphic interval (e.g., Marty et al., 2007). As we considered in the database. Apart from some data of track- will show in more detail later, peaks in the number of tracksites that the authors have studied themselves, the database sites from a given region (Fig. 20.1) may be a function of only includes published data. Not all of this published data intensive studying (sampling bias). The quality of the fossil can still be verified today, as dinosaur tracks are generally (ichnological) record or the discrepancy in the number of not collected and deposited in a museum collection. This entries and amount of data from different areas and hence is related to the fact that dinosaur tracksites are generally the quality of the biological signal may depend on the pres- large, form part of the landscape, and thus cannot easily be ence of research institutions close to the study areas, number relocated to museum collections. Moreover, most tracksites of specialists for selected branches (ichnology, in this case), are not protected against erosion and disintegration, and for outcrop area and territorial accessibility of potential data- this reason, some previously available data may today be dis- yielding rock units, and so on (e.g., Alroy et al., 2001; Smith, torted, destroyed, or not be accessible. 2001; Benson and Upchurch, 2012; Dunhill, 2012; Dunhill et In order to avoid subjective filtering at the data-entry al., 2012; Smith and Benson, 2013). A good example are the level, the basic unit adopted in the database is the tracksite, systematic track excavations carried out prior to the construc- which refers to a natural, in situ area composed of one or tion of Highway A16 in northwest Switzerland (e.g., Marty et several bedding planes (levels, horizons) with tracks. Accord- al., 2003, 2007; Marty, 2008), which have led to the documen- ingly, at a given tracksite, tracks may not only occur in just tation of a total of nearly 14,000 dinosaur tracks including 656 one single ichnoassemblage but at multiple, superimposed trackways from six tracksites, which are located geographi- levels, and true tracks may also be associated on a single sur- cally very close one to another (within a few kilometers) and face with undertracks and/or overtracks (Marty, 2008). Even which constitute one of the most abundant data entries in though the entire track-bearing interval of a given tracksite the database. may usually be considered as penecontemporaneous in geological terms, such intervals may represent up to several tens Database Structure of thousands of years (Marty, 2008), or one to several elemen- tary sequences in the concept of cyclostratigraphy (Strasser Chrono- and Lithostratigraphy et al., 1999). All tracksites considered in the database share some ba- The three categories “epoch,” “stage,” and “formation” are sic environmental parameters such as carbonate deposition, related to the age of the tracksite. For some localities, no close vicinity to the marine depositional paleoenvironments, precise dating is available, and two or more successive stages

382 Simone D’Orazi Porchetti et. al

DinoTracks.indb 382 1/22/16 11:23 AM bracket the best available range estimate. Where a formal even >>, << (much greater than, much lesser than) in case lithostratigraphic designation is lacking, “no data” figures one category is much more abundant than another. In the as an entry. case of monotypic tracksites (in terms of dietary type), H (or C) = 100% is reported. Geographical and General Setting Track sizes The five categories“ tracksite name,” “number of levels,” “locality,” country,” and “outcrop area” provide general Four categories named “average dimensions (pes track information about the tracksite. “Number of levels” indi- length) theropods/sauropods/ornithopods/other ornithischi- cates how many different levels with dinosaur tracks are ans” comprise information about track size. As all the data present at any given tracksite. “Locality” is specified by the is derived from literature, we assume that the pes length name of a nearby locality or a toponym, as reported in the is usually measured from the rear of the foot to the tip of original publication. The average dimension of the out- the most protruding toe (which is generally digit III), but crop area is expressed in square meters, but in many cases, there are many exceptions to this general rule, depending there is no precise information available about the tracksite on the type of track (e.g., sauropod or tridactyl tracks) and dimensions. on track preservation (e.g., completeness). For instance, most of the tridactyl tracks described from the Middle Jurassic of References Wyoming by Kvale et al. (2001) are preserved as isolated toe prints. In this case, we only reported the length of digit III This category comprises the references that were used as data from some selected tracks (accompanied by an asterisk in sources. A specific bibliography for the database is included. the database). Furthermore, many tracks are not true tracks but undertracks, overtracks, or penetrative tracks, and this Tracks, Trackways, and Trackmakers may distort the original dimensions and shape of the foot (Milàn and Bromley, 2006, 2008). The uniformity of the Related information is provided in nine categories. The cat- available measurements is another general issue for our da- egory “total number of tracks and trackways” reports the tabase. Many papers report discrete measurements for each total number of tracks or trackways at a single tracksite. The track encountered at a single tracksite. In other cases, the category “inferred trackmaker(s)” is the trackmaker attribu- original work includes only mean values, with no information based on the identification given by original authors. tion on the distribution of these parameters. Moreover, some When conflicting attributions are found in different papers papers give no absolute measurements but only qualitative on the same material, we report the attribution with the larg- assessment (e.g., small, medium, large). In some cases, no est consensus (see Deltapodus attribution). If no consensus is data were available at all. In the “track size” column, we found on the attribution, we report conflicting hypotheses in have tried to mitigate these different approaches as follows: the supplementary notes, supplying the original attribution if no measurements were available, the corresponding box in the “trackmakers” column. has no entry (e.g., no data). When no numerical data were The three categories “herbivores (H),” “carnivores (C),” available, the original qualitative term was reported (small or and “ratio H/C” are related to the diet of the inferred track- large, for instance). If an average length is the only available makers and the ratio between these two categories. All information, it is reported in the appropriate box. In several nonavian dinosaurs are divided in two broad ecologic cat- cases, a size range is reported. When the original article egories: “herbivores,” including Sauropodomorpha and all gave a complete list of measurements for several footprints, ornithischians, and “carnivores,” referring to all Theropoda. we calculated a mean value and added the minimum and We consider the Theropoda as largely carnivorous, follow- maximum values, as appropriate. ing the consensus on most groups of nonavian theropods (Fastovsky and Smith, 2004), even though some of them, Depositional Setting especially among coelurosaurs, are considered as herbivores (Weishampel and Norman, 1989; Barrett, 2005; Butler and This category comprises all relevant information (e.g., su- Barrett, 2008; Zanno and Makovicky, 2011). The ratio (H/C) pratidal/intertidal flat, sabkha) about the depositional pa- is expressed exclusively in relative terms. H = C is reported leoenvironment based on the indicated references. However, when these two categories are equally abundant. We adopted original works are not always accompanied by a detailed the mathematical symbols >, < (greater than, lesser than), or analysis of the geologic framework and often only provide

A Review of the Dinosaur Track Record 383

DinoTracks.indb 383 1/22/16 11:23 AM general information such as “shallow marine depositional nonavian dinosaurs. This creates a severe limiting factor: the environment.” virtual absence of present-day possibilities of comparison on this topic. Supplementary Notes Taphonomy is a further relevant issue that must be considered. Chemical-physical and microbiological factors may A further column is dedicated to supplementary notes. In- play a strong role in early consolidation of shallow marine formation is reported here that cannot be addressed in previ- carbonate deposits, thus helping to preserve vertebrate bio- ous categories, such as the current conditions of the outcrop turbation. Microbial mats are especially relevant in this pro- (destroyed, eroded) or specific information on the history of cess (Marty, Strasser, and Meyer, 2009) and may have acted the sites. like flypaper for dinosaur tracks. Early dolomitization of the trampled substrate has also been suggested as an important Discussion preservational variable (Avanzini et al., 1997). However, dinosaur tracks are also abundant in siliciclastic coastal plain Dinosaur tracksites used for this study belong to distinct facies, thus suggesting the need for further clastic-carbonate regions, but these may also have changed their paleogeo- facies comparisons before the preservational role of micro- graphic position and environmental setting between the Ju- bial mats is fully evaluated and understood. rassic and the Cretaceous. North America yields tracks from lowlands bordering an epicontinental sea (Sundance Forma- Trackmaker Attribution tion, Wyoming) during the Middle Jurassic and from coastal environments facing the Western Interior Seaway (Glen Rose Ascribing tridactyl tracks to either theropods or basal orni- Formation, Texas) during the Early Cretaceous. Many track- thopods always involves a certain degree of uncertainty (e.g., sites are from the so-called European archipelago (e.g., Ibe- Farlow et al., 2006; Castanera et al., 2013; Hübner, 2016). To rian Massif, Armorican Massif, and Massif Central), where some extent, uncertainties in trackmaker attribution may sea-level rise and fall (either linked to seafloor spreading or to also affect the tracks of large and obligate quadrupedal herbi- climatic changes) continuously and strongly modified the pa- vores (saurischians vs. ornithischians). A good example is the leogeographical and paleoenvironmental setting throughout case of Deltapodus, an ichnotaxon first attributed to a sauro- the Mesozoic. A third region encompasses the Periadriatic pod (Whyte and Romano, 1994) but today is considered as area where an alternation of carbonate platforms and pe- a stegosaurian ichnotaxon by most ichnologists (Whyte and lagic basins formed a complex patchwork during most of the Romano, 1994, 2001; Lockley and Meyer, 2000; Cobos et al., Jurassic and the Cretaceous. In this scenario, track-bearing 2010; Li et al., 2012; Xing et al., 2013, and references therein). localities are from carbonate platform environments (Trento This ambiguity in trackmaker attribution is potentially an Platform, Apulian Platform, Apennine Platform, Adriatic- intrinsic issue in ichnology and is also linked to preserva- Dinaric Platform). Central Asia has yielded large dinosaur tional quality, as a result of dynamic interactions between tracksites from Turkmenistan and Uzbekistan, representing anatomical, kinematic, and sedimentological factors (e.g., lowland shorelines of a Late Jurassic large epicontinental sea. Padian and Olsen, 1984; Gatesy, 2003). From Africa, a good, new sample comes from Madagascar, where tracks are found on the western coast of the Eastern Relative Timing of Trackway Registration Gondwana continent. Coastal deposits from northern Africa and Assessing Trackmaker Frequency (e.g., Morocco, Tunisia) in a mixed carbonate and siliciclastic facies are excluded because they do not match the paleoen- Determining the relative timing of trackway registration in vironmental requirements of this review. To the best of our a given ichnoassemblage is a difficult task. A (parallel) series knowledge, the Far East and Australia are devoid of dinosaur of trackways each with similar track dimensions may either tracks from shallow marine carbonate environments. result from a single individual passing several times or from As opposed to a considerable amount of data on dino- several individuals of similar size passing together in a group saur tracks from shallow marine carbonates, reports of ex- or a herd (Ostrom, 1972; Lockley, 1986, 1989; Lockley and tant large tetrapods within marginal marine environments Hunt, 1995; Myers and Fiorillo, 2009). However, if compared are rather rare. This may depend on two factors: first, there to many body fossil deposits, the temporal window for track- may be no exact homology between Mesozoic and modern making and preservation of tracks is comparatively short and carbonate depositional settings, especially for large epeiric may vary between a few hours and up to several days, weeks, seas that are absent today (Hallam, 1992; Harries, 2009), and or months (Cohen et al., 1993; Marty, 2008; Marty, Strasser, second, there might be no extant ecological homologues for and Meyer, 2009). Trackways are often used in census studies

384 Simone D’Orazi Porchetti et. al

DinoTracks.indb 384 1/22/16 11:23 AM as the unit for counting individuals on each individual track- 180

site (Lockley, 1998), and the same approach was adopted in 158 this work. Unfortunately, identifying trackways on a heavily 160

dinoturbated surface is sometimes impossible because of too 140 many interfering tracks. An alternative possibility may be a classification based on a tracks/area ratio, the dinoturbation 120 index (Lockley and Conrad, 1989), but this ratio is generally 100 impossible to assess if based on literature data only. Some- 94 times tracksites are not located very conveniently or, when cleaned up in order to expose tracks, the travel direction may 80 be chosen as the rationale under which a site is expanded 60 by excavation of overlying rocks. By doing so, an outstand- Number of tracksites ing long trackway may be the result, but another important 40 pattern (e.g., several parallel trackways, herding) may not be 30 detected (Marty et al., 2012). 20

0 Jurassic and Cretaceous Tracksites THEROPODS SAUROPODS ORNITHISCHIANS at a Glance: Distribution over Time, Space, and Taxonomic Groups 20.2. Total occurrences for each dinosaur group. Theropods are the most abundant, with 158 entries (left column), and sauropods follow To the best of our knowledge, a total of 212 dinosaur tracksites with 94 (central column). Ornithischians are poorly represented in the have been reported from Jurassic and Cretaceous shallow record with respect to Saurischia. A total of 30 entries (right column) is the sum of 19 entries for ornithopods (dark lower block), 8 thyreopho- marine carbonate systems worldwide. Almost every stage of rans (middle block), and 3 for other ornithischians (pale upper block). the Jurassic and the Cretaceous includes occurrences of dinosaur tracks in shallow marine carbonate systems. Excep- 212 known individual tracksites is mainly a matter of what tions are the Pliensbachian and Toarcian (Early Jurassic), the kind of question is investigated. On the other hand, abun- Aalenian and Callovian (Middle Jurassic), and the Santonian dances of dinosaur groups in the database are easily visual- and Campanian stages (Late Cretaceous). ized when data from different outcrops (tracksites) are sepa- Some 129 tracksites are known from the Jurassic and 83 rated, so this approach is followed herein (Fig. 20.2). In this from the Cretaceous. However, this number is somewhat regard, it can be argued that we must utilize the raw data on misleading, as some formations contain many individual tracksites and not infer (interject our own bias) by conclud- tracksites all located approximately at the same stratigraphic ing that some formations are “oversampled” when they may level and yielding an almost identical ichnofauna. There in fact genuinely represent depositional systems where track are also some formations with several tracksites at different registration is naturally greater due to the intrinsic dynamics stratigraphic levels, but the track-bearing layers still are very that create the sedimentary record: in other words, there is no close in space and time and are similar in composition. For objective way to pool some data sets and not others. instance, almost all known dinosaur tracksites from the Early Nevertheless, the geographic distribution of the track- Jurassic are from the Causses (France) and from the Province sites depends on the outcrop area of potential data-yielding of Trento (Northern Italy), whereas half of all Middle Jurassic rock units (i.e., shallow marine deposits), but it also strongly entries belong to the Bemaraha Formation in western Mada- reflects the intensity with which different geographic areas gascar. Not less than 49 tracksites (23% of our total database) have been studied (sampling bias). Of the 212 tracksites, 130 have been reported from the Glen Rose Formation of Texas are located in Europe, 59 in North America (United States (Albian, Early Cretaceous), which is the most prolific forma- only), 13 localities in Africa (Madagascar only), 7 localities tion in our database, followed by the Reuchenette Forma- in Asia (including the Arabian Peninsula), and 3 tracksites in tion in northwest Switzerland (Kimmeridgian, Late Jurassic), South America. No occurrences are known from Australia which yielded 25 individual tracksites. If we consider track- or Antarctica to date. The United States (59 tracksites) is the sites from the same geological formation as a single entry, richest country in terms of single track-bearing localities, 49 occurrences remain worldwide, with the bulk of them (31 followed by France (46) and Switzerland (30). In all these occurrences, or 63%) being quite evenly distributed over the countries, potential data-yielding rock units crop out over Late Jurassic and Early Cretaceous. Referring for statistical large areas and have been studied intensively since at least purposes to either these 49 occurrences by formation or all the 1980s.

A Review of the Dinosaur Track Record 385

DinoTracks.indb 385 1/22/16 11:23 AM 70 70 60 60 50 EARLY JURASSIC 50 40 MIDDLE JURASSIC 30 LATE JURASSIC 40 20 EARLY CRETACEOUS THEROPODS

Number of tracksites LATE CRETACEOUS 30 10 SAUROPODS ORNITHOPODS 0 20

Number of tracksites of Number THYREOPHORANS T. 10 ORNITHISCHIANS INDET. UROPODS THEROPODS SA INDE ORNITHOPODS THYREOPHORANSORNITHISCHIANS 0

CEOUS CEOUS TA A 20.3. Distribution through time for each trackmaker group. The y-axis is ET Y JURASSIC TE JURASSIC LA Y CRE for the number of tracksites, whereas trackmaker groups are reported EARL TE CR MIDDLE JURASSIC on the x-axis. Each column has a different color depending on the time EARL LA sampled (see legend on the right). Theropods are the most common group, except for the Late Jurassic, where sauropod tracks became dominant. 20.4. Distribution of trackmaker groups over time (epochs on the Theropod track occurrences have a peak in the Early Cretaceous, but x-axis). Theropods are well represented throughout the Jurassic and the they fall down in the next stage (Late Cretaceous), where the record is Cretaceous. The absolute peak in theropod tracks is in the Early Cretaceous, generally low for all groups. Ornithopods are best represented in the possibly in correspondence with highly prolific track-bearing forma- Early Cretaceous, whereas thyreophorans and other ornithischians have tions. Sauropods are dominant trackmakers in the Late Jurassic, being a very scarce record throughout the Jurassic and the Cretaceous. less represented before and after this epoch. Ornithopod tracks appear in the Late Jurassic with few entries until the end of the Cretaceous. Of the 212 tracksites, 158 yield tridactyl tracks assigned to Thyreophoran and generic ornithischian tracks appear as soon as the Early Jurassic, but they are very poorly represented in the record. theropods. This is the dominant trackmaker group in almost every time slice of the Jurassic and Cretaceous periods except at 17 localities, theropods dominate in number. At 2 localities, for the Late Jurassic where they are outnumbered by sauro- herbivores and carnivores were equally abundant. No data pods. A total of 103 tracksites yielded theropod tracks only. were available for 8 tracksites (Fig. 20.4). Overall, sauropods are the second most common group, When the data is sorted by lithostratigraphic units (forma- and they outnumber all other ornithischian herbivores tions), the results are slightly different, because of fewer “raw in each of the five time slices we have considered (Early, data issues” (i.e., analysis based on the number of tracksites). Middle, and Late Jurassic, and Early and Late Cretaceous). As stated earlier, this is related to the presence of multiple Among ornithischians, ornithopods are the most common tracksites from the same formation that may be geographi- group, with 19 entries, although almost half of them (9) are cally located close to each other and thus better be con- reported as dubious, presumably due to the danger of confus- sidered as one entry only (data sorted by lithostratigraphic ing them with theropod tracks. Thyreophorans are a rare but unit). The general dominance of saurischians over ornith- recurrent element, with eight entries quite evenly distributed ischians is confirmed for all time units, but an important over the considered time interval. Three sites, all from the reduction in the relative abundance of each taxonomic group Early Jurassic of France, have yielded tracks attributed to (i.e., theropods, sauropods, other ornithischians) is detected. undetermined ornithischians (Fig. 20.3). For instance, during the Early Jurassic, the count of “other There are 55 localities where herbivores and carnivores co- ornithischian” is equal to theropods and greater than sauro- occur, of which 36 yield a theropod/sauropod association and pods (Fig. 20.5). This is slightly in contrast with the results 8 a theropod/ornithischian association. Co-occurrences of obtained from the investigation based on tracksites. theropods with both sauropods and ornithischians have only been reported from 11 tracksites. Of 212 tracksites, 44 contain sauropod tracks only. Herbivores-only associations with both Observation on the Brontopodus Ichnofacies Concept saurischian and ornithischian dinosaurs are known from 3 localities only. At 14 tracksites out of 36, where sauropods and Lockley, Hunt, and Meyer (1994) first stated the presence theropods co-occur, sauropods are more abundant than the- of recurrent ichnocoenoses with sauropod (originally wide- ropods. This latter group is dominant at 12 tracksites, whereas gauge sauropods, i.e., Brontopodus and cf. Brontopodus) and sauropods and theropods are equally abundant at 2 localities large theropod tracks in shallow marine and lacustrine car- only. The remaining 10 tracksites yield no data on the H/C ra- bonate depositional environments. Most of the original data tio. Theropods co-occur with sauropods and other herbivores set (Lockley, Hunt, and Meyer, 1994) was recovered from (e.g., ornithopods and other ornithischians) at 47 localities. North America (Texas) and Europe (Switzerland and Portu- We observed a majority of herbivores at 20 tracksites, whereas gal), but occurrences from South America, Eastern Asia, and

386 Simone D’Orazi Porchetti et. al

DinoTracks.indb 386 1/22/16 11:23 AM Africa were also considered. The original definition of the 14 Brontopodus ichnofacies was later widened by including ichnoassemblages yielding Parabrontopodus and similar forms, 10 FM YIELDING THEROPODS FM YIELDING SAUROPODS

with dominance or general abundance of sauropod tracks 6 FM YIELDING ORNITHOPODS (e.g., Meyer and Hauser, 1994; Meyer and Pittman, 1994; FM YIELDING OTHER ORNITHISCHIANS Lockley and Meyer, 2000). In later revisions of the ichnofa- Number of formations 2

cies concept, Hunt and Lucas (2007) proposed conceptual S

CEOUS CEOU redefinition of the categories and of their ranking. In their JURASSIC JURASSIC TA TA YJURASSIC TE CRE CRE LA Y EARL TE interpretation, all tetrapod ichnofacies defined prior to 2005 MIDDLE EARL LA should be considered as ichnocoenoses, whereas the ichnofa-

cies label was used to characterize only five very broadly de- 20.5. Occurrences of trackmaker groups over the Jurassic and the fined universal or archetypal ichnofacies. For example, their Cretaceous, sampled by formations (FM). Differences in relative abundance (i.e., theropods vs. sauropods) are reduced if compared to the Brontopodus ichnofacies actually included clastic as well as graphs based on the tracksite unit (compare to Figs. 20.3 and 20.4). carbonate facies-related ichnoassemblages (their ichnocoe- A total of 14 formations are reported on the y-axis, time is on the noses, or ichnofacies of other workers) specifically stating x-axis. Saurischians are dominant over ornithischians in all time units, but a reduction in the relative abundance of each taxonomic group is that they were usually herbivore dominated with a low (about evident. For instance, during the Early Jurassic, the number of “other 10%) number of carnivores and high (4–8) ichnogeneric di- ornithischians” equals theropods and is greater than sauropods. versity. This definition is not in the least consistent with the carbonate platform ichnofaunas described here, which are almost as common as herbivores and, in general, localities heavily theropod and sauropod dominated (usually only 2 where H>>C are extremely rare. In any case, our data gives ichnogenera) as is characteristic of the Brontopodus ichnofa- no numerical quantification on the relative abundance of cies as originally defined (Lockley, Hunt, and Meyer, 1994). one group with respect to another. In x-axis subsequent works, Lockley (2007) and Lockley and In the light of our investigation, it is of interest to observe Gierlinski (2014) reconsidered both these approaches, dis- that Hunt and Lucas (2007) assumed for the Brontopodus tinguishing an old and a new paradigm for the ichnofacies ichnocoenosis a pattern of abundance for theropod tracks concept, referring the former to the definition of Lockley, below the threshold of 10%. Our results do not confirm this Hunt, and Meyer (1994) and the latter to that of Hunt and conclusion and in fact theropod footprints are usually abun- Lucas (2007). A recent evaluation on the meaning and use- dant also when associated to herbivores tracks. Our data are fulness of vertebrate ichnofacies was presented by Santi and more, if not fully, in agreement with the conclusions of Lock- Nicosia (2008). ley, Hunt, and Meyer (1994) as far as theropod and sauropod As we limited our analysis to a specific depositional set- tracks, when in co-occurrence, show near-equal abundance. ting within marine carbonate systems, we can test whether However, any further comment on the Brontopodus ich- correlations between a depositional environment and the nofacies must take into account those coastal environments vertebrate track record occur and whether any pattern is re- where mixed and siliciclastic deposition took place. The trievable. We can primarily state that saurischian dinosaur preference of one group for a specific environment must tracks are dominant in shallow marine/coastal deposits, and therefore be tested on a more complete data set. theropod tracks are the most abundant feature. Consequently, the question arises whether the Brontopo- Conclusions dus ichnofacies is a specific subset in the larger scenario of vertebrate tracks associated to shallow marine carbonate fa- Our data show that theropod tracks are dominant in shallow cies. The difficulty to assign some dinosaur track assemblage marine carbonate systems. Actually, theropods appear to be to one or another of the meanings of the Brontopodus ichno- over-represented if compared with herbivores, the bulk being facies was stated by Marty (2008) who noticed that the Tran- made up by small to medium-sized animals, and this may sjurane ichnocoenosis was different from previously defined be related to a higher activity level of (small) carnivores than ichnocoenoses, and specifically from the Brontopodus ichno- herbivores. coenosis as defined by Hunt and Lucas (2007). Marty (2008) Sauropod footprints are the second most abundant ich- interpreted the Transjurane as a new ichnocoenosis of the nites. There is a strong affinity of some sauropod groups to Brontopodus ichnofacies (in the definition of Lockley, 2007). coastal environments. Limiting our analysis to the ichnocoenoses where thero- Ornithischians are relatively rare in comparison with sau- pods are associated to sauropods (exclusively, or with a small rischians. Most ornithischians appear to have entered these number of ornithischians), we can state that carnivores are environments only occasionally.

A Review of the Dinosaur Track Record 387

DinoTracks.indb 387 1/22/16 11:23 AM The reasons for these habitat preferences are manifold A further step would be the statistical elaboration of the data and might be related to differences in physiology, different set, as far as a greater amount of data becomes available. behaviors, and ecological/trophic needs that were more or less adapted for these coastal environments. In order to test Acknowledgments our conclusions, in the future, these data should be plotted against the general pattern of distribution based on the body M. Avanzini (Museo delle Scienze, Trento, Italy), A. Cobos fossil record. This process might take into account the esti- (Fundacion Conjunto Paleontologico de Teruel-Dinopolis, mated abundance of these groups in terms of biomass, and Spain), C. A. Meyer (Naturhistorisches Museum Basel, Swit- their relative distribution during the Jurassic and the Cre- zerland), G. Gierlinski (Polish Geological Institute, War- taceous. The relative scarcity of data we recorded from the saw, Poland), I. de S. Carvalho (University of Rio de Janeiro, Late Cretaceous may be critical in this process, and the body Brazil) are kindly acknowledged for bibliographic input and fossil record could help drawing a more complete scenario. comments. M. G. Lockley and an anonymous reviewer are This work is a first step toward the compilation of a data- acknowledged for their insightful comments and suggestions base on the global distribution of dinosaur tracks in coastal on an earlier version of the manuscript. We would also like to environments, including those in widespread siliciclastic fa- thank the editors for their constant and valuable assistance. cies (Lockley et al., 2006, 2010, 2014; McCrea et al., 2014). In S. D’Orazi Porchetti’s research is supported by a Fundação the future, a possible extension to inland depositional set- de Amparo à Pesquisa do Estado de São Paulo (FAPESP) post- tings is envisaged. This database should be a core for further doctoral grant (process number 2013/01930-1). implementation, and should also include the Triassic period.

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