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A. Type specimen of Batrachopus (B. deweyi) from New England (traced from the type specimen 26/5 in the Pratt Museum, Amherst College) with Batrachopus from the Causses region of France for comparison.  

A. Type specimen of Batrachopus (B. deweyi) from New England (traced from the type specimen 26/5 in the Pratt Museum, Amherst College) with Batrachopus from the Causses region of France for comparison.  

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Trackways described as Batrachopus (Batrachopodidae Lull, 190426. Lull , R. S. 1904. Fossil footprints of the Jura-Trias of North America. Memoirs of the Boston Society of Natural History, 5: 461–557. View all references) from the Lower Jurassic of Europe are rare and in some cases different from the type trackways from North America. Differences...

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Context 1
... of which are evidently extra-morphological vari- ants. This is the only non-dinosaurian track considered by Haubold (1986, fig. 15.10) to be characteristic of Lower Jurassic assemblages. The only other non-dinosaurian track, from the 65 classic New England faunas, that has recently received serious scientific attention is Antipus (Coombs, 1996: Fig. 2 herein) though this may be a synonym of Batrachopous (Olsen and Padian, 1986). It is also similar to the trackway named Sten- odactylus by Hitchcock (1858) and renamed Sustenodactylus 70 by Lull (1953) (see Fig. 2), which according to Olsen and Padian (1986) is probably one of many synonyms of Batracho- pus. As reported elsewhere in ...
Context 2
... track, from the 65 classic New England faunas, that has recently received serious scientific attention is Antipus (Coombs, 1996: Fig. 2 herein) though this may be a synonym of Batrachopous (Olsen and Padian, 1986). It is also similar to the trackway named Sten- odactylus by Hitchcock (1858) and renamed Sustenodactylus 70 by Lull (1953) (see Fig. 2), which according to Olsen and Padian (1986) is probably one of many synonyms of Batracho- pus. As reported elsewhere in this volume ( Lockley et ...
Context 3
... to Batrachopus (tetradactyl and pentadactyl, re- spectively), Antipus by contrast to Batrachopus has much more 150 slender, tapering and widely-divaricating digits, that terminate in narrow claw impressions. In addition to being far more slen- der, manus digit impressions seem to be much longer in Anti- pus than they are in Batrachopus (compare Figs. 1A and 2A). The digits were evidently quite flexible as indicated by strong 155 curvature that gives the type ichnospecies its name: A. flexil- oquus. Antipus is similar to Batrachopus in having a strong outward rotation to the pes (34 • N = 4, measured between the trackway midline and the axis of pes digit III). Another feature not mentioned by ...
Context 4
... gest that Batrachopus has a relatively narrow pes, and in two of the French examples, a significantly smaller manus (greater het- eropody). The length of the Batrachopus pes is much longer in 495 some tracks than others due to a faintly impressed heel trace. For example in Fig. 1A, the type specimen, also illustrated by Olsen and Padian (1986, fig. 20.1) the pes may be twice as long as wide. However in some cases the heel is not impressed, suggesting an alternation between plantigrade and semi-digitigrade stance. ...

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Citations

... Crocodylomorph tracks are well known from the Lower Jurassic to the Cenozoic (Lockley and Meyer, 2004;Klein and Lucas, 2010;Lockley et al., 2010b) with three main crocodylomorph ichnotaxa being the most significant in terms of the number of reports: Batrachopus (mainly Lower Jurassic-Cretaceous), Crocodylopodus (mainly Upper Jurassic-Cretaceous) and Hatcherichnus (mainly Upper Jurassic-Cretaceous, see Kim et al., 2020;Lockley et al., 2020;Masrour et al., 2020 and references therein). The ichnotaxon Crocodylopodus meijidei from the Huérteles Formation (Soria, Spain) is the type of the ichnogenus Crocodylopodus and is thus a key ichnotaxa during the Mesozoic (Fuentes Vidarte and Meijide Calvo, 2001). ...
... fig. 1 and fig. A) and is the holotype according to Lockley and Meyer (2004). The specimen (Fig. 3) includes four sets of coupled manual and pedal tracks (Fuentes Vidarte and Meijide Calvo, 2001, also draw one isolated manual print partially preserved at the beginning of the trackway that is not clearly identified here). ...
... The estimated glenoacetabular distances range from 11.5 to 14.2 cm. Remarks- Lockley and Meyer (2004) noted that three holotypes (Rastro A, B, and C = MNS2002/96/2bis, MNS2003/92/8a, and MNS2002/96/4, respectively) were designated in the original description by Fuentes Vidarte and Meijide Calvo (2001) and that such a procedure is not permitted by the ICZN (1999), so they selected "Rastro A" as the holotype and designated "Rastro B" and "Rastro C" as paratypes. According to the ICZN (1999), however, the holotype "can only be fixed in the original publication and by the original author" (Article 73.1.3). ...
... Whereas the Triassic record of crocodylomorph tracks is restricted to North America and Europe (Klein & Lucas, 2010), the Jurassic crocodyliform track record is more extensive (Lockley & Meyer, 2004), spanning North America (Foster & Lockley, 1997;Lockley & Foster, 2010;Lockley, Kirkland & Milner, 2004;Lockley et al., 2018b), Europe ( 2014; Simpson et al., 2010), Europe (Segura et al., 2016;Vila et al., 2015), and Asia (Lee et al., 2019). Among Mesozoic crocodylomorph/crocodyliform tracks, the crocodyliform tracks from the Snake Creek Tracksite most closely correspond with those assigned to Hatcherichnus (Foster & Lockley, 1997;. ...
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The Upper Cretaceous 'upper' Winton Formation of Queensland, Australia is world famous for hosting Dinosaur Stampede National Monument at Lark Quarry Conservation Park, a somewhat controversial tracksite that preserves thousands of tridactyl dinosaur tracks attributed to ornithopods and theropods. Herein, we describe the Snake Creek Tracksite, a new vertebrate ichnoassemblage from the 'upper' Winton Formation, originally situated on Karoola Station but now relocated to the Australian Age of Dinosaurs Museum of Natural History. This site preserves the first sauropod tracks reported from eastern Australia, a small number of theropod and ornithopod tracks, the first fossilised crocodyliform and ?turtle tracks reported from Australia, and possible lungfish and actinopterygian feeding traces. The sauropod trackways are wide-gauge, with manus tracks bearing an ungual impression on digit I, and anteriorly tapered pes tracks with straight or concave forward posterior margins. These tracks support the hypothesis that at least one sauropod taxon from the 'upper' Winton Formation retained a pollex claw (previously hypothesised for Diamantinasaurus matildae based on body fossils). Many of the crocodyliform trackways indicate underwater walking. The Snake Creek Tracksite reconciles the sauropod-, crocodyliform-, turtle-, and lungfish-dominated body fossil record of the 'upper' Winton Formation with its heretofore ornithopod-and theropod-dominated ichnofossil record.
... As seen in the tables, the two most cited ichnogenera (Batrachopus and Crocodylopodus) do not show a definite pattern relative to the width of the trackway. Given the variability of the trackway data collected in Table 4, we think like Lockley and Meyer (2004), that the width of the trackway is not a fundamental character of ichnotaxonomic value. The width of the trackway is strongly linked to the nature of the substrate, water level and the type of progression (including variation of the travel speed) of the "crocodile". ...
... We consider that the following character set is determinative of ichnites of the ichnofamily Batrachopidae (see Lull, 1904; amended by Lockley and Meyer, 2004): quadruped trackway, narrow; the manus print is fan-shaped, digitigrade, pentadactyl, pointed and separated digits with strong divergence; "Pes with four digits, generally dinosauroid in form though plantigrade" (Lull, 1904, p. 482), triangular, larger than the manus, with pointed, relatively thin digits. The relative length of the digits follows the formula III > II ≈ IV > I. ...
... There are opinions regarding the prevention of new names for this type of ichnite, and on the defined species for them. Thus, for example, Lockley and Meyer (2004) point out that "The status of Batrachopus … remains uncertain and ambiguous", Le Loeuff et al. (2010) indicate that "A detailed review of Batrachopus is needed". When doing the comparative study with the existing species, we see that many of them have been made on a single specimen sometimes deformed and without guarantees that they are true prints or stamps, that is to say without taking into account the recommendations of Peabody (1955), Sarjeant (1994) or Romero-Molina et al. (2003). ...
Article
In this work we present the description and analysis of a palaeoichnological site (7.9TAG) that only contains crocodile tracks. The site contains well preserved and abundant crocodilian manus and pes prints; there are no belly and/or tail traces. The crocodyle tracks have been mapped and the biomorphic and morphometric characters of the tracks and trackways have been studied. Between the digit traces of the pes prints (not in manus prints) there is a graded depression that indicates the presence of a foot webbing. The number of crocodilian footprints (93) and trackways (8) of the site is sufficient to define, and to support with confidence the data shown in the article. As consequence a crocodiliform ichnogenus, Batrachopus has been defined, characterized by the presence of a trace of a probably interdigital membrane in the pes prints, the trace of a bulge on toe V, the lack of tail traces, and the very narrow gauge trackway. A comparative study is done with all the fossil crocodile footprints described so far. The 7.9TAG footprints printed on sediments of Middle?-Upper Jurassic age (Isli Formation) are the first Batrachopus ichnites described both from this age and in the African continent.
... General tracks from the level 2 surfaces were shallower with clearly defined digit and digital pad impressions (Figs. [3][4][5][6], as well as transverse creases. They also yielded the majority of well-preserved natural casts. ...
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Large well-preserved crocodylomorph tracks from the Lower Cretaceous (? Aptian) Jinju Formation of South Korea, represent the well-known crocodylomorph ichnogenus Batrachopus. The Korean sample includes multiple, narrow-gauge, pes-only trackways with footprint lengths (FL) 18–24 cm, indicating trackmaker body lengths up to ~3.0 m. Surprisingly, the consistent absence of manus tracks in trackways, with well-preserved digital pad and skin traces, argues for bipedal trackmakers, here assigned to Batrachopus grandis ichnosp. nov. No definitive evidence, either from pes-on-manus overprinting or poor track preservation, suggests the trackways where made by quadrupeds that only appear bipedal. This interpretation helps solve previous confusion over interpretation of enigmatic tracks of bipeds from younger (? Albian) Haman Formation sites by showing they are not pterosaurian as previously inferred. Rather, they support the strong consensus that pterosaurs were obligate quadrupeds, not bipeds. Lower Jurassic Batrachopus with foot lengths (FL) in the 2–8 cm range, and Cretaceous Crocodylopodus (FL up to ~9.0 cm) known only from Korea and Spain registered narrow gauge trackways indicating semi-terrestrial/terrestrial quadrupedal gaits. Both ichnogenera, from ichnofamily Batrachopodidae, have been attributed to Protosuchus-like semi-terrestrial crocodylomorphs. The occurrence of bipedal B. grandis ichnosp. nov. is evidence of such adaptations in the Korean Cretaceous.
... In the case of crocodylians and turtles, this scarcity is partly related to the semi-aquatic lifestyle of many Jurassic-Cretaceous groups and the low preservation potential in subaqueous and inundated substrates (Klein and Lucas, 2015). Mammal tracks are generally rare in the Mesozoic, and crocodylomorph tracks have mainly been reported from small terrestrial forms and by traces of swimming individuals, in particular from the Jurassic-Cretaceous of Europe and North America (Olsen and Padian, 1986;Lockley and Meyer, 2004). From the African continent, crocodylomorph tracks are scarcely known (Hadri et al., 2015;Mateus et al., 2017). ...
... A widespread crocodylomorph ichnogenus is Batrachopus (Fig. 3J). Originally described from Lower Jurassic deposits of the Newark Supergroup in eastern North America, it was thereafter identified from units of different age, ranging from the Middle Triassic to the Cretaceous (Olsen and Padian, 1986;Lockley and Meyer, 2004;Lockley et al., , 2010aLockley et al., , 2018Klein and Lucas, 2010;Le Loeuff et al., 2010). Batrachopus has a compact anterior pedal digit group I-IV with straight digits showing distinct rounded pad impressions and sometimes a small, oval, posterior impression, probably being the trace of a strongly reduced digit V, slightly resembling the imprints of Brachychirotherium from the Upper Triassic. ...
... "Leonardi (1975, p. 308) stated that it would be " a useful practice to examine recent trackways and compare them with the body and skeleton of their makers. " Like the footprints of dinosaurs, fossil tracks and traces attributed to crocodylians and their close relatives have received considerable attention, particularly in recent years (Bennett, 1992;Moratalla et al., 1995;Fuentes Vidarte and Meijide Calvo, 2001;Mazin et al., 2003;Lockley and Meyer, 2004;McCrea et al., 2004;Erickson, 2005;Pascual Arribas et al., 2005;Avanzini et al., 2007;Mateus and Mil an, 2010;Contessi and Fanti, 2012;Abbassi et al., 2015;Hadri et al., 2015;Rajkumar et al., 2015;Vila et al., 2015;Segura et al., 2016;Mateus et al., 2017). Unlike non-avian dinosaurs, which are so inconveniently extinct, about thirty crocodylian species remain (at least for now) extant (Grigg and Kirshner, 2015), permitting both laboratory and field observations of footprint creation by these animals (von Huene, 1913;Reineck and Howard, 1978;Padian and Olsen, 1984b;Kubo, 2008Kubo, , 2010bCarpenter, 2009;Farlow and Elsey, 2010;Kumagai and Farlow, 2010;Mil an and Hedegaard, 2010). ...
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We documented trackways of free-living Crocodylus acutus on beaches at the mouths of Tamarindo and Ventanas estuaries, Costa Rica. Our crocodiles had estimated total lengths of 1–3 meters or more. Manus prints have five digits, with digits I–III bearing claw marks. Pes prints have four digits, with claw marks on digits I–III. The pes is plantigrade. Claws generally dig into the substrate. Apart from claw marks, digit I and the heel of the pes are usually the most deeply impressed parts of footprints. Trackways are wide-gauge. Pes prints are usually positioned just behind ipsilateral manus prints of the same set and may overlap them. Manus and pes prints angle slightly outward with respect to the crocodile's direction of movement. Claw-bearing digits of both the manus and pes may create curved, concave-toward-the-midline drag marks as the autopodium is protracted. The tail mark varies in depth and clarity, and in shape from nearly linear to markedly sinuous. Sometimes the tail mark hugs the trackway midline, but sometimes it is closer to, or even cuts across, prints of one side. American crocodile footprints and trackways are similar to those observed in other extant crocodylian species, indicating substantial trackway conservatism across the group.
... Jurassic and Cretaceous ichnotaxa such as Antipus, Batrachopus, Crocodylopdus, and Hatcherichnus have crocodylomorph affinities. The Batrachopus and Antipus trackmakers had functionally tetradactyl feet, according to Lockley and Meyer (2004). Despite the similarity in the number of digits, in Batrachopus pes digits II and IV are sub-equal in size, and slightly smaller than digit III, while in Aracoaraichnium igen. ...
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During the Early Cretaceous, a large area in Brazil and other South American countries was covered by an extensive paleoerg field, the Botucatu Desert. The Araraquara County (São Paulo State, Brazil) contains some of the most diverse ichnological assemblages in the Botucatu Formation. Mammaliform trace fossils from the Botucatu Formation are of great interest, as they may represent the only record of Lower Cretaceous Mammaliamorpha from Brazil. These trace fossils can be grouped into two distinct classes, based on the dimensions of the footprints. The larger morphotype is described and classified as a new ichnotaxon, Aracoaraichnium leonardii igen. nov. isp. nov., with a discussion of preservational features and paleoecological aspects. This morphotype can be differentiated from Brasilichnium Leonardi, 1981 by digit morphology, with digits III and IV being larger and sub-equal in size. The tracks are mesaxonic, unlike Brasilichnium, which is ectaxonic, and the stride is proportionally smaller, suggesting a trackmaker with shorter limbs in relation to body length. Another characteristic that reinforces the new ichnotaxon is the larger size of the footprints of Aracoaraichnium igen. nov. Footprints representing intermediate ontogenetic stages between Brasilichnium and the larger morphotype are not observed, suggesting that the two track types were made by different taxa. Compared with others tetrapod ichnotaxa, the new ichnotaxon is characterized by differences in the number and morphology of the digits, track morphology, the degree of heteropody, and locomotor parameters. Tracks with good preservation quality were likely originally produced in the humid subsurface, below the surficial dry sand layer. Tracks with smaller dimensions, and trackways with the presence or absence of the manus, may be due to variable degrees of impression into subsurface layers, and substrate heterogeneity. The Aracoaraichnium igen. nov. trackmakers probably inhabited areas near the desert, making occasional incursions into desert areas, while Brasilichnium trackmakers may have been mostly nocturnal residents of the desert environment.
... Many different ichnogenera and ichnospecies are attributed to this ichnofamily, leaving the debate on a proper classification still open and under debate (see i.e., Olsen and Padian, 1986;Lockley and Meyer, 2004;Lockley et al., 2004bLockley et al., , 2010. The mostly representative morphotype, Batrachopus Hitchcock, 1845, has been tentatively attributed to 'true crocodilian or to a crocodylomorph with a pedal digit V reduced to the state seen in crocodilians' by Olsen and Padian (1986) and is described from a Lower Jurassic ichnofauna of the United States (Hitchcock, 1845;Olsen and Padian, 1986;Milner and Lockley, 2006). ...
... Ichnogenus Antipus Hitchcock, 1858, is considered a distinct morphotype from Batrachopus. They differ to one another for the digit morphology, stout digits, and with minimal digital divarication in Batrachopus, much more slender, with sharpen digits, and widely divaricated in Antipus (Lockley and Meyer, 2004) and for the inner trackway width, close to 0 in Batrachopus (the right and left pedes touch the trackway midline), and about 4 cm in Antipus (Lockley and Meyer, 2004). However, Lockley and Meyer (2004) pointed out that this difference in the trackway width could be due to the walking speed of an identical trackmaker, with Antipus representing a slower version of Batrachopus. ...
... Ichnogenus Antipus Hitchcock, 1858, is considered a distinct morphotype from Batrachopus. They differ to one another for the digit morphology, stout digits, and with minimal digital divarication in Batrachopus, much more slender, with sharpen digits, and widely divaricated in Antipus (Lockley and Meyer, 2004) and for the inner trackway width, close to 0 in Batrachopus (the right and left pedes touch the trackway midline), and about 4 cm in Antipus (Lockley and Meyer, 2004). However, Lockley and Meyer (2004) pointed out that this difference in the trackway width could be due to the walking speed of an identical trackmaker, with Antipus representing a slower version of Batrachopus. ...
... The pes track from the Serraduy Norte locality shows a different morphology and size, most probably indicating a different trackmaker and/or behaviour. The track preserves clear digital and heel impressions (plantigrade progression) and strongly resembles Crocodylopodus, particularly in the triangular and symmetrical shape of the heel impression and the orientation of digits (Fuentes Vidarte & Meijide Calvo 2001;Lockley & Meyer 2004), except for digit IV. The studied specimen is also very close to tracks attributed to Crocodylopodus meijidei by Avanzini et al. (2007, p. 148, Fig. 4D). ...
Article
Recently discovered evidence of tracks in the continental beds of the Late Cretaceous Tremp Formation in the southern Pyrenees (NE Iberian Peninsula) has been identified as scratch marks made by buoyant crocodiles. The tracks are preserved in two distinct environments and substrates (marly limestones originating in a littoral mud flat and fine-grained sandstones deposited in fluvial settings). Most of the crocodylian traces are ascribed to ichnogenus Characichnos, whereas a single plantigrade pes track is assigned to ichnogenus cf. Crocodylopodus. The crocodylian swim traces (Characichnos ichnofacies) found in the early and late Maastrichtian co-occur with Brontopodus ichnofacies attributable to terrestrial tetrapods (titanosaur sauropods, cf. Brontopodus ichnogenus; and hadrosaurid ornithopods, Hadrosauropodus ichnogenus). Analysis of the tracks allows the interpretation of palaeoenvironmental settings and track production. Thus, in lagoonal environments, swim tracks of crocodylians were produced during the rise of the water level in successive tide cycles; in fluvial settings, the swim traces of crocodylians were produced within the channel at the low-water stage. To date, there are no reports of Late Cretaceous crocodylian tracks in Europe, and the studied evidence represents the first and youngest track record of the group in the latest part of the Cretaceous (C29r) in this continent and probably in the world.
... Five digits in the manus impressions are also features of lacertilian tracks (e.g., Rhynchosauroides), although such tracks are characterized by curved and slender digits, and increase in length from digit I to IV Avanzini et al., 2010). Footprints of other small reptiles, such as crocodilians are excluded because characterized by pentadactyl manus, tetradactyl pes impressions, slender digits, and greater (about 180 ) interdigital angulation between digits I and V (Lockley and Meyer, 2004;Avanzini et al., 2007). The Tunisian tracks are small in size, have straight central digits, shorter lateral ones, and pad impressions are consistent with mammalian phalangeal formula of 2:3:3:3:3; therefore they can be tentatively referred to a mammal trackmakers. ...